Theory in Biosciences (2020) 139:191–207

https://doi.org/10.1007/s12064-020-00312-8

ORIGINAL ARTICLE

Hominid sexual nature

Christopher Mogielnicki1 · Katherine Pearl2

Received: 25 September 2019 / Accepted: 3 March 2020 / Published online: 13 March 2020
© The Author(s) 2020

Abstract
The aim of the paper is to identify psychosomatic evolutionary adaptations of hominids, which direct them at maximizing
their reproductive success, and on the basis of which their various social structures are built. Selected features of the hominid
last common ancestor were extracted; by reducing the influence of the social structure, they were defined as the hominid
“sexual nature”; these considerations were supported by the analysis of sexual jealousy as a function of socio-environmental
conditions. The “sexuality core” of a hominid female was defined as “selective polyandry”—the female selects the best males
among those available; and of a hominid male as “tolerant promiscuity”—the male strives for multi-male and multi-female
copulations with sexually attractive females. The extracted “sexuality cores” condemn hominids to a patriarchal social struc-
ture and thus to sexual coercion and jealousy. The source of male sexual jealousy is limited access to females. Hominid female
jealousy of the male results mainly from the need for protection and support. Hominids’ social structures are determined by
females’ sexual selectivity or opportunism and by their continuous or periodic proceptivity and estrus signaling. Evolution-
ary functions developed by women: out-estrus sexuality, copulation calls, multiple orgasms, allow them to obtain the best
possible spermatozoid. The institution of marriage blocks the influence of sexual selection in the species Homo sapiens.

Keywords Hominid sexuality · Hominid social structures · Sexual jealousy · Mate choice

Introduction relatives of humans, common chimpanzees (Pan troglo-

dytes, hereafter also called chimpanzees), correlates with
Due to the cultural multiplicity of reproduction systems this theory. Females, making 100% of parental investments,
in the species Homo sapiens, it is difficult to clearly deter- receive food from males and suffer sexual aggression from
mine the type of its breeding strategy. This demonstrates them (Goodall 1986). On the other hand, bonobo chimpan-
the flexibility and adaptability of the reproduction system zees (Pan paniscus, hereafter also called bonobos) have free
that depends on environmental and social conditions. Human sexual access to one another and the females, who also bear
reproductive systems can generally be divided into patri- full parental investments, are not particular about choos-
archal and matricentric (matrifocal). A vast majority of ing a partner but have priority in access to food (Stanford
human societies function under patriarchal conditions, which 1998). The diversity of reproductive strategies, not only in
include polygyny (where one man can have several wives), the species Homo sapiens, but also among the entire family
a serial monogamy (where a man can have only one wife), of hominids, tends to search for a “common sexual denomi-
and polyandry (where one woman has several husbands). In nator” for all hominids—the hominid “sexual nature.”
matricentric structures, in turn, marriages do not occur at all. Hominid (Hominidae) is a family of great apes that
According to the Trivers (1972) theory, a gender that includes species whose representatives are the largest among
invests more in the offspring will be more careful when the primates. They show intelligence, tendencies to adopt
choosing a partner, while the gender that invests less will a bipedal attitude, and the ability to produce and use tools.
compete for him/her. The behavior of genetically close This paper describes a monophyletic group of the hominid
species: the gorilla (Gorilla, hereafter also called gorilla),
* Christopher Mogielnicki
common chimpanzee (Pan troglodytes), bonobo (Pan panis-
[email protected] cus) and modern human Homo sapiens. They are all social
species. Apart from chimpanzee, bonobo and gorilla, Homo
1
Bialystok University of Technology, Bialystok, Poland sapiens’ close relative also includes the orangutan (Pongo).
2
Bialystok, Poland

13
Vol.:(0123456789)
192 Theory in Biosciences (2020) 139:191–207

These species, together with extinct human forms, form the constant in the course of evolution, common for four homi-
Hominidae family (Wilson and Reeder 2005). In this study, nid species and independent of the environmental influences;
the orangutan is not analyzed because of belonging to the they are male and female sexual functions that maximize the
subfamily Ponginae, a more distant kinship with humans and hominids’ reproductive success. To determine these “cores,”
its solitary way of life (Locke et al. 2011). the expected levels of sexual jealousy of the hominid male
Gorillas function in the terrestrial environment, in homo- and female, depending on the access to sexual partners and
genic harem groups (a male and his females), in which to environmental resources, have also been designated.
daughters and sons leave the family. Gorilla females do not
exhibit external signs of ovulation, and therefore, an estrus
is signaled by behavioral signals such as proceptive behavior Female mating predispositions
(Stewart 1977; Watts 1991). Chimpanzees live in large ter-
restrial and arboreal heterogeneous herds with a dominant Sexual behavior during infertile periods (including infer-
male, and their females signal the estrus with anal–genital tility before entering into the reproductive period) is not
swelling (Goodall 1986). Matriarchal bonobos are also ter- uncommon among apes. It occurs, e.g., in order to obtain
restrial and arboreal hominids which live in heterogeneous food from a male or to reconcile among bonobos (De Waal
herds. They do not have the dominant male; males are dom- 1987). However, Furuichi (1987) found that although bon-
inated by females. Bonobo females simulate a permanent obo females copulate when they are not maximally swollen,
state of ovulation (Stanford 1998). Promiscuity and constant more than 95% of matings were observed during maximal or
receptivity of the bonobo females caused the disappearance near-maximal swelling. It is the same for chimpanzees—97%
of the need to compete for the access to them and thus the (Goodall 1986). As for sexual receptivity, bonobo females
disappearance of the dominant male institution. The lack of are more flexible than other apes. It became a fundamental
the need to compete for the females, as a result of the abun- premise of bonobo sexuality and of the bonobo’s link to
dance of sex, is anatomically manifested in bonobos by the human behavior because, among the primates, only bonobo
reduction of canine dimorphism (Wrangham and Peterson and human females are sexually active outside of the perio-
1996; De Waal and Lanting 1997). vulatory period (Stanford 1998).
Homo sapiens is definitely dominated by militant, patri- Swollen chimpanzee females copulate with multiple
archal social structures with hierarchized males (Otter- males during the early stages of their swelling cycle. As
bein 1970; Divale 1972; Keeley 1996). It was found that many as 50 copulation bouts with eight males during 1 day
residence patterns are related to the frequency of violence were observed and the swollen females copulated with up
between groups: communities based on related women are to eight adult males within several minutes (Goodall 1986).
characterized by a lower rate of conflicts than patrilocal Bonobo females actively solicit sex from a range of males
ones (Adams 1983). The same qualitative observation was and may copulate multiple times per hour while swollen
made in a comparative study of humans and primates (Man- (Kano 1992). Gorilla females usually tend to mate only for
son et al. 1991). Homo sapiens is terrestrial and can form a few days during a 28-day cycle (probably at the time of
social structures based on both related males and females. Its ovulation), but they also show uncommon periods of pro-
females, just like bonobos, also signal the permanent state ceptivity at other times (Harcourt et al. 1980).
of the estrus. The females of all Hominidae species copulate Female copulatory vocalizations, i.e., sounds issued by a
also out of an estrus season (Savage-Rumbaugh and Wilker- female before, during, or just after a sexual intercourse, com-
son 1978; Stoinski et al. 2009a, b). mon to many primates (including the humans) are strongly
On the basis of selected features of the extant hominid associated with a promiscuous reproductive system. Females
species mentioned above, analogical features of their last from promiscuous species emit more complex copulation
common ancestor (LCA)—Hominidae clad, have been deter- calls than the females from polygynous or monogamous
mined. To determine LCA features, based on phylogenetic species. The degree of promiscuity of the females allows
relationships of living hominids presented in Scally et al. to predict how they tend to vocalize during the copulation.
(2012) and Prüfer et al. (2012), the cladistic systematic It is believed that copulatory vocalizations have two basic
analysis method was used (Hennig 1965). By reducing the functions: to encourage other males to mate and to increase
influence of the LCA social structure on the features of its the protection of the act by a copulating male (Pradhan et al.
representatives, the “sexuality cores” of the hominid male 2006).
and female have been designated. In this paper the “sexual- The analysis of the copulation calls of yellow baboon
ity core” will be, independently of the social structure, bio- females (Papio cynocephalus) showed that these sounds
logical and psychological dispositions, on the basis of which inform both about the condition of the female (the closer the
hominids’ social structures are created as a function of envi- female is to the ovulation, the more complicated sounds) and
ronmental conditions. In other words, “sexuality cores” are the social status of the male copulating with her (the higher

