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Archives of Sexual Behavior pp350-aseb-365185 December 28, 2001 8:29 Style file version July 26, 1999

Archives of Sexual Behavior, Vol. 31, No. 1, February 2002, pp. 51–62 (°
C 2002)

Sexual Partner Preference in Female Japanese Macaques

Paul L. Vasey, Ph.D.1

Received March 29, 2001; revision received August 28, 2001; accepted September 1, 2001

Whether animals ever exhibit a preference for same-sex sexual partners is a subject of debate. Japanese
macaques represent excellent models for examining issues related to sexual preference in animals be-
cause females, in certain populations, routinely engage in both heterosexual and homosexual behavior
over the course of their life spans. Multiple lines of evidence indicate that female homosexual behavior
in Japanese macaques is a sexual behavior, not a sociosexual one. Additional evidence indicates that
female Japanese macaques do not engage in homosexual behavior simply because acceptable male
mates are unavailable or unmotivated to copulate. Patterns of sexual partner choice by female Japanese
macaques that are the focus of intersexual competition indicate that females of this species choose
same-sex sexual partners even when they are simultaneously presented with a motivated, opposite-
sex alternative. Thus, in some populations of Japanese macaques, females prefer certain same-sex
sexual partners relative to certain male mates, and vice versa. Taken together, this evidence suggests
that female Japanese macaques are best characterized as bisexual in orientation, not preferentially
homosexual or preferentially heterosexual.

KEY WORDS: homosexual behavior; sexual preference; Japanese macaques.

Most researchers would agree that same-sex interac-
tions involving genital contact are widespread among an-
imals and can occur independent of any hormonal or neu-
rological manipulations (Adkins-Regan, 1988; Bagemihl,
1999; Dixson, 1998; Nadler, 1990; Vasey, 1995; Wallen
& Parsons, 1997). More contentious, however, is the issue
of whether these interactions ever reflect an actual pref-
erence for same-sex sexual partners (cf. Bagemihl, 1999;
Dixson, 1998; Wallen & Parsons, 1997).
This debate is actually fueled by a number of more
fundamental disagreements about the nature of animal
sexual behavior. First, are same-sex “sexual” interactions
in animals really sexual? Or, are they merely sociosexual
interactions that are devoid of sexual motivation and en-
acted to mediate some sort of adaptive social goal? Second,
are same-sex sexual interactions in animals manifested
in the presence of opposite-sex alternatives? Or, do they
simply occur because opposite-sex partners are difficult
to access? Third, do individuals actually choose same-
sex sexual partners when simultaneously presented with
1 Department of Psychology and Neuroscience, University of Lethbridge,
4401 University Drive, Lethbridge, Alberta, Canada T1K 3M4; e-mail:
paul.vasey@uleth.ca.
opposite-sex alternatives? Or, are animals with bisexual
potential always preferentially heterosexual? Before one
can address the larger question of whether animals ever
exhibit a preference for same-sex sexual partners, these
more fundamental issues must be addressed.
In this paper, I review my previous research on female
homosexual behavior in Japanese macaques (Macaca fus-
cata) that bears directly on the questions outlined above.
I then address how female sexual preference in Japanese
macaques might best be conceptualized on a species-wide
level.
METHODS
Subjects
Study subjects were housed at the Université de
Montréal’s Laboratory of Behavioral Primatology (est.
1992) near St-Hyancinthe, Québec. The group consisted
of three unrelated matrilines named A, B, and C in ac-
cordance with their overall dominance relationships (A >
B > C). The founding members of the study group origi-
nated from the Arashiyama East (Japan) troop of Japanese
macaques that was translocated to Texas in 1972 (Fedigan,

