Materteral and Avuncular Tendencies in Samoa

Androphilia refers to sexual attraction and arousal to adult males, whereas gynephilia refers to sexual attraction and arousal to adult females. In Independent Samoa, androphilic males, most of whom are effeminate or transgendered, are referred to as fa’afafine, which means “in the manner of a woman.” Previous research has established that fa’afafine report significantly higher avuncular tendencies relative to gynephilic men. We hypothesized that Samoan fa’afafine might adopt feminine gender role orientations with respect to childcare activity. If so, then the fa’afafine’s femininity might be a proximate mechanism for promoting their elevated avuncular tendencies. Our analyses indicated that fa’afafine had significantly higher willingness to assist in the childcare of nieces and nephews than childless women, mothers, or men, none of whom differed from each other on this measure. Thus, femininity does not appear to explain the fa’afafine’s pattern of avuncular tendencies, nor the women’s pattern of materteral (i.e., aunt-like) tendencies, relative to gynephilic men. We discuss how the fa’afafine “third” gender status might influence the expression of their elevated avuncular tendencies.

Human handedness and scalp hair-whorl direction develop from a common genetic mechanism

Theories concerning the cause of right- or left-hand preference in humans vary from purely learned behavior, to solely genetics, to a combination of the two mechanisms. The cause of handedness and its relation to the biologically specified scalp hair-whorl rotation is determined here. The general public, consisting of mostly right-handers (RH), shows counterclockwise whorl rotation infrequently in 8.4% of individuals. Interestingly, non-right-handers (NRH, i.e., left-handers and ambidextrous) display a random mixture of clockwise and counterclockwise swirling patterns. Confirming this finding, in another independent sample of individuals chosen because of their counterclockwise rotation, one-half of them are NRH. These findings of coupling in RH and uncoupling in NRH unequivocally establish that these traits develop from a common genetic mechanism. Another result concerning handedness of the progeny of discordant monozygotic twins suggests that lefties are one gene apart from righties. Together, these results suggest (1) that a single gene controls handedness, whorl orientation, and twin concordance and discordance and (2) that neuronal and visceral (internal organs) forms of bilateral asymmetry are coded by separate sets of genetic pathways. The sociological impact of the study is discussed.     

Testing multiple hypotheses for the maintenance of male homosexual copulatory behaviour in flour beetles

Diverse animal groups exhibit homosexual interactions, yet the evolutionary maintenance of such behaviours remains enigmatic as they do not directly increase reproductive success by generating progeny. Here, we use Tribolium castaneum flour beetles, which exhibit frequent male homosexual copulations, to empirically test several hypotheses for the maintenance of such behaviours: (1) establishing social dominance; (2) practice for future heterosexual encounters; and (3) indirect sperm translocation. We found no evidence that Tribolium males use homosexual copulations either to establish dominance or to practice behaviours that increase their subsequent heterosexual reproductive performance. Our results provide limited support for the hypothesis of indirect sperm translocation: when males from two genetic strains mated with females immediately following a homosexual copulation, females produced progeny sired not only by the directly mating male, but also by that male’s homosexual partner. However, this phenomenon was detected in only 7% of homosexual pairs, and in each case such indirectly sired progeny accounted for < 0.5% of females’ total progeny. Direct observations indicated that mounting males often released spermatophores during homosexual copulations. These observations suggest that homosexual copulations may be a behavioural mechanism that allows males to expel older, potentially low‐quality sperm. Additional work is needed to test this new hypothesis, and to determine whether sperm release during homosexual copulations occurs in other groups.     

Extreme skewing of X chromosome inactivation in mothers of homosexual men

Human sexual preference is a sexually dimorphic trait with a substantial genetic component. Linkage of male sexual orientation to markers on the X chromosome has been reported in some families. Here, we measured X chromosome inactivation ratios in 97 mothers of homosexual men and 103 age-matched control women without gay sons. The number of women with extreme skewing of X-inactivation was significantly higher in mothers of gay men (13/97=13%) compared to controls (4/103=4%) and increased in mothers with two or more gay sons (10/44=23%). Our findings support a role for the X chromosome in regulating sexual orientation in a subgroup of gay men.     