13
Theory in Biosciences (2020) 139:191–207 193

the male in the hierarchy, the longer the scream consisting of Johnson 1966). The women do not enter, as it is the case
a larger number of acoustic units) (Semple 2001). The pri- of men, into the refraction phase as a result of achieving
mate females during the copulation let out shouts that cannot the orgasm. For some time they remain at a high level of
be heard in any other situation. An orgasm or symptoms of excitement, which slowly decreases, and in some women
sexual pleasure are easily recognized (Small 1989). Accord- even very slowly. During this period, the woman subjected
ing to Small, macaque wander (Macaca silenus) females to erotic stimuli can easily achieve a greater number of the
attract by such calls out-group males to differentiate the gen- orgasms. Studies of mechanical stimulation of the pubic
otype in the population. In Barbary macaques (Macaca syl- mound using a vibrator showed that orgasmic sensations
vanus), the likelihood of male ejaculation is directly related were intense and achieved in a minimum of time one after
to the intensity of female vocalizations (Todt and Pohl 1984) another in the waves of the multiple orgasms, closing a sin-
and this appears to be independent of the male’s copulatory gle sexual episode (Maers and Johnson 1966).
effort (Pfefferle et al. 2008). The copulation calls are also Basson (2001) indicates that some women, during an ovu-
present in the chimpanzee (Hauser 1990; Townsend et al. latory period or after prolonged sexual abstinence, have a
2008), bonobo (Clay et al. 2011), gorillas (Nadler 1976) kind of “sexual hunger”—the intensification of sexual needs,
and Homo sapiens. Adaptation of vocalization evolved and which is called a “spontaneous desire,” and which is related
persisted in both arboreal (the chimpanzees) and terrestrial to the action of hormones (including androgens), dopamine,
(baboons) species, despite the threats that it carried. oxytocin, noradrenergic receptors in the central nervous
Matsumoto-Oda (1999) reports that during an earlier system or structures in the limbic system responsible for
estrus phase, the chimpanzee females copulate quite freely emotions. This state increases the susceptibility to sexual
with many males. However, in the later stage of the estrus, stimuli, and the rate and intensity of stimulation. Basson
when the probability of fertilization is the highest, they cop- believes that in addition to these cases, the women’s desire
ulate rather with high-ranking adult males. A hormonal anal- has a psychological background and results from the need
ysis carried out by Townsend et al. (2008) showed that the for tenderness, strengthening ties or recognition of their own
chimpanzee females vocalized much more often when they attractiveness. Only such a motivated woman opens herself
were with high-ranking males and they suppressed the calls to sexual stimulation and as a result, the woman gets the
when there were high-ranking males nearby. According to ability to generate excitement. Basson calls this desire the
the authors, the copulatory vocalizations can thus be one of “triggered desire,” or responding to the stimulus.
the potential strategies used by the female to advertise recep- An ability to achieve the orgasm by the women depends
tivity to the high-ranking males, to confuse fatherhood and on the phase of a monthly cycle: estrogens stimulate sexual
to provide future support from these socially important indi- tendencies (affect the blood supply to the vagina, labia and
viduals. In the case of women, it was also recognized that clitoris, and the hydration of the vagina), while the proges-
at least one element of these responses was under conscious terone suppresses them. Androgens, in turn, increase sexual
control, providing them with an opportunity to manipulate activity. The testosterone regulates the level of excitement
the male behavior, e.g., to accelerate partner’s orgasm, when and desire, and its concentration increases markedly a few
a woman was already weary (Brewer and Hendrie 2011). days before ovulation. The sexual fitness and attractiveness
Although all mammal females have a clitoris, there was of the women reach the highest level in the ovulatory period
a conviction confirmed by zoologists’ research that most (Rowland 2006; Bullivant et al. 2004). In this period, the
animals, with the exception of the bonobos, chimpanzees women have the orgasm most easily (Wallen and Lloyd
(Allen and Lemmon 1981; Wrangham 1993) and macaques 2011; Azadzoi and Siroky 2010; Dunn, et al. 2005). During
(Macaca arctoides) (Chevalier-Skolnikoff 1974) do not feel the ovulatory period, the women have sexual relations more
an orgasm. However, Troisi and Carosi (1998) note that than three times more often than during the postovulatory
it is possible to cause the orgasm in every primate female phase (Baker and Bellis 2014).
through artificial stimulation. Having examined the sexual- Evolutionary adaptations in the form of clear ovulation
ity of the primates, Dixson (1998) states that the primate signaling, the presence of multiple orgasms and the female
females, especially those from promiscuous reproductive copulation calls predispose species to multi-male mating.
systems (such as the macaques or chimpanzees), have the It is also known that in the case of the women, adapting to
orgasms while copulating. In contrast, the females of mostly the conditions of the multi-male mating is also indicated
monogamous gibbons (Hylobatidae) or harem gorillas do by: the ability of the female reproductive system to assess
not show any visible signs of the orgasm (Dixson 1998). immune compatibility of sperm on the basis of its chemical
In the case of the women, an orgasmic phase can occur structure (Birkhead and Pizzari 2002; Barratt et al. 2009),
repeatedly within a single sexual intercourse. An average preference of the reproductive system for the sperm deliv-
woman with optimal stimulation is able to achieve three to ered during the orgasm (Baker and Bellis 1993b). In Homo
five orgasms caused by manual stimulation (Masters and sapiens, additional factors are male evolutionary adaptations

13
194 Theory in Biosciences (2020) 139:191–207

to sperm competition, e.g., testis size (Møller 1988), ejacu- 1980). Jaeggi et al. (2013) show, however, that the bono-
late adjustment (Baker and Bellis 1993a; Shackelford et al. bos use seduction and sociosexual contacts to reduce ten-
2002), sexual arousal (men’s sexual fantasies often involve sion caused by monopolized food, which is not shown by
multiple partners) (Pound 2002), ejaculate gametes’ role the chimpanzees. Food transfer was not observed in wild
(Baker and Bellis 1988; 1989) and penis shape (Gallup et al. mountain gorillas (G. beringei) (Schaller 1963) and low-
2003; Mautz, et al. 2013). land gorillas (G. G. Gorilla) (Dixson 1981). However, it
was reported that a lowland gorilla male kept in captivity
moved a piece of an apple mouth-to-mouth through the
Male–female food transfer bars of the cage to his female neighbor (Schaller 1963).
The male–female food transfer evolves in the species
Evolutionary sexual conditions of Homo sapiens indicate along the possibility of choosing a partner by the female.
a high degree of mismatch between the women and men in Sharing food within the gender is associated with form-
terms of long-term satisfaction of mutual needs (Table 1). ing a coalition. These phenomena are statistically related
A basic “common interest” that would combine the Homo to each other (Jaeggi and Van Schaik 2011). Fruit located
sapiens male and female, and which would evolutionarily in high tree branches is important in the chimpanzees’
enforce monogamous behavior, would be collaborative diet. This type of fruit, difficult to obtain and therefore
long-term offspring care. However, great ape males are attractive, in the hands of a male can arouse the females’
not monogamous and do not bear paternal investments; the interest in it. Gorillas lead a terrestrial life and feed on
quality of offspring is measured by the mother’s ability to leaves and rhizomes, so that both genders can easily har-
provide resources (Kramer 2011). Modern men also avoid, vest from the ground. Gorilla males neither establish a
if possible, paternal investments. Therefore, as a rule, the coalition aimed at group competition for the females, nor
burden of carrying pregnancy and feeding the young rests they allow female choices, so food sharing does not occur
on the hominid females. Thus, the female who will bear in this species. However, when a gorilla male does not
the entirety of the parental investments will expect support claim rights to a female, it gives her food.
from the males with whom she has sexual relations. The Among primitive tribes, a phenomenon of sharing meat
phenomenon of male–female food sharing occurs in all spe- between related and unrelated members of the community
cies of the analyzed apes, despite the diversity of their social is popular (Kaplan et al. 1985; Le Jeune 1897). This may
structures. result directly from the inability to store it and the unpre-
Several hypotheses based on a reciprocity rule suggest dictability and relative rarity of success on the hunt of
social benefits of sharing food in the chimpanzees: “food for individual hunters. Researchers state that in societies that
sex,” “food for grooming” and “food for support” (Stanford collectively divide meat, better hunters have more chil-
et al. 1994; De Waal 1997; Mitani and Watts 2001). These dren. They indicate that there are two reasons for this rela-
hypotheses are not mutually exclusive (Crick et al. 2013). A tionship. First, more effective hunters are more desired by
comparative study showed that males gave priority to access women, which makes them have relatively easier access to
to food they possessed to females in the estrus, especially extramarital sex. Secondly, tribe members are more will-
to those they had recently copulated with (Yerkes 1941). A ing to look after the offspring of better hunters. Hunting
long-term study conducted by Gomes and Boesch (2009) skills, in connection with the habit of meat sharing, do not
implied that among the chimpanzees, “food for sex” is not contribute to the multiplication of men’s goods, but decide
a short-term food exchange aimed at copulation, but a long- about their social position, which affects both access to
term strategy for building relationships that enable future women and the quality of care for their offspring (Kaplan
mating. et al. 1984; Kaplan and Hill 1985; Hill and Kaplan 1988).
Many similarities in the food sharing are seen between In other words, it can be claimed that “the hunter hunts for
the chimpanzee and the bonobos that live wild (Kano lovers, not for wives.”

Table 1  Sexual conditioning of Homo sapiens in the context of monogamy

Woman Man

“Triggered” out-estrus sexuality, “sexual hunger” during the ovulatory period Permanent sexual readiness, the Coolidge effect (Kinsey et al.
(Basson 2001) 1948; Symons 1979; James 1981)
Long vaginal penetration necessary to achieve the orgasm (Weiss and Brody Short copulation time (Kinsey et al. 1948; Waldinger et al. 2006)
2009)
Tendency to multiple orgasms (Masters and Johnson 1966) Refraction after the orgasm