51
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C 2002 Plenum Publishing Corporation
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52
1991). The monkeys lived in five indoor rooms (25 ×
15.5 × 3.6 m3 ) and two outdoor enclosures (17 × 16 ×
4 m3 ). The rooms and enclosures were furnished with
swinging and climbing devices and loose enrichment ob-
jects (e.g., branches, pails, coconuts). Animals were fed
daily with a mixture of grains, monkey chow, fruit, and
vegetables, which was distributed in deep wood-chip litter
covering the indoor floors. Water was available at will.
During the 1993–94 mating season, the study group
consisted of 37 individuals (18 sexually active females,
ages 3.5–23 years; 5 sexually active males, ages 4.5–
9.5 years; 14 sexually immature individuals). During the
1994–95 mating season, the study group consisted of
38 individuals (1 sexually inactive female, age 24 years;
16 sexually active females, ages 3.5–24 years; 5 sexually
active males, ages 4–10.5 years; 16 sexually immature
individuals).
General Data Collection
All observations were recorded on paper in 30-min
time blocks. Group members were individually recog-
nized. During observation periods, the monkeys occupied
the two largest indoor rooms of the laboratory and the two
outdoor enclosures. Observations took place from a raised
mezzanine that looked out onto all the rooms and enclo-
sures occupied by the monkeys. As such, all members of
the study group were readily observable regardless of the
room or enclosure they occupied.
Data Collection During the 1993–94 Mating Season
Observations were made daily from dawn to dusk
(approximately 7:00–16:30) during the mating season be-
tween 20 September 1993 and 11 March 1994, excluding
a 4-day period from 25 to 29 December. All occurrences
of sexual activity (i.e., mounting, consortship formation,
sexual solicitations) were recorded during this period. Fo-
cal data and scan sample data (Altmann, 1974) were col-
lected daily for female Japanese macaques engaged in ho-
mosexual consortships, between 17 November 1993 and
13 February 1994. Focal and scan sample data were col-
lected for both consort partners simultaneously because
they were almost always interacting together. When two
homosexual consortships occurred simultaneously, I ob-
served the one for which I had the least data. I collected
129 hr of focal data. For every minute of focal data, I
recorded one scan sample of both consort partners.
Before the 1993–94 mating season, (23 March–
30 April, 1993, 45 hr; 29 July–3 August, 1993, 10 hr;
N = 55 hr) baseline data collection focused on recording
all occurrences of dyadic aggression and interventions.
Vasey
Two types of baseline data were collected following
the 1993–94 mating season. First, between 7 March and
15 April 1994, baseline data collection focused on record-
ing all occurrences of dyadic aggression and interventions
by members of the study group (N = 45 hr). Second, be-
tween 21 March and 4 May 1995, baseline data collec-
tion focused on recording all occurrences of grooming
between nonkin females (N = 36 hr). Observations for
both the pre- and postconsortship baseline periods took
place between 9:00 and 17:00.
Unless otherwise stated, the data discussed in this
paper are derived from the 1993–94 mating season.
Data Collection During the 1994–95 Mating Season
During the 1994–95 mating season, subjects were ob-
served during a baseline and experimental period. Base-
line and experimental observations were conducted on
identical days across separate 4-week periods. Thus, data
were collected during December 1994 (baseline period)
and January 1995 (experimental period) on identical days
(2, 5–24; N = 21 days/4-week period). Three hours of
observation took place every day between 9:00 and 13:00
during which all occurrences of consortship formation and
sexual solicitations were recorded. A total of 126 hr of be-
havioral sampling data were collected.
In the afternoon of January 1, 1995, an experimen-
tal subgroup was created by removing four of the group’s
five sexually active males. To control for the confound-
ing effects of male dominance acquisition over females,
the study group’s alpha male was chosen as the sole male
to remain in the experimental subgroup. Removal of the
other four males was accomplished in a few minutes by
channeling the monkeys through a network of rooms and
sliding doors. The resulting experimental and all-male
subgroups were physically and visually separated from
each other.
Definitions
Homosexual consortships occurred when two fe-
males engaged in series mounting (three or more mounts
within a 10-min period). These consortships were deemed
to have terminated if the two female partners were sepa-
rated by a distance of more than 1 m and exhibited no
mounting for 10 min. Homosexual behavior is used to
collectively refer to homosexual consortships, mounting,
and sexual solicitations.
Double foot-clasp mounts involved the mounter gras-
ping with her feet just above the mountee’s ankles and with
her hands on the mountee’s back. As such, the mounter’s
groin region was in contact with the mountee’s perineum.
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Sexual Preference in Female Japanese Macaques
Sitting mounts occurred when the mounter sat on the
mountee’s back jockey-style, while grasping the moun-
tee’s upper back with her hands and the mountee’s lower
back with her feet. In this position, the mounter’s perineum
was in contact with the mountee’s lower back. Standing
mounts involved the mounter standing bipedally with her
feet on the ground and her knees slightly bent, while gras-
ping the mountee’s lower back with her hands. As such, the
mounter’s groin region was in contact with the mountee’s
perineum. Reclining mounts occurred when the mounter
lay ventrally on the mountee’s back, using her feet to grasp
the mountee’s legs just above the ankles and her hands
to grasp fur on the mountee’s upper back. Each of these
four different types of mounts could be performed with or
without pelvic thrusts by the mounter for a total of eight
different mount types.
Hindquarter presentations were performed by po-
tential mountees who stood quadrupedally with their arms
and legs flexed and their perineum oriented towards the po-
tential mounter. Back presentations were performed by po-
tential mountees who sat with their forearms slightly bent
and their backs inclined and oriented towards the potential
mounter. Hands-on-hindquarters solicitations were per-
formed by potential mounters who grasped or placed both
hands on the hindquarters of the potential mountees.
Dominance relationships were determined by the di-
rection of submissive behavior(s), that is, displacements,
fear grimaces, rapid fleeing, piercing screams, and de-
fensive hunching. Submission had to be unidirectional
between two individuals for the recipient to be consid-
ered dominant. Dyads for which submission was bidi-
rectional were considered to have ambiguous dominance
relationships.
Interventions referred to the intrusion of an individual
(supporter) into an ongoing conflict between two oppo-
nents, during which the intruding animal supported one of
the opponents (recipient of support) against the other (tar-
get). Supporters and recipients of support were referred to
as allies and were said to have formed alliances. Oppor-
tunities for intervention included all incidences of dyadic
aggression, except displacements, during which one indi-
vidual could intervene to support another against a third.
Conservative interventions occurred when the targets of
aggression ranked below the supporter and the recipient of
support. Bridging interventions occurred when the targets
of aggression ranked below the supporter, but above the
recipient of support. Opportunities for rank changes oc-
curred when an individual participated in an intervention
against a dominant target, or one with whom they shared
an ambiguous dominance relationship.
Females from different matrilines were considered
to be nonkin, as were aunt–niece dyads. Experimental
Style file version July 26, 1999
53
