Evolution and Human Behavior 38 (2017) 155–163

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Evolution and Human Behavior

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Original Articles

Women's evaluations of other women's natural body odor depend on

target's fertility status
Kelly A. Gildersleeve a,⁎, Melissa R. Fales b, Martie G. Haselton b
a
Chapman University, USA
b
University of California, Los Angeles, USA

a r t i c l e i n f o a b s t r a c t

Article history: A large research literature indicates that men perceive women as more attractive when they are at high fertility
Initial receipt 15 May 2015 than at low fertility within the ovulatory cycle. However, it remains unclear whether women also perceive
Final revision received 16 August 2016 women as more attractive at high fertility. This study examined women's ratings of samples of natural body
odor collected from naturally-cycling women at high and low fertility within the cycle and from hormonal
Keywords:
contraceptive-using women at mid-cycle. Like men, women rated naturally-cycling women's high-fertility
Scent
Attractiveness
scent samples as more attractive than their low-fertility samples. Women rated hormonal contraceptive (HC)
Ovulatory cycle users' scent samples as more attractive than naturally-cycling women's high- and low-fertility samples, though
Hormonal contraception the difference between HC and high-fertility samples was statistically significant only when raters were treated
as the unit of analysis. These findings reveal a potentially important role for scent communication in women's
perceptions of other women and are consistent with the notion that the ovulatory cycle could influence women's
interactions with one another. The findings also highlight the need for rigorous investigations of the possible
impacts of hormonal contraception on women's attractiveness and social relationships with other women.
© 2016 Elsevier Inc. All rights reserved.

1. Introduction scents collected from women at low fertility experienced a decrease in

A large research literature indicates that men evaluate women's
faces, voices, and natural body odors as subtly more attractive on the
handful of high-fertility days leading up to ovulation, as compared
with the low-fertility days comprising the remainder of the cycle
(reviewed in Haselton & Gildersleeve, 2011, 2016; meta-analyzed in
Gildersleeve & Haselton, 2015). Findings concerning men's evaluations
of the attractiveness of women's high- versus low-fertility body scents
appear to be robust, with many studies reporting evidence that men
perceive women's high-fertility scents as more pleasant and sexier
(e.g., Cerda-Molina, Hernández-López, de la O, Chavira-Ramírez, &
Mondragón-Ceballos, 2013; Doty, Ford, Preti, & Huggins, 1975;
Gildersleeve, Haselton, Larson, & Pillsworth, 2012; Havlíček, Dvorakova,
Bartos, & Flegr, 2006; Singh & Bronstad, 2001; Thornhill et al., 2003; but
see Thornhill & Gangestad, 1999). Moreover, preliminary evidence
suggests that men respond to women's high-fertility scents with greater
levels of testosterone, as well as motivations and behaviors thought to
facilitate courtship behavior (reviewed in Makhanova & Miller, 2013).
For example, in a recent study, men who inhaled underarm or vulvar
scents collected from women at high fertility subsequently experienced
increases in testosterone and sexual interest, whereas men who inhaled
⁎ Corresponding author. Department of Psychology, Chapman University, 544 N.
Cypress St., Orange, CA 92867, USA.
E-mail address: kellygildersleeve@gmail.com (K.A. Gildersleeve).
testosterone and no change in sexual interest (Cerda-Molina et al.,
2013; also see Miller & Maner, 2010a, 2010b; but see Roney & Simmons,
2012, for a null effect of high-fertility scents versus water on men's
testosterone).
Parallel patterns have been extensively documented in nonhuman
mammals. Across diverse species, it is typical for females to experience
physical changes, including changes in scent, during the brief high-
fertility period approaching ovulation. Males typically respond to
these fertility cues with enhanced sexual interest and, in some cases, in-
creases in hormones that facilitate mating and intrasexual competition
(e.g., Amstislavskaya & Popova, 2004; Bronson & Desjardins, 1982;
Elvira, Herndon, & Wilson, 1982; Gordon, Bernstein, & Rose, 1978;
Kavaliers, Choleris, & Colwell, 2001; Perret & Schilling, 1995; Rose, Gor-
don, & Bernstein, 1972; Ziegler, Schultz-Darken, Scott, Snowdon, &
Ferris, 2005). Thus, in both humans and nonhuman mammals, cues of
fertility within the ovulatory cycle appear to play a role in regulating
male–female and, possibly, male–male social interactions.
But what about female–female social interactions? A recent study of
chacma baboons suggests that sensitivity to cues of female fertility
within the ovulatory cycle is not unique to males. In the study, female
baboons were more aggressive toward females in the high-fertility pe-
riod of the ovulatory cycle than toward females in other reproductive
states (i.e. in the low-fertility period of the cycle, pregnant, or lactating;
Huchard & Cowlishaw, 2011). Female aggressors therefore appeared to
detect cues of fertility in potential targets and increase aggression

http://dx.doi.org/10.1016/j.evolhumbehav.2016.08.003
1090-5138/© 2016 Elsevier Inc. All rights reserved.
156 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
toward the females who were likely to be most attractive to prospective
mates and therefore posed the greatest competitive threat. Given strong
evidence that men are sensitive to subtle cues of women's fertility with-
in the cycle, this finding in baboons raises the question of whether
women are also sensitive to cues of fertility in other women.
To date, only four studies have produced data relevant to addressing
this question. Although these studies provide valuable preliminary evi-
dence, their findings have been mixed, and their methods have been
limited. Three of these studies examined possible ovulatory cycle effects
on women's evaluations of other women's natural body scent. In one
study, women (n = 12) rated T-shirts worn by female stimulus donors
estimated to be closer to ovulation within the cycle as smelling more
attractive than T-shirts worn by stimulus donors estimated to be farther
from ovulation (i.e. a “between-donors” design; donor n = 41;
Kuukasjärvi et al., 2004). Likewise, in a recent study using a more pow-
erful “within-donors” design, women (n = 58) rated T-shirts worn
by women at high fertility as smelling more pleasant than T-shirts
worn by the same women at low fertility (donor n = 13, though two do-
nors provided only low-fertility T-shirts; Woodward, Thompson, &
Gangestad, 2015). However, in another study using a within-donors de-
sign, neither heterosexual women (n = 12) nor non-heterosexual
women (n = 8) rated T-shirts worn by women at high fertility as smell-
ing more attractive than T-shirts worn at low fertility (donor n = 17;
Trouton, Guitar, Carmen, Geher, & Grandis, 2012). Lastly, one study
used a within-donors design to examine possible ovulatory cycle effects
on women's evaluations of other women's facial attractiveness. Women
(n = 131) judged photographs of women taken at high fertility as more
attractive than photographs of the same women at low fertility (donor
n = 48; Roberts et al., 2004).