13
Theory in Biosciences (2020) 139:191–207 195

Research on the advantages appreciated by American Parents do not have any economic ties. Mosuo’s “walking
women from permanent and fleeting partners has shown that marriage” is based solely on love and mutual attraction.
four advantages are more highly valued among candidates Acceptance of the man, as a son-in-law, usually takes place
for a fleeting contact. These are: spending a large amount of when the girl’s mother accepts customary gifts: tea, wine,
money on a woman from the beginning, giving her valuable clothes for the girl. The language of the Mosuo does not
gifts from the beginning, leading a sumptuous lifestyle and know such words as: war, jealousy, rape and murder (Mat-
making generously his resources available to her. These four tison 2011).
virtues are expected moderately from husbands and highly A phenomenon of male–female food transfer present
from partners of a fleeting relation (Buss and Schmitt 1993). in the patriarchal chimpanzee allows dominated males, by
In matricentric human societies, like island Trobrian- establishing long-term relationships with the female, for a
ders, men bring their lovers various goods: food, tobacco reproductive success: the female will agree on the estrus
and ornaments, and women agree on sexual relations as long copulation. In exchange, bearing the full parental invest-
as a stream of gifts flows. A woman may say, “If you do not ment, she gets increased access to food. The bonobo females,
have a gift for me, then I do not agree.” The reputation of a constantly attractive and receptive, allow all the males to
Trobriand man may suffer among the women if he refuses approach them through the entire menstrual cycle, which
to give a gift to his lover, and as a result, his ability to obtain allowed matriarchy and thus the priority of the females in
another one will decrease too (Malinowski 1929). access to food. The male–female food transfer phenomenon
In Nayar’s polyandry relationships from southwest India, is also significant for Homo sapiens. In societies where there
a girl married one man, but later, in fact, she had sexual rela- are no conventional, long-term patriarchal marriages, where
tions with other men who were not related to one another. a woman chooses a sexual partner and is independent from
She lived alone and had relations with them in succession, him in an existential way, the transfer of resources to the
with their consent. In connection with the matrilineal sys- sexual partner (the woman) is expected. In primitive com-
tem of these people, children inherited from their mother’s munities, the women find it easier to make sexual contact
brother. The whole burden of child care rested on its mat- with effective hunters from whom they get meat. Meanwhile,
rilineal relatives. However, when the woman became preg- contemporary women from a monogamous American cul-
nant, she indicated a particular man as the father of the child ture expect from their fleeting partners intense transfer of
and he was obliged to support her in upbringing (Malinowski money. In matricentric human societies, women are not
1929). Under such conditions, biological paternity was not dependent on men’s resources, but they also expect support
very important and social paternity belonged to the man who from them.
paid the midwife for delivering a child. The “husbands” did
not live with their “wives,” but with their maternal relatives
whom they visited for the night. Each of the parties could Male and female sexual jealousy
end such a relationship at any time, without any special for-
malities. The visiting partner brought a small cash gift to The analysis of jealousy carried out by Symons (1979) and
the woman, while a more regular “husband” gave her (at Daly et al. (1982) provides the main theoretical basis for
the beginning of the relationship) some material for a skirt the evolutionary analysis of the jealousy. Sexual jealousy is
and then gifts on the occasion of three annual festivals (e.g., conceptualized as a functional emotion, whose basic purpose
loincloths, Beretta leaves, Areca nuts, hair and bath oils and is to preserve the bonds of the precious relationship (Buss
some types of vegetables). Not giving such a gift was a sign 2000, 2013). Dozens of empirical studies on the psychology
that the man wanted to end the relationship (Gough 1959). of the jealousy indicate that men and women do not differ in
Matricentric Mosuo people live near Lugu Hu Lake and the frequency or level of the jealousy they experience. Male
in Sichuan in Southwestern China. Every household consists jealousy, compared to the female one, seems to be more
of 8 to 20–30 members and these are multigenerational fami- sensitive to signals of sexual infidelity. Women’s jealousy,
lies, making their living by cultivating land. The relationship on the other hand, seems to be relatively more sensitive to
between a woman and a man is that they permanently live signals of emotional unfaithfulness (Buss 2013).
in their family homes, and the man accepted by the woman The hierarchy of the females that conditions access to
stays with her for the night and then returns home. The food is present in the chimpanzee species. The females are
institution of the father is not known in the Mosuo people. aggressive toward new immigrant women and even kill new-
Children born of such a relationship belong to the woman born babies of community members (Pusey and Schroepfer-
and they inherit from her, take her surname and are brought Walker 2013). Woman aggression is directed mainly against
up by her family. The role of the father is performed by other women and generally causes minor injuries. The most
their uncles. All children in a given family are brought up common victims are relatives and rivals in sex and marriage.
together. Parental separation does not affect their situation. The aggression is often a means of competing for people or

13
196 Theory in Biosciences (2020) 139:191–207

food products (Burbank 1987). The bonobo females differ mammals. Chimpanzee dominant males show jealousy
from the chimpanzees and from women; they work together mainly of females in the estrus. Sometimes they direct
to dominate adult males in competition for food (Furuichi aggression to lower-ranking males, but sometimes also to
1997; Vervaecke et al. 2000). a female for her sexual relations with others (De Waal
Table 2 presents factors generating the jealousy of a hom- 1982). In patriarchal polygyno-monogamous human soci-
inid female. Resources necessary for survival are divided eties, a man can have access and a monopoly usually on
into two groups: “unstable”—on a current basis derived a relatively small number of women in a given period of
from the natural environment (the open set), and “accumu- time (often only one), through which he can pursue his
lated”—obtained by agriculture (the closed sets). The last reproductive success. A similar situation occurs in the
line presents the expected level of the female jealousy of a dominant males of the chimpanzees or gorillas who have
male, determined on the basis of the level of factors gener- access to and monopoly on a relatively small number of
ating this jealousy. The term “female jealousy of the male” the females currently signaling ovulation. In contrast, bon-
and “male jealousy of the female” is understood here as obo males have free access to all the females, as there are
imposing sexual restrictions on the opposite gender. Table 2 no dominant males there and the females permanently sig-
indicates that the level of the female jealousy of the male nal the state of ovulation. The woman is also permanently
is closely related to the socio-environmental conditions in attractive, so in the eyes of the men she is constantly in
which she functions. a potential ovulation state. Therefore, the jealousy of the
Male sexual jealousy is a common function of the man who is holding a monopoly on the woman will also
dominant male of chimpanzees, gorillas and other herding be permanent.

Table 2  Distribution of the hominid female jealousy of the male 1980; Jaeggi et al. 2013; Schaller 1963; Dixson 1981; Jaeggi and Van
(Goodall 1986; Watts 1991; Stanford 1998; Kramer 2011; Malinow- Schaik 2011; Kaplan et al. 1984; Buss and Schmitt 1993; Smuts and
ski 1929; Mattison 2011; Gough 1959; Hill and Kaplan 1988; Kano Smuts 1993)

Female♀ Apes Homo sapiens

Social conditions Gorilla (Gorilla) Chimpanzee (P. Bonobo (P. panis- Polygyny and Polygyny and Matricentric poly-
One-male harem troglodytes) cus) monogamy (unsta- monogamy andry
Multi-male harem Matriarchy ble resources in (accumulated
open set resources—in
closed sets)

Factors
Access to environ- Yes Yes—hierarchic Yes—females Partial—females Partial or no Yes
mental resources: have priority do not get meat access
Access to preferred Limited—except Partial Unlimited Limited Limited Unlimited
sexual partners: for access to the
dominant male
Economic depend- Absent Absent Absent Absent –meat is Present Absent
ence on the male: consumed col-
lectively
Expected level of None or low Low Low Low High Low
gifts from males:
Need for protection Permanent—dur- Permanent— Absent Permanent Permanent Female is protected
from the male ing the estrus during the estrus by her brothers
against sexual and against and against
aggression: infanticide infanticide
Need for support Absent Absent Absent Present Present Low level—the off-
from the male spring of women
in the care for are taken care
offspring: of mainly by her
relatives
Expected level Low Low Absent Medium High Low
of the female
jealousy of the
male:

Factors significantly generating the female jealousy are underlined

13
Theory in Biosciences (2020) 139:191–207 197

The hypothesis of the certainty of fatherhood, as the The modern hominids function in various social struc-
source of the male jealousy presented by, for example, Alex- tures: patriarchal and matriarchal. This is why it is difficult
ander and Noonan (1979), is difficult to accept. In cases of to define their “sexual nature.” The following methodology
sexual behaviors such as inter-marital infidelity, rape, war was used to determine the “sexuality cores” of the homi-
rape, group sex, in matricentric human societies, the bono- nid male and female: it was based on the list of selected
bos and in the case of the dominated chimpanzee and gorilla features of four hominid species, the presence of these
males, the male jealousy is absent. It is also difficult to features was determined in LCA. Then, by eliminating the
assume that in the course of evolution, fatherly investments influence of the patriarchal social structure on these fea-
would be the cause of the need to increase this certainty tures, their presence was defined as the “sexual cores” of
through the jealousy. In primitive communities, especially the hominid male and female.
men with a high social position, devote little time to dealing
with their own children (having the largest number of them)
(Hewlett 1991). Hominid female’s “core of sexuality”
Table 3 contains factors generating the male jealousy in
modern hominids as a function of their socio-environmental Sexual behavior of the hominids, seen in the context of
conditions. The last line shows the expected level of the male the degree of access restrictions for the females, allows
jealousy, determined on the basis of the level of factors that to define the “cores of sexuality” specific to them. The
generate this jealousy. Based on the cases of both chim- cores are male and female sexual functions that are inde-
panzee species and the human matricentric societies, it is pendent of the environmental influences and common for
assumed that just limitation in access to the females causes all social hominids. These functions incline the hominids
a high level of the male jealousy of the female. to maximize their reproductive success. The “sexuality
core” of the hominid female, as independent of the social
structure, was extracted on the basis of the comparison of
Hominid sexual nature the selected features (Table 5).
On the basis of Tables 4 and 5, it can be assumed that
Based on the features of the modern hominids discussed the hominid female tends to achieve full sexual freedom,
above, analogous social features of their common ancestor “selective polyandry” (Fig. 1), being in safe living con-
were extracted (Table 4). These features were then treated ditions. An essential condition ensuring the sexual free-
as the basis on which the construct of the “sexual nature of dom for her is a free choice of sexual partners and time
the hominids” was built. of sexual intercourse. The quality of the father is crucial
for her because of the parental investment that she incurs
due to pregnancy and lactation and due to the reproductive
success of her genetic line in the future.