and observational evidence indicates that female Japanese
macaques treat each other as nonkin, in terms of sexual
and nonsexual affiliation, when their level of relatedness
is less than r = .25 (Chapais, Gauthier, Prud’homme, &
Vasey, 1997). This evidence suggests that female Japanese
macaques categorize individual as nonkin when they share
a level of relatedness of r < .25 (e.g., aunt–nieces).
Homosexual Behavior in Female Japanese Macaques
Japanese macaques represent excellent models for
examining issues related to sexual preference in animals
because females, in certain populations, routinely engage
in both heterosexual and homosexual behavior over the
course of their life spans. Female homosexual activity
occurs in both captive (Chapais et al., 1997; Chapais &
Mignault, 1991; Lunardini, 1989; Rendall & Taylor, 1991;
Vasey, 1996, 1998; Vasey, Chapais, & Gauthier, 1998;
Vasey & Gauthier, 2000) and free-ranging (Eaton, 1978;
Enomoto, 1974; Fedigan & Gouzoules, 1978; Gouzoules
& Goy, 1983; Takahata, 1982; Wolfe, 1984) populations
of this species. As such, these interactions cannot be dis-
missed as abnormal products of captivity.
Female homosexual behavior in Japanese macaques
occurs during temporary, but exclusive, associations with
same-sex sexual partners. These relationships, termed ho-
mosexual consortships, can be brief, lasting less than
1 hr, or they can continue for over a week (Wolfe, 1984).
During homosexual consortships, females solicit
each other to mount, and to be mounted, using vocaliza-
tions and a variety of postural and facial gestures. Sexual
solicitations include pushing, hitting, grabbing, slapping
the ground, head bobbing, screaming, lip quivering, body
spasms, and intense gazing. A female can specifically re-
quest to be mounted by presenting her inclined back or
her hindquarters to a potential mounter. Conversely, a fe-
male can specifically request to mount another female by
placing her hands on the other’s hindquarters (Vasey et al.,
1998).
Single mounts between females are rarely observed.
Instead, multiple or series mounting between consort part-
ners is the norm. During these interactions, females mount
each other in a bidirectional manner and routinely employ
up to eight different types of mounting postures (Vasey
et al., 1998; see definitions). During mounts, mounters of-
ten thrust against mountees, who in turn, commonly reach
back to grasp the mounter and gaze intently into her eyes.
Mounting between consort partners and the accom-
panying sexual solicitations can continue uninterrupted
for well over an hour. Thus, sexual behavior is clearly
the defining feature of homosexual consortships. It would
be erroneous, however, to characterize these relationships
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54
solely in terms of sex. Between bouts of sexual activity,
partners exhibit highly synchronized activity: huddling
and sleeping together, as well as grooming, following,
and defending each other for prolonged periods of time
(Gouzoules & Goy, 1983; Vasey, 1996; Wolfe, 1984).
Partners will also forage side-by-side during homosex-
ual consortships, although this form of cofeeding is not a
prominent feature of these relationships.
The frequency of female homosexual behavior varies
both within and between Japanese macaque populations.
In some populations, the behavior appears to be non-
existent. In others, every sexually mature female engages
in homosexual behavior, and they do so frequently. For
example, during the 1993–94 mating season, every sex-
ually mature female in my study group engaged in ho-
mosexual behaviour. Moreover, the majority of sexual
consortships formed (N = 95) during this period were
homosexual (55%), not heterosexual (45%). During these
consortships, females (N = 14) solicited their partners for
sex, on average, 28 times per hour of observation, and
they mounted each other, on average, 31 times per hour
of observation (N = 129 observation hours; Vasey, 1996;
Vasey et al., 1998). Some researchers have argued that this
intergroup variation can be explained in terms of “proto-
cultural” differences in sexual “traditions” among pop-
ulations (e.g., Fedigan, 1982; Itani & Nishimura, 1973;
Stephenson, 1973).
Is Homosexual Behavior a Sociosexual Adaptation?
Same-sex interactions in animals involving genital
contact have traditionally been interpreted as “sociosex-
ual” behaviors (Dixson, 1998; Wickler, 1967). Within
ethological circles, sociosexual behaviors are defined as
behaviors that are sexual in terms of their outward form,
but enacted to facilitate some type of adaptive social goal.
Despite over 40 years of intensive research on Japanese
macaques, there is not a single study demonstrating that
female homosexual behavior in this species is a sociosex-
ual adaptation. On the contrary, several studies indicate
that female Japanese macaques do not use homosexual be-
havior to facilitate sociosexual goals (Gouzoules & Goy,
1983; Vasey, 1995, 1996, 1998; Vasey et al., 1998).
Dominance Demonstration and the Regulation
of Social Tension
Same-sex mounting interactions have long been con-
ceptualized as ritualized gestures that the participants use
to communicate about their dominance relationships,
Vasey
thereby reducing the threat of incipient aggression as-
sociated with social tension (Wickler, 1967). According
to this “rank demonstration hypothesis,” mounting is an
expression of dominance, whereas allowing oneself to
be mounted expresses subordinate status relative to the
mounter (Wickler, 1967). Consequently, a test of this hypo-
thesis should demonstrate that the roles individuals
adopt during same-sex mounting interactions are differ-
entiated on the basis of their rank. In addition, mounting
interactions among same-sex individuals should be par-
ticularly common at the outset of aggressive interactions,
at which time reiterating an acceptance of the dominance
status quo should function to reduce the threat of escalated
fighting.
Several lines of evidence indicate that female
Japanese macaques do not use homosexual behavior to
communicate about their dominance relationships in the
manner predicted by the rank demonstration hypothesis
(for further details, see Vasey et al., 1998). During ho-
mosexual consortships, dominant and subordinate part-
ners typically take turns mounting and being mounted,
in a bidirectional manner. Of the 3,945 mounts observed,
56.9% were performed by dominant consort partners, and
43.1% were performed by subordinate consort partners.
This difference was not significant (Wilcoxon matched-
pairs signed-ranks test, N = 21, Z 0 = 0.2, P = .85).
Even when the catch-all category of “mounts” was sub-
divided into eight different mount types that were then
analyzed separately, it was found that neither partner was
significantly more or less responsible for mounting or be-
ing mounted (Table I).
In keeping with the spirit of the rank demonstra-
tion hypothesis, one could also predict that solicitations
to mount, and to be mounted, should be performed dif-
ferentially by dominant and subordinate consort partners.
Dominant consort partners should perform hands-on-
hindquarters solicitations more often because they func-
tion as requests to mount and prompt the receivers to
prepare to be mounted. However, dominant consort part-
ners did not perform hands-on-hindquarters solicitations
significantly more often than their subordinate counter-
parts. Of the 720 hands-on-hindquarters solicitations ob-
served, 35.8% were performed by dominant consort part-
ners, whereas 64.2% were performed by the subordinate
partners (Wilcoxon test, N = 20, Z 0 = 1.6, P = .1).
Similarly, one would predict that subordinate con-
sort partners should perform hindquarter presentations and
back presentations more often than their dominant coun-
terparts, because these signals function as requests to be
mounted. Subordinate consort partners did not, however,
perform hindquarter or back presentations significantly
more often than their dominant partners. Of the 1,225
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Sexual Preference in Female Japanese Macaques 55

