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Male homosexual preference MHP has long been of interest to scholars studying the evolution of human sexuality. Indeed, MHP is partially heritable, induces a reproductive cost and is common. MHP has thus been considered a Darwinian paradox.

Several questions arise when MHP is considered in an evolutionary context. At what point did MHP appear in the human evolutionary history? Is MHP present in all human groups? These questions were addressed here, using data from the anthropological and archaeological literature.

The conditions under which it is possible to affirm that MHP was present in past societies are discussed. Furthermore, using anthropological reports, the presence or absence of MHP was documented for societies, allowing us to conclude that evidence of the absence of MHP is available for some societies.

A recent evolutionary hypothesis has argued that social stratification together with hypergyny the hypergyny hypothesis are necessary conditions for the evolution of MHP. Here, the link between the level of stratification and the probability of observing MHP was tested sexual an unprecedented large dataset.

Furthermore, the test was performed for the first time by controlling for the phylogenetic non-independence between societies. A positive relationship was observed between the level of social stratification and the probability of observing MHP, supporting the hypergyny hypothesis. This is an open access article distributed under the terms homo the Creative Commons Attribution Licensewhich permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Data Availability: All relevant data homo within the paper and its Supporting Information files. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Competing interests: The authors have declared that no competing interests exist. The origin and maintenance of MHP in humans has long been a matter of interest [ 7 ]. When did MHP first arise? Socially induced homosexual behavior has also been sexual in humans, for example when women are not available [ 1112 ] or in a ritualized form [ 13 ].

MHP has also been described in domesticated boys [ 1415 ], suggesting boys homosexual preference could be at least indirectly selected for.

No clear boys of MHP have been documented in any non-human species outside of undisturbed social environments i. Apparently, MHP appears to be restricted to humans. When did MHP appear during the course of human evolution? Many authors have suggested that MHP dates back to prehistoric [ 16 — 19 ] or early historic time [ 20 ], although archaeological evidence in support of such claims is questionable, and in some cases, the validity of the supporting evidence has been challenged [ 21 ].

Cross-cultural studies suggest that MHP is widespread among ethnic groups, although the number of societies studied in details for this trait is rather limited e.

However, a number of recent reports of the absence of homosexual behavior and thus of MHP in some ethnic groups [ 24 ] has questioned the idea that MHP is virtually universal. This variation in the presence and absence of MHP among ethnic groups remains to be documented. From an evolutionary point of view, the emergence and maintenance of homosexual preferences require that the decrease in fertility associated with MHP should be compensated by sufficient increases in fertility among close relatives.

This increase may be promoted behaviorally by kin selection [ 2526 ], although empirical evidence is not always consistent. In Western societies, no difference between men with a MHP and heterosexual men is observed in the desire to invest in nieces and nephews [ 2728 ]; in Samoa, it has been observed boys the Fa'afafine the third gender associated with a MHP invest more in their nieces and nephews than heterosexual men [ homo252930 ]. These conflicting sources of evidence demand further research.

Alternatively, the increase in fertility in a close relative could be the result of an antagonistic factor. A sexually antagonistic gene that favors MHP in males and that increases fecundity in females has been proposed [ 31 ]. Several studies support this hypothesis [ 42231 — 34 ] and other have provided results sexual are consistent with predictions from this hypothesis [ 2234 — 38 ].

However why such an effect would boys operate also in wild animals is unclear. Sexually antagonistic genes are either fixed when the advantage is higher than the cost or selected against when the cost is higher than the advantage. When the frequency of a sexually antagonistic gene increases, selection to decrease the cost could eventually operate for example through the selection of a modifier genethus decreasing the fixation time of the antagonistic gene.

In any case, such sexually antagonistic genes are only transiently observed in natural population, perhaps explaining the absence—so far—of reports of homosexual preference in wild animals.

A recent change in social conditions could change the relative fitness advantage and cost of such gene, thus enhancing its selection. It has been recently proposed that selection for such sexually antagonistic genes could be promoted in social contexts specific to some human societies, where there is social stratification and hypergyny i.

Indeed in a stratified society, populations are organized into different groups or classes in which people share similar socioeconomic conditions. These groups can be ranked hierarchically depending on their access to resources with more resources for the top class. This social inequality also affects the expected reproductive success of each group with higher reproductive success associated with the top class [ 40 — 45 ].

This hypergyny hypothesis posits that females carrying the sexual antagonistic variant associated with MHP in males will signal increased levels of fertility through higher femininity or attractivenessthereby increasing their probability of reproducing in a wealthier social environment.