Table 3  Distribution of the hominid male sexual jealousy (Goodall 1986; Watts 1991; Stanford 1998; Kramer 2011; Malinowski 1929; Mattison
2011; Gough 1959; Hill and Kaplan 1988)
Male♂ Apes Homo sapiens
Social conditions Gorilla (Gorilla) Chimpan- Bonobo (Pan paniscus) Polygyny and Polygyny and Matri-
One-male harem zee (Pan Matriarchy monogamy (unsta- monogamy (accumu- centric
troglodytes) ble resources in lated resources—in polyandry
Multi-male open set) closed sets)
harem

Factors
Access to females Limited Limited Unlimited Limited Limited Facilitated
Level of investments Absent or low Low Low Low High Low
directed to the female
Level of investments Absent Absent Absent Low High Low
directed to offspring
Expected level of the male High (concerns High (con- Absent or low High (concerns the Very high (concerns Low
jealousy of the female the dominant cerns the male claiming the male claiming
male) dominant rights to the rights to the female)
male) female)

Factors significantly generating the male jealousy are underlined

13
198 Theory in Biosciences (2020) 139:191–207

Table 4  Social features of the hominids conditioning their sexuality (Jaeggi and Van Schaik 2011; Savage-Rumbaugh and Wilkerson 1978;
Smuts and Smuts 1993; Watts 1989; Lancaster and Lancaster 1983; Jaeggi et al. 2013; Gomes and Boesch 2009; Basson 2001)
Feature Female/male
Homo sapiens Chimpanzee (P. trog- Bonobo (P. paniscus Gorilla (Gorilla) Last common ancestor
lodytes)

Social system Mainly patriarchy, Patriarchy Matriarchy Patriarchy Patriarchal social

matricentrism structure
Community Heterogeneous Heterogeneous Heterogeneous Homogenous Heterogeneous com-
munity
Copulation periods Perovulatory; perma- Perovulatory Permanent Perovulatory Perovulatory sexual
(female sexual nent attractiveness behavior
attractiveness)
Parental investments Present/present and Present/absent Present/absent Present/absent Females bear all
­absenta parental investments
Male–female food Present Present—for building Present—male shares Absent in the wild; Males share food with
sharing long-term relation- food with a seduc- present—if a female females
ships ing him female doesn’t belong to a
male

Information given by the authors is written in an extended text

a
Data from the Central Statistical Office show that in Poland men make up approx. 96% of the total child support they are required to pay, of
whom approx. 80% do not pay the granted support

Hominid male’s “core of sexuality” fruit, while low parts of the bush mainly offer leaves and
rhizomes (low-value food), for which the gorillas had to
Features of the species on the basis of which the “sexuality develop coprophagia (Akers and Schildkraut 1985), occu-
core” of the hominid male was determined are presented in pying these niches. Gorillas’ dominant males, increasing the
Table 6. body size, got rid of other males from the herd, because they
Table 6 reveals that the “tolerant promiscuity” (Fig. 2) were able to defend it alone against threats from the environ-
will be sexually comfortable for the male: free access to ment. Therefore, both adolescent daughters and sons were
sexually attractive females and relationships with them in removed from the herd. After getting rid of internal rivals
the conditions of sperm competition. The bonobo females in accessing the females, they established a harem structure,
adapted to that need. They gave the males sexual fulfillment; reducing the need for sperm and thus the size of testicles:
however, the cost to the males was resignation from domi- the gorilla male has the smallest testicles among the homi-
nating over the females. The hominid male does not need nids (Harcourt et al. 1981). Taking into account low-value
any choice of the female: the only criterion that the female nutritional conditions, it is possible that for these primates,
should meet him again is her ability to stimulate him sexu- investing in the growth of the male’s mass and functioning in
ally—she must have clear enough signs of fertility. small, harem groups, turned out to be more energy-effective
than in-group competition for the females with other males
in the heterogeneous herd.
Discussion Sexual aggression and infanticide among mountain
gorillas (Gorilla beringei beringei) are widespread (Har-
The gorilla phenomenon court and Stewart 1987), which links them with the chim-
panzees. Infants living in the male leader group are rarely
Group violence against other communities (war) besides killed, while almost half of all the infants who leave it are
Homo sapiens is also present among patriarchal cooperat- killed shortly thereafter by other males. These data support
ing chimpanzees (Pan troglodytes) (Goodall 1986); a war the hypothesis that the murder of the offspring directs the
provides dominated males with genetic advantages. Gorillas selection responsible for the group life of the gorillas (Watts
(Gorilla), however, do not cooperate, but they can also suc- 1989; Wrangham 1979, 1982, 1987).
cessfully support the patriarchal social structure (De Waal The sexual aggression and infanticide could contribute
1982). to the disappearance of estrous swelling in gorilla females.
The gorilla is the first clad which separated from the taxon Visual estrus signaling generates inter-male competition,
leading to Homo sapiens and developed apomorphic features hence uncertainty of fatherhood. In the case of a harem herd
that are worth noting. Arboreality allows to acquire valuable consisting of one male reproducer and his females, female

13
Theory in Biosciences (2020) 139:191–207 199

Table 5  The “sexuality core” of the hominid female (Dixson 1998; Basson 2001; Wrangham 2001; Buss and Schmitt 1993; Stoinski et al.
2009a, b; Hauser 1990; Clay et al. 2011; Nadler 1976; Masters and Johnson 1966; Thornhill et al. 1995)
Feature Female
Homo sapiens Chimpanzee (P. Bonobo (P. panis- Gorilla (Gorilla) Last common Core of sexuality
troglodytes) cus) ancestor

Sexual preferences Present—selec- Present—selec- Generally Present—selec- Present—selec- Presence of sexual

tivity tivity absent—possi- tivity tivity preferences—
ble promotion of selectivity
selected males
in the estrus
season
Determinant of Social status, Social status, food Lack of data Social status Social status, food Presence of males’
attractiveness lack of sexual sharing sharing attractiveness
of the opposite aggression, gifts, features preferred
gender physicality by females
Need to protect Present Present Absent—females Present Females need to Need for protection
against sexual are sexually be protected against sexual
aggression available for the against sexual aggression
entire period of aggression
the menstrual
cycle
Sexual inclinations Polyandrism Polyandrism Polyandrism Polyandrism Polyandrism Polyandrism
Sexual drive (pro- Perovulatory Perovulatory Permanent Perovulatory Perovulatory Perovulatory sexual
ceptivity) drive
Copulation calls Present Present Present Present Present Presence of copula-
tion calls
Multiple orgasms Present Reaches orgasms Reaches orgasms Unknown Reaches orgasms Possibility of
reaching multiple
orgasms
Expected level of Depends on social Low Absent Low Low Low level of jeal-
jealousy of the ­structurea ousy of the male
male

Information given by the authors is written in an extended text

a
High level of the woman jealousy of the man—if the woman is dependent on care of one partner and needs his protection (in the patriarchal
social structure); low level of the woman jealousy of the man—if the woman is protected and economically independent, if she has an access to
multiple partners and their resources (e.g., the matricentric Mosuo people)

infidelity had to end with the murder of her young, commit-

ted by a dominant male. Therefore, it was in the interest of
the females to maintain fidelity to the dominant male and
her proceptive behavior turned out to be enough to arouse
him sexually.
Thus, it can be assumed that the hominid LCA (like
chimpanzees) had functioned in the terrestrial and arboreal
environment in the patriarchal social structures, and their
females were receptive in the perovulatory periods and sig-
naled the estrus with anal–genital swelling. The presence
of swelling indicates heterogeneity of the community—the
presence of other males in the herd next to the dominant
one (Table 4).

Hominid jealousy

In human matricentric societies, where a woman resides

Fig. 1  Selective polyandry in her mother’s home and this house protects her and her

13
200 Theory in Biosciences (2020) 139:191–207

Table 6  The “sexuality core” of a hominid male (Dixson 1998; Wrangham 2001; Mosher and Abramson 1977)
Feature Male
Homo sapiens Chimpanzee (P. Bonobo (P. panis- Gorilla (Gorilla) Last common Core of sexuality
troglodytes) cus) ancestor

Sexual preferences Absent—suffi- Absent Absent Absent Absent Lack of sexual pref-
ciently intensive erences—signs of
simulation of estrus activates
estrus activates male sexual
the male sexual behavior
behavior
Determinant of Permanent attrac- Estrous swelling Permanent estrous Proceptivity; Estrous swelling Signs of fertility are
attractiveness tiveness–simula- swelling determinant of
of the opposite tion of estrus attractiveness
gender
Sexual inclinations Promiscuity Promiscuity Promiscuity Promiscuity Promiscuity Promiscuity
Sexual drive Permanent Permanent Permanent Permanent Permanent Permanent sexual
drive
Presence of sexual Present/absenta The dominant Absent—females The dominant The dominant Lack of the sexual
jealousy male is jealous are available male is jealous male was jealous jealousy
of his harem; of his harem; of his harem;
other males—no other males—no other males—no