Table I. Comparison of Dominant and Subordinate Consort Partners’ Behaviour During Mounting Interactions
Using Wilcoxon Matched-Pairs Signed-Ranks Test

Total Dominant Subordinate N Z 0 /T P

Double-foot clasp mount, thrusting 1578 1112 466 20 1.80 .07

Double-foot clasp mount, no thrusting 1043 749 294 20 1.70 .09
Sitting mount, thrusting 749 159 590 12 53 .15
Sitting mount, no thrusting 297 105 192 14 55.5 .88
Standing mount, thrusting 40 24 16 10 39 .28
Standing mount, no thrusting 84 49 35 12 44 .73
Reclining mount, thrusting 30 11 19 6 11.5 .92
Reclining mount, no thrusting 124 35 89 9 28 .57

Note. Prime (0 ) indicates Z value corrected for ties. All other values are T values.

hindquarter presentations observed, 38% were performed Alliance Formation

by dominant consort partners, whereas 62% were per-
formed by the subordinate partners (Wilcoxon test, N =
20, Z 0 = 1.6, P = .1). Of the 463 back presentations ob-
served, 46% were performed by dominant consort part-
ners, and 54% were performed by subordinate consort
partners (Wilcoxon test, N = 17, Z 0 = 0.2, P = .81).
In accordance with the rank demonstration hypoth-
esis, one would further predict that subordinate consort
partners would facilitate mounts by their dominant part-
ners more often than the reverse, by clasping the domi-
nant mounter during mounts-in-progress. However, nei-
ther partner was significantly more or less responsible
than the other for clasping during mounts-in-progress. Of
the 1,248 incidences of clasping observed during mounts-
in-progress, 62.6% were performed by dominant consort
partners and 37.4% by subordinate partners (Wilcoxon
test, N = 20, Z 0 = 1.8, P = .08).
Additional evidence suggests that female Japanese
macaques do not use homosexual behavior to control in-
cipient aggression as predicted by the rank demonstration
hypothesis (for further details, see Vasey et al., 1998). In
Japanese macaques, a close temporal relationship does
not exist between homosexual mounting and escalating
social tension associated with incipient aggression. Only
0.004% of all mounts (N = 2,244) and 0.002% of hands-
on-hindquarters solicitations (N = 462) by dominant
partners occurred within 1 min of an aggressive interac-
tion between consort partners. Similarly, only 0.02% of
back presentations (N = 250) and 0.005% of hindquarter
presentations (N = 760) by subordinate partners occurred
within 1 min of an aggressive interaction between consort
partners.
Taken together, these multiple lines of evidence sug-
gest that female Japanese macaques do not use homo-
sexual behavior to communicate about their dominance
relationships or control social tension associated with in-
cipient aggression.
A number of primate species appear to use homosex-
ual behavior as a mechanism for alliance formation (e.g.,
Fairbanks, McGuire, & Kerber, 1977; Idani, 1991; Parish,
1994; Smuts & Watanabe, 1990). Among female Japanese
macaques, homosexual consortships and alliance forma-
tion are causally related (for further details, see Vasey,
1996). Dominant and subordinate consort partners inter-
vene for each other significantly more often during ho-
mosexual consortships than either before (Wilcoxon test
for dominant partners, N = 15, Z 0 = 3.4, P < .01; sub-
ordinate partners, N = 16, Z 0 = 3.5, P < .001) or after
(Wilcoxon test for dominant partners, N = 16, Z 0 = 3.4,
P < .01; subordinate partners, N = 16, Z 0 = 3.5, P <
.001) these relationships. During homosexual consort-
ships, females (N = 9) intervened significantly more for
their sexual partners (χ = 1.33 interventions/hr) than
for other sexually mature, nonkin females (χ = 0.09
interventions/hr; Wilcoxon test: dominant partners, N =
16, Z 0 = 3.57, P < .001; subordinate partners, N = 17,
Z 0 = 4, P < .001). As such, the tendency for consorting
females to intervene more for each other did not reflect
a generalized motivation to intervene for any sexually
mature, nonkin females. Despite this causal relationship,
patterns of partner choice and affiliation between consort
partners suggest that females were not engaging in ho-
mosexual consortships for the express purpose of alliance
formation (for further details, see Vasey, 1996, 2000b).
If the primary reason for engaging in homosexual
consortships is to mediate alliance formation, then two
key behavioral patterns should be expected to occur. First,
higher ranking females should be preferred as consort part-
ners because they represent more powerful allies and, as
such, are able to challenge targets that they dominate but
that their subordinate consort partners do not. By provid-
ing support against targets that rank above their subor-
dinate partners, but below themselves, dominant consort
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56
partners can help their partners increase in dominance
temporarily, at little risk to themselves (for data in sup-
port of this point, see Vasey, 1996, and below). Females
should, thus, prefer to consort with dominant partners,
and competition for such partners should ensue. The most
successful competitors should also be the most dominant
ones. Given the constraints against the formation of con-
sortships with close kin (Chapais et al., 1997; Chapais &
Mignault, 1991; Wolfe, 1984), this should have resulted
in a concentration of consortships between members of
the two highest ranking matrilines in the study group (i.e.,
A–B consortships vs. A–C or B–C consortships). Keep-
ing in mind that consortships between females were pos-
sible only if two individuals were receptive on the same
days, a possible 42 A–B consortships could have poten-
tially been formed during the 1993–94 mating season, of
which 42.9% actually occurred. Of a possible 12 A–C con-
sortships, 41.7% actually occurred. Of a possible 13 B–C
consortships, 38.5% actually occurred. Thus, nonkin in-
dividuals of higher rank (A–B) did not form consortships
together significantly more often than other types of con-
sorthips (i.e., A–C, B–C; χ 2 = 0.066, df = 2; P > .95).
There was also no significant difference in the number of
days over which each of the three different types of con-
sortships took place (χ for A–B dyads = 5 days; χ, A–C
dyads = 4.4 days; χ , B–C dyads = 5.8 days; Kruskal-
Wallis test: N = 28, H 0 = 0.1, df = 2, P = .93).
Second, in light of the dominance-related benefits
they can derive from homosexual consortships, subordi-
nate partners should be more responsible for showing the
affiliative behaviors that maintain these associations. This
prediction was not supported, however. During homosex-
ual consortships, the overwhelming majority of affilia-
tion flowed in a remarkably bidirectional manner, with no