Variation in the peacock’s train shows a genetic component

Female peafowl (Pavo cristatus) show a strong mating preference for males with elaborate trains. This, however, poses something of a paradox because intense directional selection should erode genetic variation in the males’ trains, so that females will no longer benefit by discriminating among males on the basis of these traits. This situation is known as the ‘lek paradox’, and leads to the theoretical expectation of low heritability in the peacock’s train. We used two independent breeding experiments, involving a total of 42 sires and 86 of their male offspring, to estimate the narrow sense heritabilities of male ornaments and other morphometric traits. Contrary to expectation, we found significant levels of heritability in a trait known to be used by females during mate choice (train length), while no significant heritabilities were evident for other, non-fitness related morphological traits (tarsus length, body weight or spur length). This study adds to the building body of evidence that high levels of additive genetic variance can exist in secondary sexual traits under directional selection, but further emphasizes the main problem of what maintains this variation.     

A Juvenile Hormone analogue enhances homosexual behaviour in female‐deprived males of the migratory locust

In laboratory colonies of crowded migratory locusts Locusta migratoria (L.), homosexual behaviour (i.e. males mounting other males) is commonly observed. Female‐deprived males of Locusta migratoria migratorioides R. & F. mount each other in a characteristic mating position, often forming a group of several insects. When allatectomized males are placed together with intact, female‐deprived males, the former are usually mounted by the latter, demonstrating some degree of control involving the corpora allata over homosexual behaviour. This may be related to the positive effects of Juvenile Hormone (JH) or Juvenile Hormone analogues (JHAs) in enhancing male sexual behaviour, as is shown in some other insects. In the present study, a potent JHA (i.e. pyriproxyfen) is injected into a group of young, crowded L. m. migratorioides males, and an equivalent group of control males is maintained in a separate cage. Both groups are deprived of females. Three times a week, during 2‐h observation periods, JHA‐injected and control males are placed together in the absence of females, and homosexual mountings, per group and per individual, are recorded every 10 min. Observations are performed for 10 weeks. Analysis of these data, including the time spent in mounting behaviour, the percentage of individuals within a group involved in the behaviour during the observation periods and the identity of both partners, reveals that the JHA‐treated males show a more intense homosexual mating behaviour than control males in all quantified parameters. This is the first report of the enhancement of homosexual behaviour by an endocrine factor in insects.     

Women’s fertility across the cycle increases the short-term attractiveness of creative intelligence

Male provisioning ability may have evolved as a “good dad” indicator through sexual selection, whereas male creativity may have evolved partly as a “good genes” indicator. If so, women near peak fertility (midcycle) should prefer creativity over wealth, especially in short-term mating. Forty-one normally cycling women read vignettes describing creative but poor men vs. uncreative but rich men. Women’s estimated fertility predicted their short-term (but not long-term) preference for creativity over wealth, in both their desirability ratings of individual men (r=.40, p<.01) and their forced-choice decisions between men (r=.46, p<.01). These preliminary results are consistent with the view that creativity evolved at least partly as a good genes indicator through mate choice.     

Courtship behaviour in Japanese macaques during heterosexual and homosexual consortships

Female Japanese macaques (Macaca fuscata) in the Arashiyama population near Kyoto, Japan, are unusual, in that they exhibit what many would consider to be male-typical sexual characteristics. Specifically, they mount other females within the context of temporary, but exclusive, sexual relationships (i.e., homosexual consortships) and they sometimes exhibit a preference for female sexual partners, even when given the choice of a sexually motivated male alternative. In this study, we examined whether female Japanese macaques also exhibited male-typical patterns of courtship behaviour during homosexual consortships. Data were collected on courtship behaviour during heterosexual and homosexual consortships in free-ranging Japanese macaques from the Arashiyama (Japan) population. We analyzed the occurrence of 12 different courtship behaviours during 3374 heterosexual inter-mount intervals and 1412 homosexual inter-mount intervals. Sex differences between heterosexually consorting males and females existed for only two of the 12 courtship behaviours we investigated: inclined-back presentations and sexual vocalizations. Dominant and subordinate homosexually consorting females were sex-typical in their expression of inclined-back presentations and sexual vocalizations. Consequently, facultative same-sex sexual partner preference, mounting and consortships do not co-occur with male-typical patterns of courtship behaviour in female Japanese macaques.     