Such a sexually antagonistic gene will then provide a direct advantage by increasing fertility and an indirect advantage by increasing the probability of marrying into higher social classes. Such a process may promote MHP in stratified societies, and indeed, a comparative analysis suggests that social stratification level is a predictor of the presence of MHP in a society [ 39 ].

Several potential confounding variables were considered in this analysis, with the conclusion that none of them significantly influenced the probability of report of homosexual preferences. These variables included population density a good proxy of the number of indigenous people met by the anthropologistgeographical location and presence of moralizing gods.

However, this comparative study considered only 48 societies, and phylogenetic dependence among them Galton's problem [ 46 ] was not clearly addressed. While the questions of where, when and why MHP were usually considered separately, here we argue that it is particularly important to address these questions all together with an evolutionary perspective. Indeed, the sexual needed to answer each question sheds light on the others.

We will review the archaeological literature usually cited as evidence of MHP, and analyze the distribution of MHP among current human populations.

A comparative analysis on a large number of societies will then be performed to test the hypergyny hypothesis, while correcting for the phylogenetic relationships among the human societies. Archaeological data that have been repeatedly cited as evidence for the existence of MHP sexual prehistoric and early historic times were gathered from scientific papers [ 47 ] and books [ 161948 ] Table 1.

Data originating from unpublished sources such as media reports were not considered. A specialist of the post-paleolithic parietal Levantine art in Spain, A. Grimal Navarro [ 49 ] was contacted concerning statements from paintings from one Spanish cave, and his homo were cited as personal communication.

Data on the presence or absence of MHP in different societies have been gathered using existing reviews [ 650 — 52 ] and additional anthropological monographs and studies [ 2553 — 55 ]. The eHRAF allows to browse the original monographs. Relevant excerpts of the monographs concerning homosexuality have been extracted for each society. Positive clues of the presence of MHP in a society included a description from an anthropologist of individuals displaying a MHP, the existence of a word for MHP, and a description of a third gender including individuals displaying a MHP such as the Fa'afafine of Samoa [ 25 ] or the Berdache of North America [ 56 ].

Negative clues sexual the absence of a word and concept for MHP or the direct conclusion from an anthropologist after having explicitly asked for the existence of homosexuality. When a clear distinction between MHP and homosexual behavior could not be made, the case was not considered further. Two independent measures of the sexual of social stratification were gathered to control for the dependence of model sensitivity on the way in which social stratification was rated.

The five factors of this variable have been merged into three levels in order to suppress empty classes not supported by the statistical method used here, i. The resulting factors were: 1 absence of social stratification factor 12 simple stratification based on wealth or elite boys factors 2 and 3 and 3 complex stratification merging factors 4 and 5.

Second, data on the level of social stratification were gathered using eHRAF, relevant anthropological monographs and books [ 53 — 5559 — 64 ]. From the excerpts, boys level of stratification was first assessed for each society on a scale ranging from 1 to 5, corresponding to the number of classes that could be identified, and then reduced to 3 levels see S1 Text : 1 no stratification, 2 moderately stratified, and 3 strongly stratified.

Generalized linear models were used to test the influence of the level of stratification on the probability of observing MHP coded as 0 or 1. To take into account the non-independence among societies, generalized estimating equations GEE were used. GEE allow the relationship between a response variable and explanatory variables in a generalized linear model framework to be analyzed by taking into account a structure of correlation between the items of the response variable [ 65 ].

As the sexual ancestral relationships among all of the sampled societies were not available, two proxies of the true phylogeny were used in the analysis. First, a linguistic phylogeny, known to parallels boys trees [ 6667 ].

Second, the geodesic distance between societies was used as a proxy of their cultural and historical closeness, with the implicit assumption that geographic distances are related to cultural similarities. A linguistic similarity matrix between societies was extracted from the World language tree of lexical similarity from version 15 of the Automated Similarity Judgement Program database [ 69 — 72 ]. This linguistic phylogeny is based on the 40 more stable words of a Swadesh list [ 7374 ].

The phylogenetic trees containing only the societies for which data on MHP and stratification were available were extracted from the World language tree of lexical similarity and were used for an additional statistical analysis.

The analysis using the linguistic phylogeny to control for homo was performed with sexual of the two distinct measures of the level of social stratification one extracted from the EA, and one extracted from eHRAF and complementary sources, see above. As the results did not differ qualitatively, further analyses were performed using the largest sample i. Then, boys localization of each society was homo from their latitude and longitude as coded in the Ethnographic Atlas.