A limitation of all of these studies is that estimations of stimulus do-
nors' positions in the cycle (and of their current fertility) were based on
methods that involve counting forward from a self-reported date of last
menstrual onset to the day of participation. In comparison with
methods that involve counting backward from a prospectively verified
date of next menstrual onset or repeat assessments of luteinizing hor-
mone within the expected fertile window, such “forward counting”
methods are low in validity (they are relatively poor at estimating
women's true fertility within the cycle) and offer low statistical power
to detect true cycle effects (see Gangestad et al., 2015; Gonzales &
Ferrer, 2015). Forward counting methods perform poorly largely be-
cause they are vulnerable to error in women's recollections of their
date of last menstrual onset. Recall error can be substantial; for example,
one study compared women's retrospective reports of their date of last
menstrual onset to the date they had prospectively reported (see
Wegienka & Baird, 2005) and found that, although 56% of the women
retrospectively reported the correct date, 19% were off by three or
more days (and note that the fertile window itself lasts only approxi-
mately 6 days; Wilcox, Weinberg, & Baird, 1995).
In addition, all but one of these past studies (Kuukasjärvi et al.,
2004) presented analyses treating raters as the statistical “unit of
analysis” and did not present analyses treating stimulus donors as
the unit of analysis. Statistically significant fertility effects based on
analyses treating raters as the unit of analysis justify the inference
that other samples of raters from the population also will probably
evaluate those particular high-fertility stimuli as more attractive
than those particular low-fertility stimuli. However, such analyses
cannot rule out the possibility that differences between high- and
low-fertility stimuli that are idiosyncratic to a particular stimulus
set have produced the illusion of a fertility effect.
For example, if a scent study similar to those described above
used a stimulus set that included one low-fertility scent sample
that smelled particularly unpleasant, analyses that collapsed across
donors in order to examine variation among raters in their evalua-
tions of high- versus low-fertility scent samples – in other words,
that treated raters as the unit of analysis – might well detect an ap-
parent fertility effect. However, this effect could be driven entirely
by the single unpleasant smelling low-fertility sample, which
might have smelled unpleasant for any number of reasons not neces-
sarily related to fertility (e.g., the stimulus donor ate garlic that day
but did not report it to the researcher).
In contrast, analyses that collapsed across raters in order to examine
variation among donors in the evaluations they received for their high-
versus low-fertility samples – in other words, that treated donors as the
unit of analysis – would be less likely to detect this potentially spurious
fertility effect against the background of between-donor variation in
high- versus low-fertility scent attractiveness. We emphasize that only
donors-as-unit analyses provide a test of the key hypothesis of interest —
namely, that women generally are more attractive at high than at low
fertility within the ovulatory cycle. Once a fertility effect has been
compellingly demonstrated by conducting donors-as-unit analyses,
raters-as-unit analyses are useful for establishing the generalizability
of the finding to other possible samples of raters.
Finally, whereas previous studies examining men's evaluations of
women's high- versus low-fertility attractiveness have typically pro-
posed that men's preference for high-fertility stimuli reflects psycholog-
ical adaptations for detecting cues of women's current fertility within
the cycle (e.g., see Singh & Bronstad, 2001), studies examining women's
evaluations of other women's high- and low-fertility attractiveness
have tended to frame such analyses as exploratory or did not explicate
a clear rationale for why we should expect women to be sensitive to
cues of fertility in other women. In fact, there are several reasons to
expect that women, like men, will perceive women as more attractive
at high fertility within the ovulatory cycle. First, given evidence that
females of related primate species likely possess psychological
mechanisms that enable them to detect cues of fertility in other females
(e.g., see Huchard & Cowlishaw, 2011), it is plausible that such
mechanisms could appear in human females as a vestigial trait passed
down from a shared ancestor in which such mechanisms were functional.
Alternatively, given that human males appear to possess psychological
mechanisms that enable them to detect cues of fertility in women, it is
plausible that such mechanisms could appear in women as a mere
byproduct of their shared physiology with men (e.g., arising due to devel-
opmental constraints).
However, we think it is more likely that selection pressures encoun-
tered by ancestral human females actively favored the psychological
mechanisms that now enable women to perceive cues of high fertility
in other women as attractive. These mechanisms may not necessarily
have been selected de novo in humans; rather, they may have been
passed down from ancestral species and maintained in human females
because of their reproductive benefits. For example, three non-mutually
exclusive possibilities are that (a) selection favored a sensitivity among
women to cues of overall reproductive quality in other women (i.e.
between-women variation in reproductive potential), (b) selection
favored sensitivity to cues of current cycle fertility in other women (i.e.
within-women, between-cycle variation in fertility), and (c) selection
favored sensitivity to cues of fertility within the cycle in other women
(i.e. within-women, within-cycle variation in fertility). Detecting cues
of any of these three types of variation might have reproductively
benefitted women in a variety of ways, including enabling them to di-
rect increased aggression at female rivals who posed a heightened com-
petitive threat in general (relative to other women), in their current
cycle (relative to other cycles), or at this point within their current
cycle (relative to other points within their cycle).
Notably, some factors – such as estradiol levels – are thought to
correlate with overall reproductive quality, current cycle fertility, and
fertility within the cycle (see Law Smith et al., 2006; Puts et al., 2014).
Therefore, if female sensitivity to cues of estradiol initially evolved to
enable women to detect between-women variation in overall reproduc-
tive quality, it might also incidentally confer an ability to detect within-
women variation in current cycle fertility or fertility within the cycle (or
vice versa. For an extended discussion of similar evolutionary explana-
tions for men's sensitivity to cues of fertility in women, see Havlíček,
 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
Cobey, Barrett, Klapilova, & Roberts, 2015; but see critiques by Roney,
Lukaszewski, Simmons, Eisenbruch, & Grillot, 2015; Gangestad &
Grebe, 2015; Haselton, 2015).
Though it is ultimately an empirical question, all of the above expla-
nations could potentially produce the same pattern of findings. There-
fore, the present study is not intended as a test between these
different accounts. Instead, the primary aim of the present study was
to use rigorous methods to determine more definitively whether
women perceive naturally-cycling women's natural body odor as
more attractive at high fertility than at low fertility. A positive finding
would indicate that there is indeed a phenomenon to be explained,
with potentially important implications for understanding how – and
why – the ovulatory cycle influences social interactions among women.