Information given by the authors is written in an extended text

a
Present—under conditions of limited access to the females (in patriarchal social structures); absent—under conditions of availability of the
females (e.g., war rape)

offspring, where there are no conventional marriages, and resources but not for the males, because they do not have
in the case of matriarchal bonobos—the female jealousy is them. In the case when the female is forced to derive the
not clear. Women in the patriarchal structures are jealous of resources or other benefits from the male, her jealousy takes
the men to whom they allow sexual access in exchange for the form of the jealousy of the male and manifests itself in
the access to the resources that these men have, while the the aggression directed against him and the females threaten-
females of the patriarchal chimpanzees compete with other ing her relationship. That is why the institution of the first
females for access to environmental resources (Table 2). wife (having most rights) is popular among the polygynous
Therefore, it can be concluded that the jealousy of the Homo sapiens societies.
hominid female is directly linked to the patriarchal social Benefits that a woman can derive from a man can, of
structure, and thus derives from interpersonal competition course, be the resources that he has but also protection
for the resources. The chimpanzee female competes for the against sexual aggression and his help in raising her chil-
dren and in the household. When the woman establishes
sexual relations with the man who fulfills her preferences,
then the benefit that she obtains from him will also be her
sexual gratification. In patriarchal societies, the woman has
extremely limited access to sexual partners. Directing the
attention of her man to other women threatens not only her
insufficient sexual satisfaction, but even its loss; her jealousy
takes the form of strictly “sexual jealousy.”
Under patrilocal conditions, the female without any rela-
tives in her community can only count on the male to whom
she belongs. Stopping the male from directing his energy
in search of new sexual objects allows him to redirect this
energy to protect and care for a jealous female and her off-
spring. The jealousy of the male who does not meet female’s
sexual preferences, but provides protection against sexual
aggression and provides support and protection (e.g., dur-
Fig. 2  Tolerant promiscuity ing wars) might ensure the survival of the female and her

13
Theory in Biosciences (2020) 139:191–207 201

offspring. In the Yanomamö tribe, if a woman does not have male, a need for his protection and support, benefits received
male protection, she is at high risk of rape and beating; her from the males in the form of valuable gifts. Under such con-
children may also be killed (Biocca 1970). Therefore, the ditions, it is in the best interests of the female to cut off the
presence of the female jealousy of the male suggests that this male from all other females—the tool that the female will
line has evolved mainly in the patriarchal and thus patrilocal use is sexual jealousy. Facilitated access to the females (as
environment, where the females without the support of rela- in matricentric human communities) significantly reduces,
tives fought for survival and reproduction in the company of and promiscuity of the females (bonobo) even eliminates the
the male to whom she belonged. sexual jealousy, both male and female.
The presence of the male jealousy only in sexual rela-
tions, in which the man claims the rights to the woman, the Hominid “cores of sexuality”
low level of man’s investment and involvement in raising
children, and his preference, if there is such a possibility, of Assuming no social structure, the hominid females’ “selec-
virgins for wives indicates that this jealousy is not related to tive polyandry” would be estrous sexual relationships with
the need for certainty of paternity that justifies investments more than one male of her choice (preferably at the same
in the offspring. However, these factors clearly indicate the time), where each of them will be attractive enough to lead
man’s tendency to monopolize the sexuality of the woman. her to an orgasm (Table 5). This will allow sperm rivalry of
Under certain conditions, the man seems to seek to suppress several high-quality males in her reproductive tracts, giving
the woman’s sexuality. These conditions are socially lim- her a chance for the highest quality offspring. Out-estrus sex-
ited access to the females, present in all patriarchal primate uality of the hominid female allows her to select the males
structures. during non-ovulatory periods. She continues to “flirt” with
Symons (1979) argues that the male sexual jealousy is the male, who activates her desire. Through the “flirt” the
more unchangeable and “obligatory” than the female one female chooses sexual partners, and by loud vocalizations
but suggests that the male sexual jealousy can be suppressed during copulation, she stimulates them sexually, informs
or inactivated in the context of polyamory. He suggests that them about the state of her ovulation and about their ability
this is possible because of male desire for sexual diversity. to take part in exciting copulation under the sperm competi-
Having sexual access to other women in exchange, a given tion conditions. The female may show a low level of jealousy
man will allow other men to have sex with his wife. It is of the males who, when flirting, give her presents.
worth adding here that in the conditions of polyamory, no The chimpanzee males, who are not dominants, give
man will feel the need to marry. females food and build relationships with them in this way.
In a traditional patriarchal resource-accumulating com- Male–female food sharing among gorillas in the wild has
munity, the man is obliged to invest in his woman and her not been found. A gorilla dominant male has no competi-
children because they are economically dependent on him. tion in the herd and has a monopoly on the females, and he
These investments will be an additional factor (not a source) does not have to seek their favors. However, in captivity,
heightening his sexual jealousy toward his woman—the man when the female was separated from the male by bars (so
does not want someone else to meet his sexual needs through she did not belong to him), he gave her an apple. American
the woman he maintains; he does not want to invest in some- women (probably not only them), expect valuable gifts from
body else’s offspring and, at his own expense, increase the candidates for a fleeting contact. In the case of the women
reproductive success of other men. In matricentric commu- from the matricentric societies, the end of the inflow of gifts
nities, where there are no conventional long-term marriages from the man means the end of mutual relations. Therefore,
and therefore, access to the women is facilitated, and where in conditions of independence from the male, the hominid
the man does not invest heavily in either the woman or in his female expects male’s gifts to improve the quality of her life
offspring, the male jealousy does not apply. and facilitate her reproductive success.
The above analysis indicates that the patriarchal social The female Coolidge effect was found in rodent species
structures condition the escalation of the jealousy, in both in which the females benefited from copulation with many
the hominid female and male (Table 3). The dominance of males (Lisk and Baron 1982; Lester and Gorzalka 1988;
the males entails the ownership of the females, so mutual Ventura-Aquino and Fernández-Guasti 2013). Therefore, the
free sexual access is limited. Limited access to sexual part- hominid female should also be concerned with the Coolidge
ners is a sufficient factor to induce a high level of the jeal- effect. The more high-quality males she flirts with, the more
ousy in the males claiming rights to the females. The female gifts she gets from them and the more her quality of life
jealousy of the male, in the case when she has free access increases. Cost-effectiveness of polyandrous behavior of the
to environmental resources, would not be justified. How- hominid female is matched with her multiple orgasms and
ever, the male becomes substantially conditions the female’s copulatory vocalizations. If the hominid female did not con-
being due to such factors as: economic dependence on the cern the Coolidge effect, the presence of women’s out-estrus

13
202 Theory in Biosciences (2020) 139:191–207

“triggered” sexuality would not be justified. The woman applicable at the expense of giving up meeting psychological
would also be deprived of the possibility of semen selec- and biological needs of both genders.
tion. However, she is equipped with appropriate tools such If hypothetical hominid species gained free access to the
as orgasms. Being limited to one male significantly reduces environmental and sexual resources it needed, then both gen-
the number of gifts received by the female, and when he ders would use only their “cores of sexuality”: the males
claims to own her, she does not receive gifts at all—as in would be tolerant and promiscuous, and the females would
the case of the gorillas. choose the best of them. However, these “cores” do not coin-
A hominid male “core” has been identified as the “tol- cide with each other and create a social conflict in the form
erant promiscuity” (Table 6). Tolerability reduces male’s of the males rejected by the females.
reproduction chances by the sperm competition but it gives A hypothetical, primary heterogeneous community of the
him exciting promiscuous copulations (for men it is more hominids, in which no social structure yet exists, will be
attractive to participate in a promiscuous copulation than in considered below. The males are not yet able to cooperate,
a sexual act of a heterosexual couple) (Mosher and Abram- the females clearly signal ovulation, both genders have free
son 1977). Group sex participants (swingers) use rules to access to food, and both have the aforementioned “sexu-
create more freedom and safety to act (Harviainen and Frank ality cores” (in the form of “tolerant promiscuity” of the
2018). Therefore, the hominid male will be inclined to both: male and “selective polyandry” of the female). By adopting
monogamous sex—increasing his reproductive chances, and an early stage of social evolution, it is not a very numer-
multi-male sex—eliminating the need for inter-male aggres- ous herd. Both young males and females migrate between
sion and giving him more satisfaction. However, it should be flocks. During the estrus periods, the females will seek to
noted here that the hominid male is inherently jealous of the mate only with the most attractive males. So a small group
females to which he claims rights: the LCA has been func- of the preferred and a larger group of the rejected males by
tioning in the patriarchate and thus in conditions of limited the females will be formed. All the males are promiscuous
access to the females. and they have high sexual needs, so they will demand copu-
The hominid male will be also inclined to both, multi- lation. The refused males will begin to apply sexual coercion
male and multi-female group sex. The multi-male group to the females and aggression toward the privileged males.
sex will stimulate his testicles to produce large amounts of Competition for access to the females under the rape condi-
sperm for rivalry in the female reproductive tract, as it is the tions will begin. In this situation, the females will seek pro-
case of the chimpanzee or bonobo (Møller 1988). Copula- tection of the preferred males. These males are attractive to
tion with multiple females at the same time, analogically to the females—so they are also healthy and strong. Therefore,
a multi-male act, will be a strong sexual stimulus, causing they will restrict access to the females from the unwanted
testicles to produce semen, which he has to provide all the males. Under conditions of competition for the females, a
females with. The more females in the act, the higher the discussion about mutual inter-male sexual tolerance is no
degree of excitement of the male. longer valid. In a herd with not many members, the strongest
male (unable to cooperate) will protect all the females in the
“Cores of sexuality” and the patriarchate estrus, and thus will gain a monopoly on access to them, as it
is the case of gorillas. In numerous human and chimpanzee
Analyzing contemporary views on sexual strategies for communities, patriarchy is maintained due to the coopera-
human mate choice Easton et al. (2015) divide both women’s tion of the males and thus to the male hierarchy.
and men’s strategies into short-term and long-term mating. The above brief analysis of the “primary hominid” behav-
The “sexuality cores” designated here do not indicate any ior reveals that having particular “cores of sexuality,” under
hominid long-term mating tendency. This strategy is associ- conditions of periodical estrus signaling, condemns the
ated with limiting sexual freedom of the partner, so it is not a hominid females to sexual aggression and generates a need
comfortable strategy and cannot be a part of “human nature.” for their protection. Therefore, the whole hominid family is
This strategy neither occurs in bonobos nor in matricentric condemned to a patriarchal social structure and, as a result,
Homo sapiens. Among chimpanzees, there is fierce competi- to sexual coercion and adultery which allows the females to
tion for access to fertile females, while among gorillas nei- make a choice. Patriarchy, combined with the selectivity of
ther gender chooses a partner. Long-term mating is therefore the females, is the cause of extreme sexual restrictions for
a strategy only characteristic of patriarchal Homo sapiens the males. This paves the way for searching other sexual
with long-term property rights on the sexual partner and tension unloading channels such as war rapes, prostitution,
extremely limited mutual choice of partners that results from harem tendencies. Polygyno-monogamous tribal patriarchy,
these rights. Therefore, this strategy is only the outcome of in conditions of the permanently simulated estrus, will also
restrictive Homo sapiens patriarchal social conditions and is generate marital rape and marital prostitution.