statistically significant differences between dominant and
subordinate consort partners (Chapais & Mignault, 1991;
Vasey, 1996; Vasey et al., 1998). For example, 49% of
approaches (N = 2386) were performed by dominant
members of the consortships and 51% by their subor-
dinate partners (Wilcoxon test, N = 21, Z 0 = 0.82, P =
.41). Similarly, 45.2% of sexual solicitations (N = 3649)
were performed by dominant consort partners and 54.8%
by their subordinate partners (Wilcoxon test, N = 21, Z 0 =
0.3, P = .77). In addition, dominant females intervened
in support of their subordinate consort partners during
29.2% of the opportunities to do so (N = 411), whereas
subordinate females intervened for their dominant consort
partners during 8.7% of available opportunities (N = 601;
Wilcoxon test, N = 16, Z 0 = 1.4, P = .16). The sole dif-
ference in affiliative behavior involved grooming, which
the dominant consort partners performed significantly
more often than their subordinate counterparts. Out of
Vasey
1,328 instantaneous scan samples during which groom-
ing was recorded, dominant females groomed their sub-
ordinate consort partners 86.2% of the time, and subor-
dinate partners groomed 13.8% of the time (Wilcoxon
test, N = 18, Z 0 = 3.7, P = .001). Thus, dominant con-
sort partners were actually more responsible for maintain-
ing homosexual consortships than were their subordinate
partners.
In sum, although homosexual consortships and al-
liance formation are causally related, patterns of partner
choice and affiliation between consort partners suggest
that females were not engaging in these relationships for
the express purpose of alliance formation. As such, associ-
ations with useful allies during homosexual consortships,
let alone powerful ones, would be variable at best. This
suggests that homosexual behavior was not designed by
natural selection as a sociosexual adaptation for alliance
formation.
Acquisition of Alloparental Care
Research on various bird species, particularly gulls
and terns, suggests that females will form pair-bonds with
other females to obtain alloparental care for their depen-
dant offspring (for reviews, see Bagemihl, 1999; Diamond,
1989). Several lines of evidence suggest that female
Japanese macaques do not engage in homosexual con-
sortships to obtain alloparental care for their immature
offspring (for further details, see Vasey, 1998).
Grooming and alliance support are the most overt ex-
pressions of nonsexual affiliation in Japanese macaques,
but these activities did not characterize interactions be-
tween females and their consort partners’ immature off-
spring. For example, consorting females (N = 17) were
within arm’s reach of their same-sex partners’ immature
offspring during 9.7% of the scan samples in which they
were observed (N = 6,190 scan samples). Despite this
proximity, they virtually never groomed these immature
individuals (0.0005% of scan samples).
During homosexual consortships, females (N = 16)
had various opportunities to intervene in support of their
same-sex consort partners’ immature offspring when the
latter were involved in conflicts (N = 134 opportunities
for support, χ = 8.4). None of the females supported their
partners’ immature offspring in this manner. It is, however,
striking that during 10.5% of these conflicts, females in-
tervened against their partners’ immature offspring (χ =
0.88). This difference was significant (Wilcoxon test, N =
6, T = 21, P < .05). These interventions occurred dur-
ing 42.9% of all homosexual consortships for which one
or both partners had immature offspring (N = 14), and
37.5% of homosexually consorting females (N = 16)
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Sexual Preference in Female Japanese Macaques
intervened against their partners’ immature offspring
when given the opportunity.
In addition, females aggressed their partners’ imma-
ture offspring significantly more often during homosex-
ual consortships (N = 73 aggressive interactions) than
either before (N = 22 aggressive interactions; Wilcoxon
test, N = 10, T = 47, P < .05) or after (N = 15 aggres-
sive interactions; Wilcoxon test, N = 13, T = 88.5, P =
.001). These aggressive interactions occurred during
85.7% of all homosexual consortships for which one or
both partners had immature offspring (N = 14), and
70.6% of homosexually consorting females aggressed
their partners’ immature offspring at least once (N = 17,
χ = 4).
In sum, there was no evidence that female Japanese
macaques provided alloparental care to the immature off-
spring of their same-sex consort partners, and this was not
for lack of opportunities to do so. In fact, the relationship
between females and their same-sex consort partners’ im-
mature offspring could best be described as agonistic, not
alloparental, in nature. This points to the conclusion that
female homosexual behavior in Japanese macaques has
not been designed by natural selection as a sociosexual
adaptation for the acquisition of alloparental care from
same-sex sexual partners.
Acquisition of Male Mates
Parker and Pearson (1976) proposed that female ho-
mosexual behavior functions to increase the reproductive
success of the mounting female. By mimicking the copu-
latory pattern of rival males, Parker and Pearson argue that
the mounting female can attract male sexual partners and
increase her chances of insemination. If the primary rea-
son for engaging in homosexual consortships is to attract
male mates, three key behavioral patterns should be ex-
pected to occur. First, female homosexual behavior should
only occur in the presence of males. Second, female ho-
mosexual behavior should only occur when the mounting
female is fertile. Third, mounting females should cease
homosexual mounting after receiving sexual solicitations
from a male.
Several lines of evidence indicate that female
Japanese macaques do not use homosexual behavior to at-
tract male mates. First, consorting females often attempt
to spatially and visually separate themselves from other
group members (Gouzoules & Goy, 1983). Second, fe-
male homosexual behavior in female Japanese macaques
frequently occurs following conception, when the issue
of insemination is moot (Fedigan & Gouzoules, 1978;
Gouzoules & Goy, 1983; Vasey, 1998; Wolfe, 1984). Third,
females engaged in homosexual consortships tend to
Style file version July 26, 1999
57
ignore or even threaten males who solicit them for sex
(Vasey, 1998; Wolfe, 1984).
Taken together, the available evidence indicates that
female Japanese macaques do not engage in homosexual
behavior to attract male mates.
Is Homosexual Behavior in Female
Japanese Macaques Sexual?
An implicit theme in much of the literature is the