Patterns of infant handling and relatedness in Barbary macaques (<Emphasis Type="Italic">Macaca sylvanus</Emphasis>) on Gibraltar

Among papionin primates, the Barbary macaque (Macaca sylvanus) shows the most extensive interactions between infants and group members other than the mother. Two different types of interactions occur: (1) long-lasting dyadic interactions between a handler and an infant, and (2) brief triadic interactions between two handlers involving an infant. Previous investigations showed that infant handling by males is best explained as use of infants to manage relationships with other males. In contrast, no adaptive explanation for infant handling by females emerged. Here, we compared the infant-handling pattern between subadult/adult males and subadult/adult females in a free-ranging group of 46 Barbary macaques on Gibraltar to test whether the relationship management hypothesis also applies to female handlers. We further investigated the infant-handling pattern of juveniles and used microsatellite markers to estimate relatedness between infant handlers and the infant’s mother. We found that males, females and juveniles all participated extensively in triadic interactions using infants of above-average related females. In contrast, only males and juveniles were highly involved in dyadic interactions with infants of related females, while females rarely handled infants other than their own. The pattern of infant handling was entirely compatible with the predictions of the relationship management hypothesis for males and mostly so for females. Moreover, our genetic analysis revealed that males and females differ in their partner choice: while females preferred to interact with related females, males had no significant preference to interact with related males. We further discuss the observed above-average relatedness values between infant handlers and the infant’s mother in the light of kin-selection theory.     

Male mate choice in the gynogenetic–sexual complex of crucian carp,<Emphasis Type="Italic"> Carassius auratus</Emphasis>

The asexual all-female Japanese crucian carp, Carassius auratus langsdorfii (Teleostei: Cypriniformes), reproduces gynogenetically, relying on the sperm of males of the sexual "host," C. auratus subspp. Theoretically, frequency-dependent mating preference of males to conspecific females can lead to the coexistence of asexual and sexual fish, if all else is equal. Our specific questions are whether males prefer conspecific females over asexual females and whether individuals show dominance hierarchies that potentially cause frequency-dependent mating preference. In an individual choice experiment, a tank was partitioned into three compartments with the middle one for a single male and the two outer ones for a sexual and an asexual female. The males of C. auratus bürgeri demonstrated a significant preference for ovulated conspecific females over ovulated asexual females. In contrast, in a group mating experiment, a single experimental tank included two males, a sexual female, and an asexual female together, and males chased and mated with both asexual and sexual females equally. Male mate preference was weak in group mating, which is typical in natural populations. Males and females of crucian carp showed no apparent agonistic behavior to each other in the group mating experiment. This is different from other gynogenetic complexes with the dominance hierarchy of males showing strong frequency-dependent mating preference (e.g., Poeciliopsis). We conclude that male mate preference is unlikely to be a strong frequency-dependent force maintaining the coexistence of asexual–sexual complexes of Japanese crucian carp. Received in revised form: 5 February 2001 Electronic Publication     

Do male mealworm beetles, <Emphasis Type="Italic">Tenebrio molitor</Emphasis>, sustain the honesty of pheromone signals under immune challenge?