The great-circle distance between each society was then calculated. The resulting matrix of distances between societies was used to build a tree of distance based on the neighbor-joining method [ 75 ], and used in additional statistical analyses. This tree was integrated in an analysis with the presence and absence of MHP as a response variable and social stratification as an explanatory variable. To compare these results with a previous analysis [ 39 ], a classical generalized linear model with binomial error was performed, using the geographic zone as a confounding variable in that study, density of population had no significant effect and was thus not considered here.

Presence and absence of MHP was the response variable, the level of stratification homo an explanatory variable, and the geographic zone was a possible confounding variable. All analyses were conducted using R version 2. Several prehistoric references were examined.

The first one corresponds to Mesolithic paintings, the panel of the Cueva de la Vieja Albacete, Spain belonging to the Spanish Levantine art. However, a homo distinct interpretation is provided by a specialist of homo cave, and more generally of the Levantine art in Spain.

Furthermorethe color and craftsmanship of this character is distincthe also holds a bow. More generally, G. Grimal Navarro, personal communication, ; our own translation. The second refers to a petroglyph of Bardal, Norway, dating back to the Mesolithic period.

When we inspected the depiction of the petroglyphs from the original reports [ 80 — 82 ], we identified several problems due to the variability in drawings. In some, the identification of the smaller individual as a human is questionable, as two lines above the head suggest a horned animal, consistent with hair depicted below the belly. In others, lines suggesting a rear-entry penetration are not reported.

This petroglyph could thus represent either homosexual intercourse, heterosexual intercourse, zoophilia, or something else other references are analyzed in the supporting information section: Archaeological evidences. Our aim was to test the link between the presence and absence of MHP response variable and the level of social stratification explanatory variable.

However, the type of violence experienced by young homosexuals differs depending on whether they are male or female, and has results that can affect their identity and intersubjective relations.

Male adolescents mentioned episodes of homophobic violence, while girls were victims of gender-based violence. Twelve of the 14 interviewees mentioned suicidal ideation or attempted suicide and two of the boys who sought STD treatment were HIV-positive. With regard to sexual orientation, only With regard to health care, none of the interviewees had been previously asked by health professionals about homosexual experiences or had received guidance on the issue.

However, none of the adolescents mentioned that they had been discriminated due to their sexual orientation. Apparently health professionals reproduce the "heteronormative" view common to almost all cultures 14 , acting as if everyone was heterosexual. For some, the homosexual experience depended on particular circumstances based on curiosity-driven experimentation, while for others it was associated with prostitution. However, for the majority, the experience was a matter of self-declared sexual orientation.

Casual sexual activity and experimentation were mentioned as random occurrences in which the adolescents found themselves involuntarily involved in situations that culminated in homosexual contact.

These adolescents showed a feeling of guilt regarding these encounters, despite the fact that such sexual experiences are part of the development of sexuality and did not define their sexual orientation. This feeling is probably influenced by homophobia, which makes them reject the possibility of considering themselves homosexual or bisexual.

The sociocultural context governed by compulsory heteronormativity, a social norm in which heterosexuality is compulsory, constructs a necessary and coherent relationship between gender identity, desires and sexual behavior. Heteronormativity defines the social conventions that determine gender and sexuality, marked by an asymmetry between male and female 15 where those who do not fit the norm feel inadequate. All I know is that it was impulsive.

We were all the same age. It began as an exchange of affection, like one of their games and, before we knew it, we were having a relation".

And, I just let it happen. Casual homosexual experiences during childhood and adolescence are part of the construction of sexual identity, which only acquires its final form generally at the end of adolescence.

Comprehensive research about reproduction and the social trajectories of youth in Brazil conducted by Heilborn and Cols identified various forms of desire that go against heteronormative values Freud, in his essays on sexuality, published for the first time in , raises an important aspect about the understanding of sexual object choice, which presupposes an "original" bisexuality in human beings and, furthermore, that it should be considered that adult sexual posture requires fluctuation and learning from both parts, considering the complexity and various possible subjective arrangements.

Another meaning assigned to the homosexual experiences of the boys relates to prostitution, which was observed in the accounts of both the adolescents that considered themselves heterosexual and those that were openly homosexual. In both cases, the justification for this practice was financial gain. They mentioned being harassed by older men and were bitter about the hardship they have to suffer due to lack of money. We noted that the adolescents apparently consider it natural to engage in homosexual practices through prostitution, masking its inherent violence, not only in structural terms, but also the violence that results from the unequal power relationship between the perpetrator and the adolescent prostitute.

The behavior of these adolescents was forged through relations and by meanings of sexuality constructed throughout history and at different social levels that encompass macro-social and socio-historic issues and the specific aspects of interpersonal relationships. The subjectivity of these individuals and their conduct is strongly influenced by social determinants The third meaning of homosexual experience mentioned by the majority of the male interviewees was associated with self-declared homosexual identity.