13
Theory in Biosciences (2020) 139:191–207 203

The exceptions from the patriarchal fate are: the bonobo, which female he fertilizes—each of this male’s offspring
whose females, resigning from selectivity, satisfy the pro- will inherit his tendency to opportunism. Under conditions
miscuous needs of the males, so they do not compete for of free access to the females, the hominid male will not
access to them; the Homo sapiens matricentric communities, compete for them, but he will be inclined to (exciting for
characterized by the lack of males’ monopoly on the females him) copulations in the conditions of sperm rivalry and
and, as a result, by facilitated access to them. The females high demand for sperm. In turn, the possibility of choice is
from both species simulate the state of permanent ovulation, a key condition for the hominid female: the female chooses
which suggests that estrus simulation is a necessary condi- the best males from those available (“selective polyandry”)
tion but not enough to reduce a patriarchal social system. and their sperm will compete in her reproductive tracts for
Homo sapiens mostly functions in patriarchal structures; access to the ovum. She will bear a full parental invest-
therefore, simulation of the permanent ovulation, without ment in the future.
stretching the female’s sex drive for the entire menstrual The natural (free) mating cycle of a Homo sapiens female
cycle, does not guarantee social structures without the male consists, therefore, of the following stages:
hierarchy.
The female’s needs are mutually exclusive in patriarchal 1. Outside the ovulatory period, following the “triggered
structures. The need for care creates the sexual monopoly on sexuality,” from her surroundings the female chooses the
the female in care of the male. This monopoly is the cause males who arouse her desire (also by means of gifts) and
of her limited availability and thus, of her limited sexual by flirting, she informs them about her potential sexual
freedom. Patriarchal structures also restrict women’s free accessibility;
choice, limiting it to infidelity. Meeting sexual needs of both 2. In the first phase of the ovulation period, the woman may
genders in an optimized way provides matricentric social have a tendency to perform a mating dance, announcing
structures of Homo sapiens, where her relatives take care of the surrounding men her estrous state. A spontaneous
the woman and her children and thanks to that, she can enjoy mating dance of the woman was observed on the 12th
sexual freedom. From the males, she only expects physi- day of her menstrual cycle. It was performed publicly,
cal attractiveness, gifts and lack of aggression. Matricentry in conditions of reduced sexual restrictions (the authors’
does not provide the men with conditions for total promis- own observation).
cuity because giving them facilitated access to the women 3. In the estrus phase, the female copulating with the male
imposes on them meeting women’s preferences. However, of her choice loudly vocalizes, invoking the other males
the lack of a male hierarchy causes that these communi- befriended with her and by this, stimulating her partner
ties break down under the influence of aggressive organ- (group copulation with invoked males is more exciting
ized actions of patriarchal groups, focused on the proprietary to him than monogamous mating, loud vocalization
acquisition of the women. also “raises” the social status of the copulating male).
By vocalization, the female increases the vigilance of
“her” males and therefore, also her own safety; unwanted
Summary males will not be allowed to her by the presence of the
chosen ones;
The analysis of the hominids sexual behavior carried out 4. During copulation, the female using the potential of
in this paper indicates that their social structures manifest multiple orgasms achieves the orgasm with each of sub-
various uses of socio-environmental conditions to meet their sequently selected males, sucking their sperm into the
biological, existential and emotional needs. This behavior is genital tract. In such conditions, the ovum is fertilized
based on the common to all the hominids “cores of sexual- by the “best of the best” spermatozoid.
ity.” These “cores” are evolutionary constant psychosomatic
mechanisms that direct the organism at the maximization By the presence of out-estrus sexuality, the hominid
of its reproductive success. Due to a huge disparity regard- female, while selecting the males, can be guided by the “trig-
ing parental investments between the hominid males and gered sexuality”—she continues flirting with the male who
females, these “cores” differ from each other. activates her desire, also by bestowing her. The women’s
The results of this analysis correlate with the Trivers “triggered sexuality” should therefore be treated as an evo-
(1972) theorem. However, it has been also shown here lutionary tool inherited from common ancestor: the females
that the “tolerantly promiscuous” hominid male, by his from all four hominid species also copulate in out-estrus
very nature, does not choose female partners at all. He is periods. This is used to determine the quality of the males,
satisfied if the female is able to sexually stimulate him. their selection, and it is aimed at selecting those, who will
The hominid male (including the man) does not show be able to give her one of the orgasms during multi-male
paternal instincts, so it does not make him any difference estrous copulation.

13
204 Theory in Biosciences (2020) 139:191–207

Table 7  Hominid social structures and the females’ sexuality

Feature Female sexual prefer- Ovulation signaling Female sexual drive Group aggression of Social structure
ences (sexual attractiveness of males (war)
the females)

Species
Gorilla (Gorilla) Present Proceptive behavior dur- Perovulatory Absent—lack of male Patriarchy
ing estrus cooperation
Bonobo (P. paniscus) Absent (opportunism) Simulators of the perma- Permanent Absent—lack of male Matriarchy
nent estrus hierarchy
Chimpanzee (P. trog- Present Estrous anal–genital Perovulatory Present Patriarchy
lodytes) swelling
Homo sapiens Present Simulators of the perma- Perovulatory Present Patriarchy
nent estrus Absent—lack of male Matricentrism
hierarchy

Free multi-male group sex as a function of sexual selec- Open Access This article is licensed under a Creative Commons Attri-
tion seems to be the basis for the effective evolution of bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
the hominids. The institution of marriage, both traditional as you give appropriate credit to the original author(s) and the source,
(polygynous) and Euramerican (monogamous), blocking the provide a link to the Creative Commons licence, and indicate if changes
natural polyandrous needs of the women, to a large extent were made. The images or other third party material in this article are
blocks the influence of sexual selection in the species Homo included in the article’s Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
sapiens. If the women had existential security (e.g., thanks the article’s Creative Commons licence and your intended use is not
to relatives) and sexual freedom (as in matricentric com- permitted by statutory regulation or exceeds the permitted use, you will
munities), they could realize their “core of sexuality,” and need to obtain permission directly from the copyright holder. To view a
for each of their pregnancies they would acquire the best copy of this licence, visit http://creat​iveco​mmons​.org/licen​ses/by/4.0/.
genetic material.
The hominid females determine the social structures
through their sexual selectivity or opportunism, proceptivity, References
and the way of estrus signaling (continuous or periodic). The
Adams DB (1983) Why there are so few women warriors. Behav Sci
need for selection of the hominid female periodically signal- Res 18(3):196–212
ing ovulation determines the patriarchal social structure (the Akers JS, Schildkraut DS (1985) Regurgitation/reingestion and
common chimpanzee, the gorilla). The need for selection of coprophagy in captive gorillas. Zoo Biol 4(2):99–109
the females feeling perovulatory desire and simulating the Alexander RD, Noonan KM (1979) Concealment of ovulation, parental
care, and human social evolution. In: Chagnon NA, Irons W (eds)
permanent state of ovulation, in conditions of wars and thus, Evolutionary biology and human social behavior. Duxbury Press,
of the male hierarchy, also results in the patriarchal structure North Scituate, pp 436–453
(Homo sapiens). On the other hand, the absence of wars and Allen ML, Lemmon WB (1981) Orgasm in female primates. Am J
thus, of the male hierarchy (the hierarchy falls as a result Primatol 1(1):15–34
Azadzoi KM, Siroky MB (2010) Neurologic factors in female sexual
of adultery of the females) allows such females to create function and dysfunction. Korean J Urol 51(7):443–449
structures based on their kinship (Homo sapiens). Sexual Baker RR, Bellis MA (1988) “Kamikaze” sperm in mammals? Anim
opportunism of the females being permanently receptive, Behav 36:936–939
proceptive and simulating the permanent state of ovulation, Baker RR, Bellis MA (1989) Elaboration of the kamikaze sperm
hypothesis: a reply to Harcourt. Anim Behav 37:865–867
by breaking the male dominance, creates a matriarchal type Baker RR, Bellis MA (1993a) Human sperm competition: ejaculate
of the community (the bonobo) (Table 7). adjustment by males and the function of masturbation. Anim
If the women were not only permanently sexually attrac- Behav 46:861–885
tive, but also stretched the sexual drive for the entire men- Baker RR, Bellis MA (1993b) Human sperm competition: ejaculate
manipulation by females and a function for the female orgasm.
strual cycle, as the bonobo females do, then undoubtedly, in Anim Behav 46(5):887–909
the species Homo sapiens, neither marriage nor war would Baker R, Bellis MA (2014) Human sperm competition: copulation,
appear. Moreover, women’s estrous sexual desire indicates masturbation and infidelity. Hard Nut Books, Newcastle
that in the history of the hominin line evolution, a matriar- Barratt CL, Kay V, Oxenham SK (2009) The human spermatozoon: a
stripped down but refined machine. J Biol 8(7):63
chal structure, equivalent to that of bonobo’s, could not have Basson R (2001) Human sex-response cycles. J Sex Marital Ther
taken place. 27(1):33–43