A subsidiary aim of the present study was to conduct exploratory
analyses to compare women's evaluations of the attractiveness of
naturally-cycling women's body odor with their evaluations of the
body odor of women using combined oral hormonal contraceptives
(HC). Shifts across the ovulatory cycle in women's attractiveness are
thought to be mediated by reproductive hormones, including estradiol
and progesterone (see Puts et al., 2013). Combined oral hormonal con-
traceptives typically contain a synthetic estrogen, ethinyl estradiol
(though some newer forms contain a natural estrogen), and one of sev-
eral synthetic progestins. The introduction of these exogenous hor-
mones into women's bodies reduces their secretion of endogenous
estradiol, progesterone, and other hormones and also suppresses
naturally-occurring cyclic variation therein (e.g., see Fleischman,
Navarrete, & Fessler, 2010). It follows that hormonal contraceptives
could influence women's scent attractiveness.
In order to address this question, two studies examining shifts across
the cycle in women's attractiveness have included a subsample of HC-
users as a comparison group. One study (also discussed above) found
that, unlike naturally-cycling women, HC-users did not experience a
shift across the cycle in scent attractiveness (Kuukasjärvi et al., 2004).
However, this did not result in an on-average difference in scent attrac-
tiveness between HC-users (n = 42) and naturally-cycling women
(n = 39). The second study – which tracked female lap dancers' tip
earnings over a period of 60 days – likewise found that, unlike
naturally-cycling women, HC-users did not exhibit a shift across the
cycle in tips received from male strip club patrons (within-women de-
sign; Miller, Tybur, & Jordan, 2007). In this case, HC-users earned less
money in tips on average than did naturally-cycling women. However,
sample sizes were small (n = 7 and 11, respectively, though many ob-
servations were nested within each woman).
Lastly, one recent within-women study found that women's male
romantic partners rated them as less attractive when they were using
hormonal contraceptives than when they were naturally-cycling and
at high fertility or naturally cycling and at low fertility within the
cycle. However, because all participants completed these three sessions
in the same order, an order effect cannot be ruled out (n = 14; Cobey,
Buunk, Pollet, Klipping, & Roberts, 2013). In sum, findings in the extant
literature are consistent with the notion that HC-using women – unlike
naturally-cycling women – do not experience changes across the cycle
in their attractiveness, but it remains unclear whether women are per-
ceived as less (or more) attractive when using hormonal contraception
(reviewed in Alvergne & Lummaa, 2010).
In the present study, we collected samples of natural body odor from
naturally-cycling women at high fertility and at low fertility within the
cycle, based on luteinizing hormone test results and prospectively veri-
fied dates of menstrual onset. We also collected body odor samples from
HC users at mid-cycle. A large sample of women evaluated all of the
samples for scent attractiveness and intensity. We then conducted
two sets of analyses, treating scent donors and raters as the statistical
unit of analysis, respectively. We conducted both types of analyses in
order to ensure that significant findings will generalize to other possible
stimulus donors (and sets of high- and low-fertility scent stimuli), as
well as to other possible raters.
157
2. Methods
2.1. Scent donors and sample collection procedures
A total of 33 women (mean age = 19.97, SD = 1.87) – 21 naturally-
cycling women and 12 hormonal contraceptive users – participated as
scent donors. Scent donors reported the following ethnicities: African
American (n = 3), Asian (n = 8), Caucasian (n = 11), Hispanic (n =
5), multiple ethnicities (n = 2), and not reported (n = 4). Scent donors
collected samples of their natural body odor by wearing cotton gauze
pads under their arms for 24 hours. During this period, they followed
strict behavioral guidelines designed to minimize the contamination
of their samples by non-natural scents. For example, deodorant and an-
tiperspirants were prohibited. Scent samples were frozen until the rat-
ing session to prevent odors from dissipating (Lenochova, Roberts, &
Havlíček, 2009; Roberts, Gosling, Carter, & Petrie, 2008). To control for
possible effects of body side, only right-arm scent samples were used
in the present study. Procedures used to collect and store scent samples
were identical to those used in our previous study. Please see
Gildersleeve et al. (2012) for a more detailed description.
The 21 scent donors who were naturally cycling had not used hor-
monal contraception or hormonal medications in the past three months
and reported regular menstrual cycles with an average length between
25 and 34 days. Fourteen of these women had participated as scent do-
nors in our previous study (Gildersleeve et al., 2012), and their high-
and low-fertility scent samples were re-used in the present study. We
chose to re-use the scent samples of this particular subsample of
women because they had reported excellent compliance with the
scent collection guidelines (see “Ideal Donors” from Gildersleeve et al.,
2012; the one remaining Ideal Donor's samples were lost and therefore
could not be re-used in the present study). Recruitment methods for
these 14 women are described in detail in our previous study
(Gildersleeve et al., 2012).
The remaining seven women provided new scent samples
specifically for this study. Initially, we recruited 30 women to provide
new scent samples for this study and for a related study being conduct-
ed concurrently (odor sample collection was combined for the two
studies). However, 18 of those women did not provide usable scent
samples because they did not show compelling evidence of an LH
surge (see below; n = 12), used the LH tests incorrectly (n = 1), expe-
rienced an irregular cycle that precluded us from accurately scheduling
their high- and low-fertility sessions (n = 1), withdrew voluntarily
(e.g., due to conflicts with exams; n = 3), or had missing scent collec-
tion compliance information (n = 1). Of the 12 pairs of usable samples
that remained, five women's samples were allocated to the related
study. The other seven women's samples were allocated to this study
(though one of the women lost her high-fertility sample and therefore
supplied only a low-fertility sample).
The primary purpose of using an additional seven women's new
scent samples in the present study (rather than using only the 14
pairs of samples from our previous study) was to ensure sufficient
power for statistical analyses treating scent donors as the unit of analy-
sis. We previously found significant fertility effects on women's scent at-
tractiveness in a sample of 15 pairs of scent samples (see Gildersleeve
et al., 2012). Therefore, we reasoned that 20 pairs of samples would
be more than sufficient for the present investigation. Thus, in total,
this study used odor samples provided by 21 naturally-cycling women.
We scheduled women's high- and low-fertility scent collection ses-
sions using the backward counting method based on prospectively re-
ported dates of menstrual onset. In other words, donors prospectively
reported the date of their next menstrual onset after completing both
sessions (see Gildersleeve et al., 2012). We counted back from this
date to determine their current position within the cycle. To verify
that the sessions fell on true high- and low-fertility days of the cycle,
scent donors also completed urine tests for luteinizing hormone for
five consecutive days surrounding their high-fertility session. These
158 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
methods have been shown to yield higher-validity estimates of
women's fertility within the cycle than the two most common methods
in this research area — namely, backward counting based on predicted
(rather than prospectively reported) dates of menstrual onset and for-
ward counting based on retrospectively recalled dates of menstrual
onset (see Gangestad et al., 2016). Furthermore, we counterbalanced
the order of women's high- and low-fertility sessions (52% of the
women completed their high-fertility session first).