Immune function is potentially costly and traded-off against the expression of sexual signals, thus, making sexual advertisement a condition-dependent and honest indicator of current immunocompetence. We conducted a series of laboratory experiments using mature mealworm beetles, Tenebrio molitor, to examine whether an immunological challenge (nylon implant(s), microbial lipopolysacharides, microlatex beads) could affect the attractiveness of male pheromones or male’s survival in ad libitum and ‘water-only’ food regimes or whether pheromones signal more long-term qualities of males. We treated another set of newly-emerged males with a single nylon implant and gave first ad libitum food but later only water. We used several immune challengers and feeding regimes to examine if the expected trade-off between immune function and sexual signaling would be expressed differently depending on the resource level and the type of the immune challenge. None of the treatments affected the female preference for male pheromones or the male longevity except for the acute mortality caused by two-implant treatments. However, males lost less body mass when immune challenged and given an opportunity to feed. We conclude that females seem to prefer certain males, and the effect of immune challenge in males cannot systematically override these preferences.     

Baboon sexual swellings and male response: A possible operational mammalian supernormal stimulus and response interaction

Experimental work using coagulated masturbatory seminal emissions from nine adult male chacma baboons (Papio ursinus)as a dependent variable revealed that males show a greater sexual arousal response to female conspecifics exhibiting exaggerated perineal swellings. The menstrual cycles of the “superswollen” females did not differ in length or patterning from the controls. Since additional experimental work, including behavioral observations on females with normal and supersized swellings, failed to reveal differences in female proceptivity, the male’s increased arousal response appears dependent upon the perineal size per se.The males’ responses suggest that the supersized swellings may be an example of an operational visual supernormal stimulus and response interaction.     

Dispersal of the alien raccoon dog Nyctereutes procyonoides in Southern Brandenburg, Germany

The raccoon dog Nyctereutes procyonoides, an East Asian canid species, was introduced into the European part of the former USSR since 1928. Within 50 years (1935–1984), it colonised a territory of 1.4 million km² in Europe. A telemetry study took place in Southern Brandenburg in a 60 km² sized study area with a typical mosaic structured East German agricultural landscape. For catching raccoon dogs, 20 trap boxes were set there in an area of 46 km², and between February 2001 and July 2004, 15 (5 males, 10 females) adult and 46 (25 males, 21 females) juvenile raccoon dogs were eartagged and adults additionally fitted with radio collars (Biotrack, 150–151 MHz). Data on dispersal behaviour was collected by the relocation points of 11 juveniles (6 males, 5 females). Four juvenile males dispersed even more than 40 km from their trapping places. Additionally, dispersal of two adult males could be documented. This behaviour probably indicates that the German raccoon dog population still is in a process of colonising. This canid’s ability for colonising spacious and distant areas during comparative short periods of time and its preference for habitats with richness of water possibly make this species to be an important vector of fox tapeworm Echinococcus multilocularis—a very dangerous zoonosis.     

The ratio of 2nd to 4th digit length and male homosexuality

Sexual orientation may be influenced by prenatal levels of testosterone and oestrogen. There is evidence that the ratio of the length of 2nd and 4th digits (2D:4D) is negatively related to prenatal testosterone and positively to oestrogen. We report that (a) 2D:4D was lower in a sample of 88 homosexual men than in 88 sex- and age-matched controls recruited without regard to sexual orientation, (b) within the homosexual sample, there was a significant positive relationship between mean 2D:4D ratio and exclusive homosexuality, (c) overall, there was a decrease in 2D:4D from controls to homosexual men to bisexual men and (d) fraternal birth order, a positive predictor of male homosexuality, was not associated with 2D:4D in a sample of 240 Caucasian men recruited without regard to sexual orientation and 45 homosexual men.Further work is needed to confirm the relationships between 2D:4D and sexual orientation. However, these and other recent data tend to support an association between male homosexuality and high fetal testosterone. Very high testosterone levels may be associated with a sexual preference for both men and women.     