notion that same-sex interactions in animals involving
genital contact are devoid of any sexual motivation, de-
spite the fact that they appear to be sexual in terms of
their superficial form. Consequently, homosexual behav-
ior, especially female homosexual behavior, is argued by
some researchers to be a phenomenon that is virtually
unique among humans, with almost no counterparts in
nonhuman animals (e.g., Dixson, 1998; for discussion,
see Bagemihl, 1999; Nadler, 1990; Vasey, 1995, 2000a;
Wallen & Parsons, 1997).
Various types of evidence indicate that female ho-
mosexual behavior in Japanese macaques is sexual. First,
homosexual interactions follow many of the same patterns
as heterosexual copulation (e.g., Fedigan & Gouzoules,
1978; Gouzoules & Goy, 1983; Wolfe, 1984). Although
heterosexual copulation should not be the yardstick by
which all other types of sexual behaviors are measured,
heterosexual copulation is considered, a priori, by many
researchers, to be sexual (e.g., Dixson, 1998; for a discus-
sion, see Bagemihl, 1999; Vasey, 2000a). Consequently,
demonstrating that homosexual behavior parallels hetero-
sexual behavior in many aspects of its expression repre-
sents powerful evidence for some researchers that the for-
mer is, indeed, a sexual behavior. With this caveat in mind,
it is interesting to note that like heterosexual behavior,
(1) female homosexual behavior is never observed
outside of the species’ fall–winter mating season;
(2) females that engage in homosexual behavior ex-
hibit a reddening of the face and perineum that
is indicative of increased sexual receptivity;
(3) female homosexual behavior occurs within the
context of temporary, but exclusive, sexual pair-
bonds called consortships;
(4) female homosexual behavior is temporally linked
to the bidirectional exchange of sexual solicita-
tions between partners (Vasey, 1996; Vasey et al.,
1998). These sexual solicitations appear to be
identical to those exchanged in heterosexual
contexts;
(5) females employ certain mounting postures in a
homosexual context (e.g., double-foot clasp
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58
mounts with thrusting), which appear to be in-
distinguishable from those that males employ in
a heterosexual context (Vasey et al., 1998); and
(6) female mountees do not try to push or shake off
the mounter; instead, they typically brace their
bodies to support the mounters. Often, the moun-
tees will facilitate mounts-in-progress by reach-
ing back and clasping the mounters, while gazing
into the mounters’ eyes. The same pattern occurs
during many heterosexual mounts.
When considering the issue of sexual motivation, it
is also important to note that patterns of affiliation dur-
ing homosexual consortships do not mimic generalized
patterns of social affiliation that occur outside of these as-
sociations. Rather, patterns of affiliation between females
during homosexual consortships represent radical depar-
tures from normal patterns of affiliation that are widely
recognized as social. This suggests that homosexual con-
sortships are not social relationships, but rather, are sexual
ones. For example,
(1) female Japanese macaques exercise incest avoi-
dance with close female kin (r ≤ .25; for more
details, see Chapais et al., 1997). Consequently,
mounting, sexual solicitations, and consortships
are never observed between mothers and daugh-
ters, sisters, or grandmothers and granddaughters
(Chapais et al., 1997; Chapais & Mignault, 1991;
Wolfe, 1984). In contrast, these close categories
of female kin commonly engage in other types
of affiliative behavior with each other, such as
grooming, huddling, and interventions (Chapais,
1992; Koyama, 1991; Kurland, 1977).
(2) female consort partners intervene in conflicts to
support each other at consistently high rates dur-
ing homosexual consortships. Outside the con-
sortship period, these same females rarely sup-
port each other, if at all (for more details, see
Vasey, 1996; see above).
(3) during homosexual consortships, a dramatic in-
crease occurs in the number of bridging interven-
tions performed by dominant consort partners for
their subordinate counterparts (for further detail,
see Vasey, 1996). This pattern is highly unusual
because outside of homosexual consortships,
nonkin female Japanese macaques almost never
perform bridging interventions for each other;
instead, they perform conservative interventions
(Chapais, 1992; Chapais, Girard, & Primi, 1991).
Yet, when opportunities for support are con-
sidered, dominant consort partners are as likely
to perform bridging interventions (55.2% of
Vasey
N = 67 opportunities for support) for their sub-
ordinate partners as conservative interventions
(31.7% of N = 205 opportunities for support;
Wilcoxon test, N = 14, Z 0 = 0.6, P = .55).
(4) normal dominance relationships are temporarily
destabilized during homosexual consortships be-
cause, subordinate consort partners receive sup-
port from their dominant partners against targets
that normally rank above them, or with whom
they share ambiguous dominance relationships
(for further detail, see Vasey, 1996). Over half
of all subordinate consort partners (57.1%; N =
13) rose in rank during the consortship period.
They temporarily increased in dominance dur-
ing 40% of the opportunities to do so, and fell
in dominance during 1.5% of the opportunities
to do so (N = 65). This difference is significant
(Wilcoxon test, N = 13, Z 0 = 2.3, P < .05).
(5) dominant consort partners groom their subordi-
nate counterparts significantly more often than
the reverse (for more details, see Vasey, 1996;
see above). Outside of homosexual consortships,
the opposite pattern holds true: female Japanese
macaques overwhelmingly direct grooming up
the hierarchy, not down (Chapais, Gauthier, &
Prud’homme, 1995; Koyama, 1991).
(6) during homosexual consortships, females are al-
most exclusively in proximity with their nonkin
consort partners (Gouzoules & Goy, 1983; Vasey,
1996, 1998; Wolfe, 1984). Outside the consort-
ship period, however, female Japanese macaques
spend most of their time in proximity with close
kin (Kurland, 1977; Singh, D’Souza, & Singh,
1992).
Finally, clitoral contact and stimulation occur during
many homosexual mounts. During sitting mounts, for ex-
ample, mounters rub their clitorises repeatedly and vigor-
ously against the back of mountees. During some double-
foot clasp and standing mounts, mounters thrust against
the mountees’ clitorises. In addition, mounters and moun-
tees rub their clitorises with their tails during many same-
sex mounts (Vasey, 1996; Vasey et al., 1998; Wolfe, 1984).
Taken together, these various lines of evidence sug-