On average, scent donors provided their high- and low-fertility scent
samples 16.24 days (SD = 2.59) and 5.43 days (SD = 3.61), respectively,
before the onset of their next menstrual period. These days fall within
expected high- and low-fertility phases of the cycle, respectively. In ad-
dition, on average, women showed evidence of a luteinizing hormone
surge on the same day as their high-fertility session (mean = 0.29
days before their high-fertility session, SD = 1.52, range = 2 days before
to 2 days after their high-fertility session). A luteinizing hormone surge
indicates that ovulation will occur within the next 24–48 hours (Testart
& Frydman, 1982), with rare exceptions (e.g., see Killick & Elstein, 1987),
and past research indicates that women's fertility is highest on the day
before and the day of ovulation (Wilcox et al., 1995). Thus, it is highly
likely that women's high- and low-fertility scent collection sessions
fell within the correct cycle phases.
The remaining 12 scent donors reported using a 28-day combined
oral hormonal contraceptive and had used that contraceptive for at
least three months prior to participating in the study (Lo-Ogestrel,
Apri, Zovia, Ortho Tri-Cyclen, Lutera, Enpresse,Tri Sprintec, Lo Loestrin
FE, and Gildess). Initially, we recruited 13 hormonal contraceptive
users to participate as scent donors (our target sample size was 20,
but only 13 eligible women were identified by the participant recruit-
ment cutoff date). However, one of these women voluntarily withdrew
due to scheduling conflicts. We scheduled the 12 hormonal contracep-
tive users to provide scent samples just once, about 14 days after the
start of their last round of placebo pills (approximately halfway be-
tween menstrual periods). Hormonal contraceptive users provided
their scent samples an average of 13.25 days (SD = 1.71) after the
first day of their most recent round of placebo pills, according to their
retrospective (n = 5) or prospective (n = 7) report of that date.
2.2. Scent raters and rating procedures
A total of 96 women participated as scent raters. Raters were recruit-
ed in-person from a community Lesbian and Gay Pride festival while
conducting a related study and via fliers posted at a university campus.
A brief prescreening interview was used to determine their eligibility.
Women were deemed eligible if they reported that they met all of the
following inclusion criteria: between the ages of 18 and 40 years,
smoked cigarettes no more than two times per week (smoking is asso-
ciated with olfactory deficits; Katotomichelakis et al., 2007), non-
impaired sense of smell, not pregnant, had not given birth within the
last year, not breastfeeding, not post-menopausal or currently
experiencing symptoms of menopause, had not used emergency hor-
monal contraception (“Plan B”) in the last three months, and had not
used other hormonal medications in the last three months (except for
hormonal contraception, which was acceptable).
Of the 96 women who participated in the scent rating sessions, four
women's data were excluded from analysis because their questionnaire
responses indicated that they actually did smoke cigarettes three or
more times per week (n = 3) or might have an impaired sense of
smell (due to a deviated septum surgery; n = 1). One additional
woman opted out of the scent rating task (but participated in other por-
tions of the study). Thus, the analyses presented below are based on 91
raters (mean age = 22.2 years, SD = 4.8). Raters reported the following
ethnicities: African American (n = 1), Asian (n = 30), Caucasian (n =
21), Hispanic (n = 15), multiple ethnicities (n = 21), and other (n = 3).
Rating sessions were held on two consecutive days as part of a larger
study. Raters could arrive at any time between 10 a.m. and 5 p.m. on the
first day or between 10 a.m. and 2 p.m. on the second day. In addition to
the main scent rating task (described in detail below), raters completed
a task designed to examine changes in hormones and motivations in re-
sponse to human scent stimuli, questionnaires collecting demographic
(e.g., age, ethnicity, socioeconomic status) and biographical information
(e.g., relationship and sexual history, sexual orientation, recent stress
experienced in their romantic relationship [if applicable], hormonal
contraceptive use), and a saliva sample for genotyping. These other
tasks were unrelated to the present investigation, and the results will
be reported elsewhere.
On the evening before the first rating session, we transferred the
scent samples into 2 oz. plastic bottles and then returned them to the
freezer. On the morning of both rating sessions, we removed the bottles
from the freezer, removed their caps, and allowed the samples to thaw
for two hours before the raters began to arrive. Bottles were left
uncapped and at room temperature throughout both rating sessions.
At the end of each rating session, we returned the scent stimuli to
the freezer.
For both sessions, we set up 21 scent rating stations around a large
classroom. At each station, we placed two or three bottles, labeled A,
B, and C. These bottles typically contained a high-fertility scent sample
from a naturally-cycling woman, a low-fertility scent sample from the
same woman, and if there was a third bottle, a scent sample from a hor-
monal contraceptive user. In three cases only (see below), one of the
bottles instead contained an unworn gauze pad for the purpose of ex-
amining how raters perceived gauze pads scented with women's body
odor as compared with unworn pads. We used random assignment to
determine which donors' scent samples would be presented at which
scent rating station and to determine which samples would be assigned
to bottles A, B, and C at a given station. The scent station setup was iden-
tical across both rating sessions.
All raters evaluated all of the scent samples, and the order in which
raters traveled from one scent rating station to the next was pre-
randomized. In a previous study in which men evaluated pairs of
high- and low-fertility scent samples from 18–32 women, we found
that men's ability to accurately discriminate between the same
woman's high- and low-fertility samples on an A-B-X task did not de-
cline the more trials they completed; if anything, accuracy slightly in-
creased (r = 0.09; Gildersleeve et al., 2012). Therefore, our past
research does not suggest that rater fatigue substantially impacted
women's evaluations of high- versus low-fertility scent samples in the
present study. In addition, it was common for multiple raters to be in
the classroom completing scent ratings at the same time. To create pri-
vacy and minimize distraction, scent rating stations were spaced apart
by at least a few feet and surrounded by cardboard carrels.
Raters completed two kinds of judgments at each scent rating station:
forced choices between samples and individual ratings of each sample.