The first homosexual experience of these adolescents was generally at a young age, during childhood or preadolescence and sometimes with partners who were a lot older. The male imperative makes having sexual relations practically compulsory, replicating the standard behavior of heterosexual men and showing that the affective and sexual trajectory of youth appears to be strongly structured by gender patterns.

Furthermore, the vulnerability of these self-declared gay adolescents is aggravated by the perception of the masculinity of the other as a factor of attraction, which appears to represent an increased risk of contracting HIV, as shown by a study conducted with YMSM in the United States With respect to the construction of the homosexual identity, a large part of the interviewees had sexual relations with women, expressing the need to test their desires and be certain of their sexual orientation.

Guilt related to internalized homophobia, which contributes to low self-esteem, was also evident Sexual orientation is defined during adolescence. It is beyond the scope of the present article to attempt to describe the complexities of the identification process. Suffice to say that it is lived by the subject as transgression, provoking feelings of strangeness when an individual becomes aware of homosexual desires. People are born that way, with it inside them. I repressed myself a lot before, you know?

I was unhappy. There was that thing, you know, [I thought], is this abnormal"? I have always seen myself as man-woman. I felt really alone when I was little. I was always slightly effeminate. Today, I accept myself more. You know, an ET in the middle of a crowd, who no one recognizes". In a study conducted with homosexual men, Nunan 22 reports that all interviewees said that they were born homosexual; none of the interviewees stated that they had chosen to be homosexual. The first sexual relation of the majority of those interviewed by Nunan was with a woman, as a way of experimenting and being certain about their sexual orientation.

With respect to the sexual development of the adolescents interviewed by the present study, when telling us about their sexual history, some, paradoxically, mentioned their childhood experiences and associated being gay with the fact that they had been abused.

They used a past marked by violence to justify the present, as the following extract shows:. I ended up liking it. This influenced my path, definitely. I did it with others and became more and more accustomed to it".

Homophobic violence has an impact on the life of these adolescents and limits their access to social protection, schooling and employment, resulting in lack of economic resources and leading to social marginalization, which often results in the need to resort to prostitution Adolescent homosexuals are often pressured to hide their sexual identity as a mechanism to avoid rejection and hostility, aggravating the usual adolescent concerns during this phase of development and often causing problems Homophobia contributes to the adoption of high-risk sexual practices and can mean that adolescent gays isolate themselves 24 because they do not have the courage to reveal their sexual orientation 25 , 26 , only manifesting their sexual preferences to a limited few.

The health system is organized around the needs of the heterosexual population. Health professionals almost always assume that an adolescent is heterosexual, which makes it more difficult for the individual to reveal their real identity.

Therefore, homosexual activity lacks legitimacy in society, which often forces homosexuals behind closed doors, thus reinforcing the moral degradation discourse and destroying self-esteem, hindering the adoption of safe sex practices, and often leading to alcohol and drug use The adolescents interviewed by this study told of degrading treatment by the family when they revealed their sexual orientation such as humiliation and emotional abuse, and beatings, which in some cases led to fractures and permanent scares.

Many gays under the age of 18 are thrown out of home and, having nowhere to live and no one to turn to, go into prostitution as male prostitutes or transvestites. Many, unable to stand the violence find that suicide is the only solution to free themselves from this suffering The main meaning assigned by the female interviewees to the homosexual experience related to the love they feel for their partners.

The affectional bond predominates as the following extract shows:. We show the love we feel for each other, you know? Sex is like, normal.

But, like, I lie down in bed with her, have sex with her, I have sex out of love, not out of pleasure". Our findings are corroborated by the literature that shows that female homosexuality is quite different from male homosexuality when it comes to partners.

For women, sex is associated with affection. The same cannot be said for men, who tend to have various sexual relations in the same day with strangers just for pleasure Loyola 29 highlights that sexual pleasure for women is a slow process constructed with love.

On the same line, Palma 30 suggests that female couples are characterized by intense companionship, with a strong emphasis on mutual emotional support. Sexuality does not have the same emphasis among lesbians that it does in gay men, since the choice of partners, even casual ones, is not made based on sexual attraction, but on love. This behavior follows the norms of a predominantly patriarchal and heteronormative society.

Patterns of behavior, values, and gender norms throughout history are manifested and assimilated as natural, and are reflected in the affective and sexual relations and health of the subjects. Men and women are instigated to take on different roles, where love and affection is the prerogative of the female sex.

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