13
Theory in Biosciences (2020) 139:191–207 205

Biocca E (1970) Yanoáma: the narrative of a white girl kidnapped by Goodall J (1986) The chimpanzees of Gombe: patterns of behavior. The
Amazonian Indians. Dutton, New York Belknap Press of Harvard University Press, Cambridge
Birkhead TR, Pizzari T (2002) Postcopulatory sexual selection. Nat Gough EK (1959) The Nayars and the definition of marriage. J R
Rev Genet 3(4):262–273 Anthropol Inst G B Irel 89(1):23–34
Brewer G, Hendrie CA (2011) Evidence to suggest that copulatory Harcourt AH, Stewart KJ (1987) The influence of help in contests on
vocalizations in women are not a reflexive consequence of dominance rank in primates: hints from gorillas. Anim Behav
orgasm. Arch Sex Behav 40(3):559–564 35(1):182–190
Bullivant SB, Sellergren SA, Stern K, Spencer NA, Jacob S, Men- Harcourt AH, Fossey D, Stewart KJ, Watts DP (1980) Reproduction in
nella JA, McClintock MK (2004) Women’s sexual experience wild gorillas and some comparisons with chimpanzees. J Reprod
during the menstrual cycle: identification of the sexual phase Fertil (Supplement) 28:59–70
by noninvasive measurement of luteinizing hormone. J Sex Res Harcourt AH, Harvey PH, Larson SG, Short RV (1981) Testis
41(1):82–93 weight, body weight and breeding system in primates. Nature
Burbank VK (1987) Female aggression in cross-cultural perspective. 293(5827):55–57
Behav Sci Res 21(1–4):70–100 Harviainen JT, Frank K (2018) Group sex as play: rules and transgres-
Buss DM (2000) The dangerous passion: why jealousy is as neces- sion in shared non-monogamy. Games Cult 13(3):220–239
sary as love and sex. Simon & Schuster, New York Hauser MD (1990) Do chimpanzee copulatory calls incite male: Male
Buss DM (2013) Sexual jealousy. Psihol Teme 22(2):155–182 competition? Anim Behav 39:596–597
Buss DM, Schmitt DP (1993) Sexual strategies theory: an evolu- Hennig W (1965) Phylogenetic systematics. Annu Rev Entomol
tionary perspective on human mating. Psychol Rev 100(2):204 10(1):97–116
Chevalier-Skolnikoff S (1974) Male-female, female-female, and Hewlett BS (1991) Demography and childcare in preindustrial socie-
male-male sexual behavior in the stumptail monkey, with ties. J Anthropol Res 47(1):1–37
special attention to the female orgasm. Arch Sex Behav Hill K, Kaplan H (1988) Tradeoffs in male and female reproductive
3(2):95–116 strategies among the ache: Part 1. In: Betzig L, Borgerhoff-
Clay Z, Pika S, Gruber T, Zuberbühler K (2011) Female bonobos use Mulder M, Turke P (eds) Human reproductive behavior: a Dar-
copulation calls as social signals. Biollet 7(4):513–516 winian perspective. Cambridge University Press, Cambridge, pp
Crick J, Suchak M, Eppley TM, Campbell MW, de Waal FB (2013) 277–289
The roles of food quality and sex in chimpanzee sharing behavior Jaeggi AV, Van Schaik CP (2011) The evolution of food sharing in
(Pan troglodytes). Behaviour 150(11):1203–1224 primates. Behav Ecol Sociobiol 65(11):2125–2140
Daly M, Wilson M, Weghorst SJ (1982) Male sexual jealousy. Ethol Jaeggi AV, de Groot E, Stevens JMG, van Schaik CP (2013) Mecha-
Sociobiol 3:11–27 nisms of reciprocity: testing for short-term contingency of
De Waal FBM (1982) Chimpanzee politics: sex and power among apes. grooming and food sharing in bonobos and chimpanzees. Evol
Jonathan Cape, London Hum Behav 34:69–77
De Waal FBM (1987) Tension regulation and nonreproductive func- James WH (1981) The honeymoon effect on marital coitus. J Sex Res
tions of sex in captive Bonobos (Pan paniscus). Natl Geogr Res 17(2):114–123
3(3):318–335 Kano T (1980) Social behavior of wild pygmy chimpanzees (Pan panis-
De Waal FBM (1997) The chimpanzee’s service economy: food for cus) of Wamba: a preliminary report. J Hum Evol 9(4):243–260
grooming. Evol Hum Behav 18:375–386 Kano T (1992) The last ape: Pygmy chimpanzee behavior and ecology.
De Waal FBM, Lanting F (1997) Bonobo: the forgotten ape. University Stanford University Press, Stanford
of California Press, Berkeley Kaplan H, Hill K (1985) Hunting ability and reproductive success
Divale WT (1972) Systemic population control in the middle and upper among male Ache foragers: Preliminary results. Curr Anthropol
palaeolithic: inferences based on contemporary hunter-gatherers. 26(1):131–133
World Archaeol 4(2):222–243 Kaplan H, Hill K, Hawkes K, Hurtado A (1984) Food sharing among
Dixson AF (1981) The natural history of the gorilla. Columbia Uni- Ache hunter-gatherers of Eastern Paraguay. Curr Anthropol
versity Press, New York 25(1):113–115
Dixson AF (1998) Primate sexuality. Comparative studies of the Pro- Kaplan H, Hill K, Cadeliña RV, Hayden B, Hyndman DC, Preston RJ,
simians, monkeys, apes, and human beings. Oxford University Smith EA, Stuart DE, Yesner DR (1985) Food sharing among
Press, Oxford ache foragers: tests of explanatory hypotheses. Curr Anthropol
Dunn KM, Cherkas LF, Spector TD (2005) Genetic influences on 26:223–246
variation in female orgasmic function: a twin study. Biollet Keeley LH (1996) War before civilization. Oxford University Press,
1(3):260–263 New York
Easton JA, Goetz CD, Buss DM (2015) Human mate choice, evolu- Kinsey AC, Pomeroy WB, Martin CE, Sloan S (1948) Sexual behavior
tion. In: Wright JD (editor-in-chief) International encyclopedia in the human male, vol 1. Saunders, Philadelphia
of the social & behavioral sciences, 2nd edn., vol 11. Elsevier, Kramer KL (2011) The evolution of human parental care and recruit-
Oxford, pp 340–347 ment of juvenile help. Trends Ecol Evol 26(10):533–540
Furuichi T (1987) Sexual swelling, receptivity, and grouping of Lancaster JB, Lancaster CS (1983) Parental investment: the hominid
wild pygmy chimpanzee females at Wamba, Zaire. Primates adaptation. In: Ortner D (ed) How humans adapt: a biocultural
28(3):309–318 odyssey. Smithsonian Press, Washington, DC, pp 33–56
Furuichi T (1997) Agonistic interactions and matrifocal dominance Le Jeune P (1897/2009) Les relations des Jesuites. Toronto Public
rank of wild bonobos (Pan paniscus) at Wamba. Int J Primatol Library, Toronto
18(6):855–875 Lester GL, Gorzalka BB (1988) Effect of novel and familiar mating
Gallup GG, Burch RL, Zappieri ML, Parvez RA, Stockwell ML, Davis partners on the duration of sexual receptivity in the female ham-
JA (2003) The human penis as a semen displacement device. ster. Behav Neural Biol 49(3):398–405
Evol Hum Behav 24(4):277–289 Lisk RD, Baron G (1982) Female regulation of mating location and
Gomes CM, Boesch C (2009) Wild chimpanzees exchange meat for acceptance of new mating partners following mating to sexual
sex on a long-term basis. PLoS ONE 4(4):e5116 satiety: the Coolidge effect demonstrated in the female golden
hamster. Behav Neural Biol 36(4):416–421