gest that female homosexual behavior in Japanese
macaques is a sexual behavior, not a social one, or even
a sociosexual one. Moreover, the overwhelmingly bidi-
rectional flow of sexual solicitations, mounts, clasps, ap-
proaches, and interventions within consortships (Chapais
& Mignault, 1991; Vasey, 1996; Vasey et al., 1998)
suggests that these relationships are based on a mutual
sexual attraction between the partners.
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Sexual Preference in Female Japanese Macaques
Are Female Japanese Macaques That Engage
in Homosexual Behavior Unable
to Obtain Male Mates?
It is commonly thought that homosexual behavior
in animals is more frequently manifested when individu-
als have difficulty accessing opposite-sex mates and are,
thus, obliged to accept same-sex sexual partners (e.g.,
Nadler, 1990; Wallen & Parsons, 1997). Several factors
might compromise an individual’s ability to access an
opposite-sex mate. Most obviously, opposite-sex mates
might simply be scarce or, in some cases, entirely ab-
sent. Alternatively, opposite-sex mates might be present
but disinterested in sexual activity.
My colleague and I conducted a behavioral experi-
ment to test whether female Japanese macaques engage
in homosexual behavior because they are unable to access
male mates (for more details, see Vasey & Gauthier, 2000).
The experiment involved two phases (see Methods). We
began with a period of baseline observation on the in-
tact study group that had a female-biased operational sex
ratio (i.e., number of sexually active males to sexually ac-
tive females) typical for this species (5 males:16 females).
Following this, we removed four of the group’s sexually
active males to create an experimental subgroup with an
operational sex ratio that was heavily skewed towards fe-
males (1 male:11 females). Thus, the experimental sub-
group contained all of the adult females and all of the
immature individuals present during the baseline period,
but only one of the sexually active males. We found that
females engaged in significantly more homosexual con-
sortships in the context of this female-skewed operational
sex ratio, as compared to the baseline period (Wilcoxon
test, N = 6, T = 21, P < .05). Thus, homosexual activ-
ity increased among Japanese macaques as the number of
adult males in the population decreased. Increases in fe-
male homosexual activity during the experimental period
of the study could not be attributed to some change in the
reproductive status of the subjects between the baseline
and experimental periods, because none of the subjects
conceived during the period of data collection. Three of
the females conceived prior to the study period, and the
others did not conceive at all during the 1994–95 mating
season.
On the face of it, it would appear that females were
engaging in more homosexual behavior simply because
they lacked male mates. However, this was not an ade-
quate explanation to account for the increased level of
female homosexual activity observed at this time. During
the experimental period of the study, the sole male was
frequently available and motivated to engage in hetero-
sexual activity. For example, on 71% of the days during
Style file version July 26, 1999
59
the experimental period of the study (N = 21 days), the
sole male who was present solicited females for sex, but
they simply ignored him and opted to engage in homosex-
ual activity instead.
We reasoned that females might be rejecting solicita-
tions by the sole available male in favor of same-sex sexual
partners because they did not favor him as a mate under any
circumstances. If this were the case, one might conclude
that for all intents and purposes such females really had
no heterosexual options during the study’s experimental
period. However, the data did not support this conclusion.
The majority of females that rejected a sexual solicitation
from the sole available male in favor of a same-sex sex-
ual partner also formed heterosexual consortships with the
same male at some other point during the study (83.3%;
N = 6). This indicates that females who engaged in ho-
mosexual behavior in the presence of the sole available
male also recognized him, not only as a potential mate,
but also as an acceptable one.
Taken together, this research suggests that, in most
instances, female Japanese macaques do not engage in
homosexual behavior simply because they are unable to
obtain sexually motivated and acceptable male mates.
Are Female Japanese Macaques
Preferentially Heterosexual?
Do individuals actually choose same-sex sexual part-
ners when simultaneously presented with opposite-sex al-
ternatives? Or, are animals with bisexual potential always
preferentially heterosexual? I examined these issues by an-
alyzing patterns of sexual partner choice by anovulatory
female Japanese macaques that were the focus of inter-
sexual competition (for more detail, see Vasey, 1998). In-
tersexual competition for female sexual partners occurred
when a sexually motivated male and female (competitors)
simultaneously sought access to the same female sexual
partner (focus of competition). Competitors were consid-
ered to be sexually motivated, vis-à-vis the focus of com-
petition, if they solicited her for sex while competing in-
tersexually for her or if they solicited her for sex at some
point during the same day.
Intersexual competition for female sexual partners
was initiated when a sexually motivated male intruded on
a homosexual consortship, targeting one of the females
as the focus of competition and the other as his com-
petitor (N = 120 intrusions). Intrusions took the form of
approaches and sexual solicitations directed at the focus
of competition, as well as displacements and aggression
directed at the female competitor. They functioned to in-
crease the males’ access to an anovulatory female sexual
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60
partner (focus of competition), while decreasing or pre-
venting her consort partner from doing the same.
Intrusions on homosexual consortships did not reflect
a generalized motivation among males to disrupt any type
of affiliation between nonkin females. Males (N = 4) in-
truded on homosexual consortships for female sexual part-
ners 4.3 times more often (0.6 intrusions/hr, χ = 0.15)
than they interfered with nonkin female grooming dyads
observed outside the mating season (0.14 grooming inter-
ferences/hr, χ = 0.04). The tendency to disrupt females
engaged in sexual affilation, but not those engaged in so-
cial affiliation, strongly suggests that intrusions are cor-
rectly identified as sexual, not social, competition.