Male barn swallows use different signalling rules to produce ornamental tail feathers

The evolution of secondary sexual characters is the subject of controversial debate between those defending their role as ‘viability indicators’ and those arguing that ornaments are purely ‘attractive traits’ selected by females. Recent theoretical studies suggest that these hypotheses are not mutually exclusive, as both viability and attractiveness can contribute to improve the reproductive success of progeny and could thus simultaneously underlie female choices. If that is the case, strategies of cheaper advertisement, allowing the expression of larger ornaments for the same cost, could proliferate even in species in which honest signalling of viability prevails. Under this scenario, different males could invest a different amount of resources per ornament unit of expression, thus using different signalling rules. We studied the relationship between tail feather length (a trait that is the subject of a female mate preference) and feather mass (a measure of investment in feather production) in a barn swallow Hirundo rustica population. Different males used different and consistent signalling rules when developing ornamental feathers. That is, to produce a feather of a given length, each male used a constant amount of resources across different years, but this amount varied between males. Although the amount of material invested in feathers (feather mass) is a condition-dependent trait, the organization of this material in ornamental feathers (i.e. the signalling rules) was not. Neither survival nor risk of feather breakage was related to the signalling rules. Thus, these results suggest that both ‘viability’ and ‘runaway’ mechanisms are independent determinants of the evolution of ornamental sexual feathers in the barn swallow. A preference for long tails will ensure that females either obtain a sire with high viability, or one transferring the capability to produce longer and more attractive tails at a lower cost of production to its offspring.     

Female mate preferences in Drosophila simulans: evolution and costs

Female mate preference is central to sexual selection, and all indirect benefit models require that there is genetic variation in female preference. This has rarely been tested however, with relatively few studies documenting heritable variation in female preference and even fewer that have directly selected on mate preference to unequivocally show that it can evolve. Additionally, costs of mate preference are poorly understood even though these have implications for preference evolution. We selected on female preference for ebony‐males in replicate Drosophila simulans lines, and generated a rapid evolutionary response in both replicates, with the proportion of females mating with ebony‐males increasing from approximately 5% to 30% after five generations of selection. This increase was independent of changes in ebony‐males as only females were included in our selection regime. We could detect no cost to mate preference itself other than that associated with the fitness consequences of mating with ebony males.     

Comparative response to citrus foliage and citrus fruit odour by wild and mass-reared sterile Mediterranean fruit fly males of a genetic sexing strain

In greenhouse behavioural assessments conducted at Seibersdorf, Austria that allowed flies the choice of different types of potted host trees and plants, it was found that mature laboratory mass-reared Mediterranean fruit fly males, Ceratitis capitata (Wied), (Dipt. Tephritidae), strain Vienna-42 (temperature-sensitive lethal genetic sexing strain) showed the same preference for orange foliage, over other plants types and parts, as wild males. They aggregated in leks, released pheromone, and courted approaching females almost exclusively on orange foliage. Even though a number of other trees and plants were available to Vienna-42 males, less than 1% of these sexual activities occurred on the foliage or other plant parts of non-Citrus trees and not a single mating occurred there. In choice tests carried out in Chios, Greece in large field cages housing naturally planted orange trees, it was determined that mature and sterile Mediterranean fruit fly males of the same sexing strain responded to the odour emanating from the flavedo (due to superficial cuts made in the fruit peel containing the oil glands) of ripening sweet, sour oranges and lemons, in a similar intensity, order of fruit preference and time of day as did mature males of the local wild population. Males of both strains were frequently observed contacting with their mouth parts the oils exuding from the cuts made in the flavedo of the orange peel, or exhibiting homosexual activity on, or near the cuts. The results show that mass-reared, sterile males of the Vienna-42, genetic sexing strain behave in a comparable way to wild males in an important component of habitat location (odour of host foliage and host fruit) and hence are competitive with them in these respects. None of the standard international quality control tests for sterile Mediterranean fruit flies evaluates this important habitat location behaviour, which is a prerequisite for the successful encounter of the sexes in a sterile release programme.     