For the forced choices, participants were instructed to hold Bottle A just
under their nose, being careful not to touch it to their face, and to take a
hearty sniff; repeat this procedure for Bottle B; and then mark on their rat-
ing form which of the two bottles – A or B – smelled more attractive to
them. They repeated this task to compare all possible pairs of samples at
that station (e.g., A vs. B, A vs. C, and B vs. C). For the individual ratings of
each sample, participants were instructed to again smell Bottle A and rate
it on three dimensions: pleasantness, sexiness, and intensity (1 = Very Un-
pleasant/Very Unsexy/Very Unintense; 9 = Very Pleasant/Very Sexy/Very
Intense). They repeated this procedure for all bottles at that station. They
then moved to the next station on their rating form, and so on.
Raters completed scent ratings at their own pace, usually finishing
within 30 to 60 min. Importantly, raters were not informed of the pur-
poses of the study in advance of their participation. They were told
only that they would be smelling and rating gauze pads that had been
worn by women and that the study had been designed to examine the
role of scent in women's attractions. In response to a debriefing ques-
tionnaire at the end of the study, none of the raters accurately guessed
the precise hypotheses under investigation.
 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163 159

3. Results
For the 14 pairs of samples that were reused from our previous study
(Gildersleeve et al., 2012), ratings received from the male raters in our
previous study correlated highly with ratings received from the female
raters in the present study (pleasantness at high fertility, r = .85; pleas-
antness at low fertility, r = .81; sexiness at high fertility, r = .82; sexi-
ness at low fertility, r = .75; intensity at high fertility, r = .85;
intensity at low fertility, r = .90). This suggests that leaving the bottles
uncapped during the rating sessions, freezer storage, and freeze–thaw
cycles did not substantially affect the quality of the scent samples
(also see Lenochova et al., 2009).
To examine differences between women's evaluations of naturally-
cycling women's high- and low-fertility scent samples and hormonal
contraceptive users' samples, we conducted two sets of analyses: First,
we treated scent donors as the unit of analysis, and second, we treated
scent raters as the unit of analysis.
3.1. Stimulus donors as the unit of analysis
Ratings of scent pleasantness and sexiness were highly positively
correlated with each other at high fertility (r = 0.99) and at low fertility
(r = 0.99). However, in keeping with prior studies using these
measures (e.g., Singh & Bronstad, 2001), we report these separately
rather than as composites. Scent pleasantness and sexiness were also
negatively correlated with ratings of scent intensity at high fertility
(rpleasant, intense = −.60; rsexy, intense = −.52) and at low fertility
(rpleasant, intense = −.87; rsexy, intense = −.83).
Twenty forced-choice trials involved comparisons between
naturally-cycling donors' high-fertility samples and those same
women's low-fertility samples. As shown in Fig. 1, on average, donors'
high-fertility samples were chosen as more attractive than their low-
fertility samples by 60% of the raters (SD = 22%). A one-sample t-test in-
dicated that this rate was significantly greater than chance (an expected
rate of 50%), t(19) = 2.14, p = .046, d = .48. An independent t-test
showed that this rate did not differ depending on the order in which
donors had provided their high- and low-fertility scent samples,
t(18) = −.34, p = .74. In addition, as shown in Fig. 2, donors' high-
fertility samples were rated as smelling significantly more pleasant
(t(19) = 2.40, p = .03, d = 0.55) and sexier (t(19) = 2.30, p = .03,
d = 0.53) but marginally less intense (t(19) = −2.02, p = .06, d = −.47)
than their low-fertility samples.
Twelve forced-choice trials involved comparisons between
naturally-cycling donors' high-fertility samples and hormonal contra-
ception users' samples. On average, naturally-cycling women's high-
fertility samples were chosen as more attractive than hormonal
contraception users' samples by only 42% of the raters (SD = 23%). In
other words, hormonal contraception users' scent samples were slightly
preferred over naturally-cycling women's high-fertility samples, but
this rate did not exceed chance, t(11) = −1.17, p = .27, d = −.34.
This rate did not differ depending on the order in which naturally-
cycling donors had provided their high- and low-fertility scent samples,
t(10) = −.98, p = .35. Hormonal contraception users' samples and
naturally-cycling women's high-fertility samples were not rated differ-
ently in terms of pleasantness (t(30) = 0.98, p = .34, d = 0.38), sexi-
ness (t(30) = 1.04, p = .31, d = 0.39), or intensity (t(30) = 0.75,
p = .46, d = −0.29; here, positive d values indicate higher ratings of
hormonal contraception users' samples).
An additional 12 forced-choice trials involved comparing naturally-
cycling women's low-fertility samples to hormonal contraception
users' samples. On average, naturally-cycling women's low-fertility
samples were chosen as more attractive than hormonal contraception
users' samples by only 29% of the raters (SD = 25%). In other words,
hormonal contraception users' scents were preferred over naturally-
cycling women's low-fertility scents at a rate significantly greater than
chance, t(11) = 2.85, p = .02, d = .82. This rate did not differ depending
on the order in which naturally-cycling donors had provided their high-
and low-fertility scent samples, t(10) = −.72, p = .49. Furthermore,
hormonal contraception users' samples were rated as smelling signifi-
cantly more pleasant (t(31) = 2.41, p = .02, d = 1.04) and sexier
(t(31) = 2.40, p = .02, d = 1.01) but less intense (borderline signifi-
cant; t(31) = −2.05, p = .05, d = −0.84) than naturally-cycling
women's low-fertility samples.
3.2. Raters as the unit of analysis
On the 20 forced-choice trials involving comparisons between
naturally-cycling women's high-fertility samples and those same
women's low-fertility samples, raters (N = 91) judged high-fertility
samples to smell more attractive than low-fertility samples for 61% of
the donors, on average (SD = 10%). This rate was significantly greater
than chance (50%), t(90) = 10.23, p b .001, d = 1.07. Furthermore,
raters judged women's high-fertility samples to smell significantly
more pleasant (t(90) = 12.78, d = 0.67) and sexier (t(90) = 8.85,
d = 0.38) but less intense (t(90) = −13.35, d = −0.61) than their
low-fertility samples, all p values b.001.