Following 62% of intrusions, intersexual competition
for female sexual partners terminated. In such cases, the
consorting females would (1) ignore the male, who would
eventually leave, (2) move away from the male altogether,
(3) separate and then rejoin to continue their consortships,
or (4) terminate their consortships. Intersexual compe-
tition for female sexual partners could, however, esca-
late with the female competitor posing a counterchallenge
against the intruding male. During counterchallenges, the
female competitor attempted to maintain exclusive access
to her consort partner, the focus of competition, while pre-
venting the male competitor from doing the same. Coun-
terchallenges took the form of approaches and sexual so-
licitations directed at the focus of competition, as well
as displacements and aggression directed at the intrud-
ing male competitor. During some counterchallenges,
female competitors would also interpose themselves be-
tween the male competitor and the focus of competition.
Overall, 38% of intrusions by male competitors pro-
voked counterchallenges by female competitors (N = 45
counterchallenges).
Fig. 1. Choice of competitors by foci of competition following intersexual competition
interactions.
Vasey
Proceptive females did not perform counterchallen-
ges simply because they were in a heightened state of
arousal and thus, more assertive. Rather, proceptive fe-
males (N = 9) retaliated, in kind, against males who ag-
gressed or displaced them significantly more often when
they were engaged in homosexual consortships (χ =
17.4%), as compared to when they were not (χ = 0.01%;
one-tailed Wilcoxon test, N = 5, T = 15, P = .03). Fur-
thermore, females were never seen to retaliate against
males who disrupted their grooming interactions with
nonkin females outside the mating season. The females’
tendency to challenge males who disrupted their sexual,
but not their social, affiliation strongly suggests that coun-
terchallenges are correctly identified as sexual competi-
tion and are not part of a more ubiquitous pattern of social
competition.
Following episodes of intersexual competition, fe-
males that were the foci of competition could choose be-
tween two sexually motivated competitors, one male and
the other female, who were simultaneously available. Con-
sequently, these interactions represented an excellent “nat-
ural experiment” for testing whether females preferred
certain female sexual partners relative to certain male
alternatives when simultaneously given a choice. In the
overwhelming majority of cases (N = 120), the foci of
competition chose to continue mounting with their female
sexual partners significantly more often than they chose
to begin mounting with the male competitor (Wilcoxon
test, N = 14, T = 105, P = .002) or to cease mounting
altogether (Wilcoxon test, N = 13, T = 91, P = .002;
Fig. 1).
Taken together, this research suggests that in those
populations where female Japanese macaques engage in
both homosexual and heterosexual behavior, females are
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Archives of Sexual Behavior pp350-aseb-365185 December 28, 2001 8:29
Sexual Preference in Female Japanese Macaques
not preferentially heterosexual. Females will choose
same-sex sexual partners, even when they are simulta-
neously presented with sexually motivated, opposite-sex
alternatives.
Sexual Preference in Female Japanese Macaques
In light of the evidence presented above, I now return
to the question with which I opened this paper: Do animals
ever exhibit a preference for same-sex sexual partners? If
female Japanese macaques are the animals under consid-
eration, the answer to the question appears to be a qualified
“yes.” Although female Japanese macaques demonstrate
a preference for particular same-sex sexual partners, it
would be incorrect to refer to female Japanese macaques
as “homosexual” or to suggest that they are “preferen-
tially homosexual.” This is because, in those populations
where female Japanese macaques engage in homosex-
ual behavior, they almost invariably do so in combina-
tion with heterosexual behavior over the course of their
life spans. Exclusive preference for same-sex sexual part-
ners over the entire life span has been documented in
Japanese macaques, but such cases are exceptionally rare
(McDonald-Pavelka, 1993, p. 108). It is possible that in
certain populations some females manifest exclusive ho-
mosexual behavior throughout an entire mating season.
As a rule, however, females who engage in homosexual
behavior also engage in heterosexual behavior during the
same mating season, and, indeed, they sometimes even
alternate between these activities on the same day.
How then might female sexual preference in Japanese
macaques best be characterized? In some populations,
individual females, when faced with a pool of potential
sexual partners, prefer certain same-sex sexual partners
relative to the available opposite-sex mates. It is impor-
tant to stress, however, that the reverse is equally true:
the same females will prefer certain male mates rela-
tive to the available female sexual partners. As such, fe-
males do not invariably prefer males over females, nor vice
versa.
The likelihood that a female sexual partner will be
chosen over a male increases whenever preferred female
sexual partners are abundant in the population. Conse-
quently, it is not a lack of males per se that promotes the
expression of female homosexual activity in the context
of female-skewed operational sex ratios; rather, it is the
relative paucity of preferred male mates coupled with a
relative abundance of preferred female sexual partners.
The precise nature of individual sexual preferences
will vary greatly from one female to the next and will de-
pend largely on idiosyncratic predilections for particular
Style file version July 26, 1999
61
sexual partners with particular characteristics, be they
male or female. Taken together, this evidence suggests
that, in some populations, female Japanese macaques are
best characterized as bisexual in orientation, not preferen-
tially homosexual or preferentially heterosexual.
ACKNOWLEDGMENTS
I thank Michael Bailey, Ray Blanchard, Bernard
Chapais, Lee Ellis, Carole Gauthier, Stefani Kovacovsky,
Richard Pawsey, Sergio Pellis, Jim Pfaus, Anne Russon,
Ken Zucker, and two anonymous reviewers. Various stages
of this research were funded by the Universite de Montreal,
the Sigma Xi Society, the Fonds pour la formation des
chercheurs et l’aide de recherche (FCAR) Quebec, the
University of Lethbridge, and the Natural Science and En-
gineering Research Council (NSERC) of Canada.
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