Male dominance and genetically determined reproductive success in the mandrill (Mandrillus sphinx)

Darwin referred to the adult male mandrill (Mandrillus sphinx) as the most brightly coloured of all mammals, citing the brilliant red and blue pigmentation of the face, rump, and genitalia as extreme examples of evolution by sexual selection. Considerable controversy exists concerning possible effects of sexually selected phenotypes via intermale competition on reproductive success. Behavioural and genetic studies of a large, semi-free ranging mandrill colony in Gabon have now demonstrated that clear-cut relationships exist between male secondary sexual development, social dominance, copulatory behaviour, and reproductive success in this primate species. Two morphological variants of adult male were identified; “fatted” males, with maximum secondary sexual coloration, which occupied dominant positions in the social group, and “non-fatted” males, with muted secondary sexual adornments, smaller testes and lower plasma testosterone levels, which lived as peripheral/solitary individuals. DNA fingerprinting analyses on infants born over five successive years showed that only the two most dominant, fatted males in the group had fathered offspring. Throughout the annual mating season these males attempted to mate-guard and copulate with females during periods of maximal sexual skin tumescence. Male rank and mating success were strongly positively related and the alpha male sired 80 – 100% of the resulting offspring during three consecutive years. Non-fatted adult males and group associated subadult males engaged in infrequent, opportunistic matings and did not guard females. Loss of alpha status resulted in a fall in reproductive success, but the effect was gradual; the deposed alpha male continued to father 67% and 25% of infants born during the next two years. Thus, whilst claims that male dominance determines mating success and paternity in primates have caused considerable debate, these results on mandrills provide unequivocal evidence for the existence of such effects.     

Breeding-season relationships of immature male rhesus monkeys with females. I. Individual differences and constraints on partner choice

Several aspects of breeding-season relationships with mature females are described for free-ranging immature male rhesus monkeys (Macaca mulatta)on Cayo Santiago. At puberty, immature males are still groomed by mature females, as they were as infants, but no longer receive active protection from females other than their mothers. A further, potentially beneficial, aspect of their associations with females lies in the opportunities to observe closely the tactical social and sexual interactions of receptive females with adult males. However, immature males themselves rarely copulate with females. Developmental constraints may explain the finding that most females observed by immature males were relatives but that high-ranking males showed a particular preference for adolescent relatives, whereas the preference of low-ranking males was for adult relatives. A further finding was that while there was a tendency for immature males to devote a greater proportion of their total observation time to females that ranked above their mothers than to those of inferior rank, these were also the females from which males received the most aggression, while most of the females with which the males groomed or copulated were lower ranking than the males’ mothers. Of three hypotheses concerning three possible types of social constraint which might account for such a finding, the data supported only one—that particular levels of aggression within these relationships are compatible only with certain other elements (observations), and not with others (grooming, copulation).     

Water temperature and pH influence olfactory sensitivity to pre-ovulatory and post-ovulatory ovarian pheromones in male Barilius bendelisis

The attractive response and sexual activity elicited by pre-ovulatory steroid sulphate and post-ovulatory 15K-PGF pheromones are greater in wild caught tubercular males and immature males which express breeding tubercles on the snout (at 12–13 days post androgen implant) than in non-tubercular and non-androgen implanted males of freshwater fishBarilius bendelisis. This shows that circulatory androgens exert an activational effect on olfactory receptors of male fish. Wild caught tubercular males and androgen implanted juvenile males exhibit a high responsiveness to steroid sulphate at the water temperature and pH which fish experience during the pre-spawning phase. The male’s sensitivity to 15K-PGF is almost equally high at the water temperature and pH which they experience in wild during the both pre-spawning and spawning periods. This suggests that the differential olfactory sensitivity to the two classes of pheromones in androgen implanted males is due to the varied temperature and pH of water, and that during the breeding season the male’s olfactory sensitivity to PGF pheromone is more widespread than to the steroidal pheromone. An increased and decreased olfactory sensitivity in mature males to sex pheromones and L-alanine respectively during the breeding phase is in agreement with the hypothesis that pheromonal stimuli dominate over feeding stimuli to promote spawning success.