On the 12 forced-choice trials involving comparisons between
naturally-cycling women's high-fertility samples and hormonal contra-
ception users' samples, raters (N = 91) judged high-fertility samples to
smell more attractive than samples from hormonal contraception users
on only 42% of the trials, on average (SD = 11%). In other words, raters
preferred hormonal contraception users' scents over naturally-cycling

Forced Choice Comparing Attractiveness of Scent Samples from Naturally-cycling Women

at High and Low Fertility and from Hormonal Contraception Users
100
90 *
80
70
60 *
50 Chance (50%)
40
30 NC High Fertility > NC Low Fertility
20
NC High Fertility > HC Users
10
NC Low Fertility > HC Users
0
Percentage of Trials Chosen as More Attractive
* p < .05

Fig. 1. Percentages of forced-choice trials on which naturally-cycling (NC) women's high-fertility samples were chosen as more attractive than their low-fertility samples, naturally-cycling
women's high-fertility samples were chosen as more attractive than hormonal contraception (HC) users' samples, and naturally-cycling women's low-fertility samples were chosen as
more attractive than hormonal contraception users' samples, treating scent donors as the unit of analysis. Error bars depict standard deviations. Asterisks depict significance levels for
comparisons against chance (50%).
160 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
Ratings of Naturally-cycling and Hormonal Contraception-using Women’s Scent Samples
9
8
7 * * NC Low Fertility
6
1
Pleasantness
†p .10, * p < .05
Fig. 2. Mean ratings of scent samples provided by naturally-cycling (NC) women at high and low fertility and by hormonal contraception (HC) users, treating scent donors as the unit of
analysis. Error bars depict standard deviations.
women's high-fertility scents, and this rate was significantly greater
than chance, t(90) = −6.42, p b .001, d = −.67. Furthermore, raters
judged the samples of women using hormonal contraception to smell
significantly more pleasant (t(90) = 7.14, d = 0.31) and sexier
(t(90) = 5.63, d = 0.20) but less intense (t(90) = −6.59, d = −0.23)
than naturally-cycling women's high-fertility samples, all p values
b.001.
On the 12 forced-choice trials involving comparisons between
naturally-cycling women's low-fertility samples and hormonal contra-
ception users' samples, raters (N = 91) judged low-fertility samples
to smell more attractive than samples from hormonal contraception
users on only 29% of the trials (SD = 13%). Thus, raters also preferred
hormonal contraception users' scents over naturally-cycling women's
low-fertility scents at a rate significantly greater than chance,
t(90) = −14.77, p b .001, d = 1.55. Furthermore, raters judged the sam-
ples of women using hormonal contraception to smell significantly
more pleasant (t(90) = 14.68, d = 0.96) and sexier (t(90) = 10.47,
d = 0.57) but less intense (t(90) = −14.07, d = −0.81) than
naturally-cycling women's low-fertility samples, all p values b.001.
Raters completed three forced-choice trials involving unworn gauze
pads that had been randomly paired with a high-or low-fertility sample.
As a result of having one more low-fertility than high-fertility sample,
raters completed two trials comparing low-fertility samples to unworn
gauze pads and a single trial comparing a high-fertility sample to an un-
worn gauze pad. On average, raters (N = 89; two women failed to com-
plete this trial) judged the low-fertility samples to smell more attractive
than unworn gauze pads on 54% (SD = 33%) of the trials, a rate that did
not significantly exceed chance, p = .21. A majority – 67% – of the raters
(N = 91) judged the high-fertility sample to smell more attractive than
the unworn gauze pad. This rate was significantly greater than chance,
p = .001. In general, raters judged the unworn pads to be at around
the midpoint of the scale for pleasantness (M = 4.99, SD = 1.32),
slightly below the midpoint for sexiness (M = 4.01, SD = 1.70), and
below the midpoint for intensity (M = 3.07, SD = 1.56).
3.3. Exploratory analyses
We conducted several exploratory analyses to examine variation in
scent attractiveness among donors and in scent preferences among
raters. First, we examined within- versus between-women variation in
scent qualities among naturally-cycling scent donors. A recent paper
claimed that between-women variation is greater than within
women-variation in women's scent attractiveness, possibly rendering
within-women changes in scent attractiveness across the cycle mean-
inglessly small (see Havlíček et al., 2015). To parse the variation in
† * NC High Fertility
* * HC Users
Sexiness Intensity
scent ratings into between- and within-women portions, we conducted
three two-level multilevel models, with scent samples at Level 1 nested
within women at Level 2. Fertility (1 = high, 0 = low) was a Level-1
predictor. The outcome variable was mean scent rating (pleasantness,
sexiness, or intensity), collapsing across raters (i.e. treating donors as
the unit of analysis). Contradicting the notion that within-women vari-
ation in scent attractiveness is subtle, 71% of the variance in scent pleas-
antness ratings, 73% of the variance in sexiness ratings, and 70% of the
variance in intensity ratings was within-women. Moreover, 19% (pleas-
antness), 18% (sexiness), and 13% (intensity) of this within-women var-
iance was accounted for by whether women were at high or low fertility
within the cycle when they provided their scent samples.
We also conducted several exploratory analyses to examine
variation in scent preferences among scent raters. For example, a re-
viewer suggested that hormonal contraception use could alter female
raters' perceptions of other women's scents. Twenty-one raters in this
study reported using some form of hormonal contraception, 67 report-
ed being naturally-cycling, and three did not answer the question. The
key findings from the full sample of raters were robust across the sub-
samples of naturally-cycling and HC-using raters.
However, do naturally-cycling raters' scent preferences vary de-
pending on their own fertility within the cycle? One previous study
found evidence that naturally-cycling women's responses to other
women's high- versus low-fertility scents might depend on their cur-
rent levels of estrogen and progesterone within the cycle (Woodward
et al., 2015). In the present study, a subsample of 46 naturally-cycling
raters reported regular cycles and provided the start date of their last
menstrual period. We therefore used the forward counting method to
estimate their current cycle day and then assigned continuous actuarial
estimates of fertility based on the values reported by Wilcox, Dunson,
Weinberg, Trussell, and Day Baird (2001). Correlational analyses did
not reveal any significant associations between raters' estimated fertili-
ty and their preference on the forced-choice task for high- over low-
fertility samples or for HC-users' samples over naturally-cycling
women's high-fertility samples (both p values N.10). However,
the higher their estimated fertility was, the less strongly raters preferred
HC-users' samples over low-fertility samples, r = .31, p = .03. We urge
caution in interpreting this finding. As discussed above, the forward
counting method used here is relatively low in validity (see Gangestad
et al., 2015). This finding could be a Type 1 error.