13
206 Theory in Biosciences (2020) 139:191–207

Locke DP, Hillier LW, Warren WC, Worley KC, Nazareth LV, Muzny Pusey AE, Schroepfer-Walker K (2013) Female competition in chim-
DM, Yang SP, Wang Z, Chinwalla AT, Minx P, Mitreva M, Cook panzees. Philos Trans R Soc B 368:20130077
L, Delehaunty KD, Fronick C, Schmidt H, Fulton LA, Fulton RS, Rowland DL (2006) Neurobiology of sexual response in men and
Nelson JO, Magrini V, Pohl C, Graves TA, Markovic C, Cree A, women. CNS Spectr 11(S9):6–12
Dinh HH, Hume J, Kovar CL, Fowler GR, Lunter G, Meader Savage-Rumbaugh ES, Wilkerson BJ (1978) Socio-sexual behavior in
S, Heger A, Ponting CP, Marques-Bonet T, Alkan C, Chen L, Pan paniscus and Pan troglodytes: a comparative study. J Hum
Cheng Z, Kidd JM, Eichler EE, White S, Searle S, Vilella AJ, Evol 7(4):327–344
Chen Y, Flicek P, Ma J, Raney B, Suh B, Burhans R, Herrero J, Scally A, Dutheil JY, Hillier LW, Jordan GE, Goodhead I, Herrero J,
Haussler D, Faria R, Fernando O, Darré F, Farré D, Gazave E, Hobolth A, Lappalainen T, Mailund T, Marques-Bonet T, McCa-
Oliva M, Navarro A, Roberto R, Capozzi O, Archidiacono N, rthy S, Montgomery SH, Schwalie PC, Tang YA, Ward MC, Xue
Della Valle G, Purgato S, Rocchi M, Konkel MK, Walker JA, Y, Yngvadottir B, Alkan C, Andersen LN, Ayub Q, Ball EV, Beal
Ullmer B, Batzer MA, Smit AF, Hubley R, Casola C, Schrider K, Bradley BJ, Chen Y, Clee CM, Fitzgerald S, Graves TA, Gu
DR, Hahn MW, Quesada V, Puente XS, Ordoñez GR, López-Otín Y, Heath P, Heger A, Karakoc E, Kolb-Kokocinski A, Laird GK,
C, Vinar T, Brejova B, Ratan A, Harris RS, Miller W, Kosiol C, Lunter G, Meader S, Mort M, Mullikin JC, Munch K, O’Connor
Lawson HA, Taliwal V, Martins AL, Siepel A, Roychoudhury A, TD, Phillips AD, Prado-Martinez J, Rogers AS, Sajjadian S,
Ma X, Degenhardt J, Bustamante CD, Gutenkunst RN, Mailund Schmidt D, Shaw K, Simpson JT, Stenson PD, Turner DJ, Vigi-
T, Dutheil JY, Hobolth A, Schierup MH, Ryder OA, Yoshinaga lant L, Vilella AJ, Whitener W, Zhu B, Cooper DN, de Jong P,
Y, de Jong PJ, Weinstock GM, Rogers J, Mardis ER, Gibbs RA, Dermitzakis ET, Eichler EE, Flicek P, Goldman N, Mundy NI,
Wilson R (2011) Comparative and demographic analysis of Ning Z, Odom DT, Ponting CP, Quail MA, Ryder OA, Searle
orang-utan genomes. Nature 469(7331):529 SM, Warren WC, Wilson RK, Schierup MH, Rogers J, Tyler-
Malinowski B (1929) The sexual life of savages in North-Western Smith C, Durbin R (2012) Insights into hominid evolution from
Melanesia. Routledge, London the gorilla genome sequence. Nature 483(7388):169–175
Manson JH, Wrangham RW, Boone JL, Chapais B, Dunbar RIM, Schaller GB (1963) The mountain gorilla: ecology and behavior. Uni-
Ember CR, Irons W, Marchant LF, McGrew WC, Nishida T, versity of Chicago Press, Chicago
Paterson JD, Smith EA, Stanford CB, Worthman CM (1991) Semple S (2001) Individuality and male discrimination of female copu-
Intergroup aggression in chimpanzees and humans [and Com- lation calls in the yellow baboon. Anim Behav 61(5):1023–1028
ments and Replies]. Current Anthropology 32(4):369–390 Shackelford TK, LeBlanc GJ, Weekes-Shackelford VA, Bleske-Rechek
Masters WH, Johnson VE (1966) Human sexual response. Little, AL, Euler HA, Hoier S (2002) Psychological adaptation to
Brown, Boston human sperm competition. Evol Hum Behav 23:123–138
Matsumoto-Oda A (1999) Female choice in the opportunistic mating Small MF (1989) Female choice in nonhuman primates. Am J Phys
of wild chimpanzees (Pan troglodytes schweinfurthii) at Mahale. Anthropol 32(S10):103–127
Behav Ecol Sociobiol 46(4):258–266 Smuts BB, Smuts RW (1993) Male aggression and sexual coercion of
Mattison SM (2011) Evolutionary contributions to solving the “Mat- females in nonhuman primates and other mammals: evidence and
rilineal Puzzle”. Hum Nat 22(1–2):64–88 theoretical implications. Adv Study Behav 22(22):1–63
Mautz BS, Wong BB, Peters RA, Jennions MD (2013) Penis size inter- Stanford CB (1998) The social behavior of chimpanzees and bonobos:
acts with body shape and height to influence male attractiveness. empirical evidence and shifting assumptions. Curr Anthropol
Proc Natl Acad Sci 110(17):6925–6930 39(4):399–420
Mitani JC, Watts DP (2001) Why do chimpanzees hunt and share meat? Stanford CB, Wallis J, Mpongo E, Goodall J (1994) Hunting decisions
Anim Behav 61:915–924 in wild chimpanzees. Behaviour 131(1):1–18
Møller AP (1988) Ejaculate quality, testes size and sperm competition Stewart KJ (1977) The birth of a wild mountain gorilla (Gorilla gorilla
in primates. J Hum Evol 17(5):479–488 beringei). Primates 18:965–976
Mosher DL, Abramson PR (1977) Subjective sexual arousal to films of Stoinski TS, Perdue BM, Legg AM (2009a) Sexual behavior in female
masturbation. J Consult Clin Psychol 45(5):796–807 western lowland gorillas (Gorilla gorilla gorilla): evidence for
Nadler RD (1976) Sexual behavior of captive lowland gorillas. Arch sexual competition. Am J Primatol 71(7):587–593
Sex Behav 5(5):487–502 Stoinski TS, Rosenbaum S, Ngaboyamahina T, Vecellio V, Ndagiji-
Otterbein KF (1970) The evolution of war: a cross-cultural study. mana F, Fawcett K (2009b) Patterns of male reproductive behav-
HRAF Press, New Heaven iour in multi-male groups of mountain gorillas: examining theo-
Pfefferle D, Brauch K, Heistermann M, Hodges JK, Fischer J (2008) ries of reproductive skew. Behaviour 146(9):1193–1215
Female Barbary macaque (Macaca sylvanus) copulation calls do Symons D (1979) The evolution of human sexuality. Oxford University
not reveal the fertile phase but influence mating outcome. Proc R Press, Oxford
Soc B Biol Sci 275:571–578 Thornhill R, Gangestad SW, Comer R (1995) Human female orgasm
Pound N (2002) Male interest in visual cues of sperm competition risk. and mate fluctuating asymmetry. Anim Behav 50(6):1601–1615
Evol Hum Behav 23(6):443–466 Todt D, Pohl R (1984) Communicative strategies in estrous Barbary
Pradhan GR, Engelhardt A, van Schaik CP, Maestripieri D (2006) The ape females (Macaca sylvanus) during copulation behaviour:
evolution of female copulation calls in primates: a review and a advertising, triggering, affiliating. Verhandlungen der Deutschen
new model. Behav Ecol Sociobiol 59(3):333–343 Zoologischen Gesellschaft 7:225
Prüfer K, Munch K, Hellmann I, Akagi K, Miller JR, Walenz B, Koren Townsend SW, Deschner T, Zuberbühler K (2008) Female chimpan-
S, Sutton G, Kodira C, Winer R, Knight JR, Mullikin JC, Meader zees use copulation calls flexibly to prevent social competition.
SJ, Ponting CP, Lunter G, Higashino S, Hobolth A, Dutheil J, PLoS ONE 3(6):e2431
Karakoç E, Alkan C, Sajjadian S, Catacchio CR, Ventura M, Trivers RL (1972) Parental investment and sexual selection. In: Camp-
Marques-Bonet T, Eichler EE, André C, Atencia R, Mugisha bell B (ed) Sexual selection and the descent of man. Chicago,
L, Junhold J, Patterson N, Siebauer M, Good JM, Fischer A, Aldine, pp 136–179
Ptak SE, Lachmann M, Symer DE, Mailund T, Schierup MH, Troisi A, Carosi M (1998) Female orgasm rate increases with male
Andrés AM, Kelso J, Pääbo S (2012) The bonobo genome dominance in Japanese macaques. Anim Behav 56(5):1261–1266
compared with the chimpanzee and human genomes. Nature Ventura-Aquino E, Fernández-Guasti A (2013) Reduced proceptiv-
486(7404):527–531 ity and sex-motivated behaviors in the female rat after repeated

13
Theory in Biosciences (2020) 139:191–207 207

copulation in paced and non-paced mating: effect of changing Wrangham RW (1982) Mutualism, kinship and social evolution. In:
the male. Physiol Behav 120:70–76 Bertram BCR (ed) Current problems in sociobiology. Cambridge
Vervaecke H, De Vries HAN, Van Elsacker L (2000) Dominance and University Press, Cambridge, pp 269–289
its behavioral measures in a captive group of bonobos (Pan panis- Wrangham RW (1987) The significance of African apes for recon-
cus). Int J Primatol 21(1):47–68 structing human social evolution. In: Kinze WG (ed) The evolu-
Waldinger MD, Quinn P, Dilleen M, Mundayat R, Boolell M, Sch- tion of human behavior: primate models. SUNY Press, Albany,
weitzer DH (2006) A multi-national population survey of intra- pp 51–71
vaginal ejaculation latency time. J Sex Res 43(1):29 Wrangham RW (1993) The evolution of sexuality in chimpanzees and
Wallen K, Lloyd EA (2011) Female sexual arousal: genital anatomy bonobos. Hum Nat 4:47–79
and orgasm in intercourse. Horm Behav 59(5):780–792 Wrangham RW (2001) Out of the Pan, into the fire: how our ancestors’
Watts DP (1989) Infanticide in mountain gorillas: new cases and a evolution depended on what they ate. In: De Waal FBM (ed) Tree
reconsideration of the evidence. Ethology 81(1):1–18 of origin: what primate behavior can tell us about human social
Watts DP (1991) Mountain gorilla reproduction and sexual behavior. evolution. Harvard University Press, Cambridge, pp 119–143
Am J Primatol 24:211–225 Wrangham RW, Peterson D (1996) Demonic males: apes and the ori-
Weiss P, Brody S (2009) Original research–psychology: women’s gins of human violence. Houghton, Mifflin and Company, Boston
partnered orgasm consistency is associated with greater dura- Yerkes RM (1941) Conjugal contrasts among chimpanzees. J Abnorm
tion of penile-vaginal intercourse but not of foreplay. J Sex Med Soc Psychol 36(2):175–199
6(1):135–141
Wilson DE, Reeder DM (eds) (2005) Mammal species of the world: a Publisher’s Note Springer Nature remains neutral with regard to
taxonomic and geographic reference (Vol. 1). The Johns Hopkins jurisdictional claims in published maps and institutional affiliations.
University Press, Baltimore
Wrangham RW (1979) On the evolution of ape social systems. Soc Sci
Inf 18(3):336–368

13