Lastly, because raters were recruited at a Lesbian and Gay Pride fes-
tival, the sample was diverse in terms of sexual orientation. Raters iden-
tified as lesbian (n = 22), bisexual (n = 16), heterosexual (n = 48),
other (n = 4), and not reported (n = 1). This variation afforded us
the unique opportunity to examine possible associations between
 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
women's scent preferences and their sexual orientation, as well as other
aspects of their sexual and relationship histories. The key findings from
the full sample of raters were robust across the subsamples of lesbian,
bisexual, and heterosexual women. Furthermore, correlational analyses
did not reveal any significant associations between women's continu-
ous ratings of their past or present same-sex (versus other-sex) sexual
attraction, fantasy, or behavior as assessed using the Klein Sexual
Orientation Grid (Klein, Sepekoff, & Wolf, 1985) and their preference
for high- over low-fertility scents, HC-using women's samples over
naturally-cycling women's high-fertility samples, or HC-users' samples
over naturally-cycling women's low-fertility samples (all p values N.5).
There also were no significant correlations between women's lifetime
number of female sex partners (after removing one outlier who report-
ed having had 600 partners), lifetime number of sexual relationships
with female partners, lifetime number of romantic relationships with
female partners, length of their longest sexual or romantic relationship
with a women, or having lived with a female partner and their
preference for naturally-cycling women's high- over low-fertility
scents, HC-users' scents over high-fertility scents, or HC-users' scents
over low-fertility scents (all p values ≥.10). Thus, the exploratory
analyses did not reveal any evidence that women's evaluations of
other women's scents depend on their own attraction to or sexual histo-
ries with women.
4. Discussion
A growing body of evidence indicates that men are sensitive to cues
of women's current fertility within the ovulatory cycle, that they typi-
cally perceive cues of high fertility as attractive, and that exposure to
such cues may have downstream consequences for their hormones, mo-
tivations, and behavior. Comparatively little is known about whether
women also are sensitive to cues of fertility in other women, despite
multiple rationales for predicting this (discussed above). The present
findings indicate that, like men, women perceive cues of high fertility
in other women as attractive. Specifically, women rated other women's
high-fertility natural body scents as more pleasant, sexier, and perhaps
less intense than their low-fertility scents. Importantly, because these
findings were robust across analyses treating stimulus donors and
raters as the unit of analysis, we can conclude that these effects are un-
likely to depend on idiosyncratic features of the particular female scent
stimuli included in this study and, instead, are likely to generalize to the
population of cycling female targets and female perceivers.
Given evidence for changes in women's natural body odor across the
cycle, an important question is which hormones mediate these changes.
Estradiol and progesterone fluctuate across the natural ovulatory cycle.
Estradiol levels climb throughout the follicular phase (the first half of
the cycle, approaching ovulation), peak, and then drop just prior to ovu-
lation. Estradiol levels remain relatively low throughout the luteal phase
(the second half of the cycle, after ovulation), increasing slightly in the
mid-luteal phase, and then decreasing until menstruation, when the
cycle begins again. In contrast, progesterone levels are low throughout
the follicular phase, begin to climb just after ovulation, reach their
peak around the middle of the luteal phase, and then decrease until
menstruation. Thus, relatively high estradiol levels (and high estradiol
relative to progesterone levels) mark impending ovulation and the
highest-fertility phase of the cycle, whereas relatively high progester-
one levels (and high progesterone relative to estradiol levels) mark a
low-fertility phase of the cycle.
In one previous study, naturally-cycling women were photographed
and audio-recorded twice within the ovulatory cycle. At a within-
woman (rather than between-women) level, low progesterone – a hor-
mone profile characterizing the follicular, relatively high-fertility phase
of the cycle – predicted both facial and vocal attractiveness. In addition,
high estradiol relative to progesterone – a hormone profile characteriz-
ing the late follicular, highest-fertility phase – predicted vocal attrac-
tiveness (Puts et al., 2013). Thus, estradiol, progesterone, and the ratio
161
of estradiol to progesterone are prime candidates for possible hormonal
mediators of changes across the cycle in naturally-cycling women's
scent attractiveness. We encourage future studies in this area to include
multiple assessments of these hormones in conjunction with assess-
ments of women's scent attractiveness across the cycle.
Another important question is what odorous compounds contribute
to changes in women's scent attractiveness across the cycle. Apocrine
glands are densely clustered in the axillae (underarms). These sweat
glands produce “nutrient-rich” sweat, which feed bacteria living on
the skin and hair follicles. Axillary (underarm) sweat is odorless when
first secreted; however, it is quickly broken down by bacteria into a
wide variety of volatile (and, hence, scented) compounds, which we
perceive as natural body odor. Importantly, the apocrine glands appear
to be hormonally regulated. These glands become active at puberty (de-
spite being present even before birth; see Hays, 2003) and express an-
drogen and estrogen receptors (see Beier, Ginez, & Schaller, 2005).
Although it was outside of the scope of the present investigation, we en-
courage future research to examine changes across the cycle in the
chemical composition of women's axillary odor that might underpin
changes in scent attractiveness. In addition, we note that past research
has found evidence for changes across the cycle in the attractiveness
of women's vulvar/vaginal scents, as well (e.g., Cerda-Molina et al.,
2013). However, underarm/torso scents and vulvar/vaginal scents
appear to have different chemical compositions (e.g., Doty, 1981).
Therefore, we also encourage future research to examine changes across
the cycle in the chemical composition of women's vulvar/vaginal scents.
Are changes across the cycle in women's scent attractiveness of the
magnitude reported in this study likely to have a meaningful effect on
perceptions of attractiveness in real life? One important observation in
the present study is that approximately 13% of the total variance in
women's scent attractiveness (pleasantness and sexiness) was
accounted for by their current position within the ovulatory cycle. This
suggests that changes across the cycle could have a meaningful impact
on attractiveness perceptions — perhaps especially in close relation-
ships or other contexts involving regular, close contact with cycling
women. Furthermore, even the highly valid methods used to estimate
women's fertility in the present study are imperfect. Thus, the effect
sizes we report here could underestimate true fertility effects on
women's scent attractiveness.
On the other hand, in this study, women evaluated samples of
women's high- and low-fertility body odors side by side in a controlled
laboratory setting, and the odor samples themselves were collected
using strict protocols to minimize possible contamination by nonhuman
scents. It is possible that in more a naturalistic setting, day-to-day vari-
ation in women's diet and activities, the weather, ambient odors, and so
on would introduce so much “noise” as to render subtle fertility-related
changes in women's scent attractiveness undetectable. Then again, fer-
tility effects on perceptions of women's scent attractiveness might have
been more pronounced among our ancestors (or modern populations
that differ from the U.S. in this respect), for whom products for masking
body scent were less widely available and who therefore had more day-
to-day exposure to others' natural body odors. To provide a clearer pic-
ture of the magnitude of these effects and their possible moderation by
environmental factors, future research should examine changes across
the cycle in women's body odor attractiveness in a variety of contexts.
This research also raises questions about how women respond hor-
monally and behaviorally to cues of fertility in other women. To date,
two studies have directly addressed this question (Maner & McNulty,
2013; Woodward et al., 2015). In the first of these studies, women
(n = 25) smelled a T-shirt worn at high fertility or at low fertility by
one of four naturally-cycling women (Maner & McNulty, 2013).
Women who smelled a high-fertility T-shirt maintained testosterone
levels over the course of the study, whereas women who smelled a
low-fertility T-shirt exhibited declining testosterone levels over the
course of the study. We note that, because each participant smelled a
single T-shirt from just one of four donors, the researchers could not
162 K.A. Gildersleeve et al. / Evolution and Human Behavior 38 (2017) 155–163
conduct analyses treating donors as the unit of analysis. Therefore, it re-
mains unknown whether other samples of women's high- and low-
fertility scents would have had the same effect on female perceivers.
Nonetheless, the authors' proposal that the greater levels of testoster-
one they observed among women exposed to cues of high fertility in
other women might serve to facilitate competitive aggression is worthy
of further empirical investigation.
In the second study (also discussed above), naturally-cycling
women smelled a set of T-shirts worn by women at high fertility or
low fertility and rated their pleasantness (Woodward et al., 2015).
They then reported how they would respond to hypothetical transgres-
sions committed by young or elderly women (e.g., a [young/elderly]
woman swerving in front of them on the freeway). Young but not elder-
ly women were expected to be perceived as potential rivals. Compared
with women who smelled low-fertility T-shirts, women who smelled
high-fertility T-shirts exhibited marginally less aggressive responses to
young female transgressors relative to elderly transgressors. This find-
ing ran counter to the prediction that women would direct more com-
petitive aggression specifically at plausible rivals exhibiting cues of
high fertility (Maner & McNulty, 2013). Furthermore, there was no
overall effect of condition on women's testosterone; women who
smelled high-fertility T-shirts did not exhibit greater testosterone levels
than did women who smelled low-fertility T-shirts. However, the find-
ings suggested that women with relatively high estradiol levels and low
progesterone levels – a hormone profile indicating that they themselves
might be at high fertility within the cycle – responded to other women's
high-fertility scents (versus low-fertility scents) with greater testoster-
one than did women with hormone profiles indicating lower fertility.
One implication, then, is that women's hormonal and behavioral re-
sponses to cues of fertility in other women might depend on their
own fertility within the cycle.
Both of the above studies were limited by small samples of stimulus
donors and by relying on error-prone self-reported dates of last
menstrual onset to estimate each stimulus donor's position within the
ovulatory cycle. Nevertheless, these studies take important first steps
toward understanding the possible impacts of fertility cue detection
on female–female social interactions in humans.
As discussed above, women's sensitivity to cues of other women's
current fertility within the ovulatory cycle could have arisen via
several different evolutionary paths. Women's tendency to perceive
cues of high fertility in other women as attractive could reflect adap-
tations favored by selection specifically for that function, byproducts
of adaptations favored by selection for some other function, or non-
functional vestigial mechanisms passed down from an ancestral
species. Given the availability of mathematical modeling and phylo-
genetic techniques for comparing the relative likelihood of different
evolutionary paths, this is another feasible and important direction
for future research.
A subsidiary aim of this study was to conduct exploratory analyses to
compare women's evaluations of naturally-cycling women's high- and
low-fertility scents with those of hormonal contraception-using
women. We found that women rated hormonal contraceptive-using
women's body odor as more pleasant and sexier but less intense than
naturally-cycling women's low-fertility and high-fertility body odor,
though the latter set of effects reached statistical significance only
when raters were treated as the unit of analysis. These findings are in
contrast to the null results reported by a previous study comparing
evaluations of the attractiveness and intensity of scent samples
provided by naturally-cycling versus hormonal contraceptive-using
women (described above; Kuukasjärvi et al., 2004). We emphasize
that the present findings should be considered preliminary, given
their between-women, correlational nature and the small sample of
hormonal contraceptive users. The hormonal contraceptive users in
this study might have differed from the naturally-cycling women in a
variety of ways prior to beginning hormonal contraception, and such
pre-existing differences could create the illusion of HC effects.
We encourage future work to carefully examine the possible effects
of hormonal contraception, including the effects of different formula-
tions of hormonal contraception. It is not yet known precisely how ethi-
nyl estradiol or the many synthetic progestins found in hormonal
contraceptives compare with endogenous hormones in terms of their
effects on women's attractiveness. For example, ethinyl estradiol, the
synthetic estrogen found in most combined oral hormonal contracep-
tives, is a more potent estrogen than endogenous estradiol (Blair et al.,
2000). It is also known to decrease free testosterone by increasing
sex hormone binding globulin (e.g., see Hugon-Rodin, Gompel, & Plu-
Bureau, 2014). In addition, synthetic progestins vary widely in their
binding affinities for different steroid hormone receptors and therefore
can have progestogenic, androgenic, anti-androgenic, and/or estrogenic
effects (see Sitruk-Ware, 2004). Given this wide pharmacological varia-
tion, we think it is highly unlikely that all hormonal contraceptives have
the same effect on women's attractiveness (or other aspects of their
psychology and behavior). We encourage future research to examine
which hormonal contraceptives are associated with a more low
fertility-like state, high fertility-like state, or unique state unlike either.
Ultimately, true experiments in the form of double-blind randomized
trials with larger samples of women are needed to establish whether as-
sociations between women's attractiveness and use of hormonal con-
traception or their use of one formulation of hormonal contraception
versus another reflect true causal effects of hormonal contraception.
In sum, our findings support a potentially important role for scent in
women's perceptions of other women, raising the intriguing question of
how fertility cue detection might affect female–female social interac-
tions in competitive and other contexts. This study also highlights a
need for rigorous research examining possible hormonal mediators of
shifts in women's attractiveness across the cycle and, relatedly, the im-
pact of different formulations of hormonal contraception on women's
attractiveness, social behavior, and relationships. More generally, this
study adds to a rich body of evidence indicating that human social
behavior – like that of virtually all other mammals – is influenced by
the female ovulatory cycle.
Supplementary materials
Supplementary data to this article can be found online at doi: http://
dx.doi.org/10.1016/j.evolhumbehav.2016.08.003
Acknowledgments
We are grateful to the many research assistants who assisted with
the scent rating sessions and to Kristine Chua for her additional
assistance with data organization and entry. This research was supported
by a grant from the American Institute for Bisexuality (AIB) and a UCLA
Academic Senate grant awarded to Martie Haselton.
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