Sex differences in sexual partner acquisition, retention, and harassment during female homosexual consortships in Japanese macaques

Female Japanese macaques (Macaca fuscata) in certain populations are unusual in that they exhibit male-typical patterns of mounting behavior and sexual-partner preference. The goal of this study was to determine whether female Japanese macaques, from one such population, employ male-typical behavioral tactics to disrupt existing homosexual consortships, as well as to acquire and retain same-sex sexual partners. "Harassment" of homosexual consortships occurred when a sexually motivated, third-party male or female interrupted a consorting female couple by displacing or aggressing them. Sexual harassment was a male-typical strategy for disrupting existing homosexual consortships, but was rarely performed by females. "Intrusions" occurred when a male or female competitor attempted to acquire exclusive access to a female engaged in a homosexual consortship by targeting that female as the focus of competition and her partner as his/her competitor. "Sexual coercion" occurred when one individual alternately sexually solicited and aggressed another individual as part of the same behavioral sequence during an intrusion. Males employed consortship intrusions and sexual coercion when they attempted to acquire female sexual partners that were already engaged in homosexual consortships, but females rarely did so. However, females did employ male-typical patterns of aggressive competition and sexual coercion to retain same-sex sexual partners when confronted with male competitors' attempts to usurp those partners. These results indicate that female sexual activity during homosexual consortships is not uniformly "masculine" in expression, but rather is a mixture of male- and female-typical behaviors.     

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.     

Mounting Interactions between Female Japanese Macaques: Testing the Influence of Dominance and Aggression

The goal of the work reported here was to determine whether female Japanese macaques (Macaca fuscata) participated in same-sex mounting interactions during homosexual consortships to communicate about asymmetries in their dominance relationships and to reduce aggression. Focal data were collected during 21 homosexual consortships involving 14 females living in a captive, mixed-sex group of 37 individuals. We identified eight types of mounts, one solicitation used specifically to request to mount (hands-on-hindquarters solicitation), two solicitations used specifically to request to be mounted (hindquarter and back presentations), and one behavior employed to facilitate mounts-in-progress (clasping). We tested whether dominant consort partners (1) mounted more and (2) requested to mount more than their subordinate partners and whether subordinate consort partners (1) requested to be mounted more and (2) facilitated mounts-in-progress more than their dominant partners. Finally, we examined whether mounting was temporally linked to the onset of aggressive interactions between consort partners and whether it functioned to defuse incipient aggression. None of these predictions was supported. All types of mounts, mount solicitations, and clasping occurred bi-directionally within consort dyads. Mutual sexual attraction and gratification provided the proximate motivation for these mounting interactions and, in turn, for the formation and maintenance of their homosexual consortships.     

Hatching sex ratio and sex specific chick mortality in common terns Sterna hirundo

Bias in sex ratios at hatching and sex specific post hatching mortality in size dimorphic species has been frequently detected, and is usually skewed towards the production and survival of the smaller sex. Since common terns Sterna hirundo show a limited sexual size dimorphism, with males being only about 1–6% larger than females in a few measurements, we would expect to find small or no differences in production and survival of sons and daughters. To test this prediction, we carried out a 2-year observational study on sex ratio variation in common terns at hatching and on sex specific post hatching mortality. Sons and daughters hatched from eggs of similar volume. Post hatching mortality was heavily influenced by hatching sequence. In addition, we detected a sex specific mortality bias towards sons. Overall, hatching sex ratio and sex specific mortality resulted in fledging sex ratios 8% biased towards females. Thus, other reasons than body size may be influencing the costs of rearing sons. Son mortality was not homogeneous between brood sizes, but greater for two-chick broods. Since adults rearing two-chick broods were younger, lighter and bred consistently later than those rearing three-chick broods, it is suggested that lower capacity of two-chick brood parents adversely affected offspring survival of sons. Though not significantly, two-chick broods tended to be female biased at hatching, perhaps to counteract the greater male-biased nestling mortality. Thus, population bias in secondary sex ratio is not limited to strongly size dimorphic species, but species with a slight sexual size dimorphism can also show sex ratio bias through a combination of differential production and mortality of sons and daughters.     

The reproductive ecology of resident manta rays (Manta alfredi) off Maui, Hawaii, with an emphasis on body size

In resident manta rays (Manta alfredi) off Maui, sexual maturity appears delayed until growth exceeds 90% of maximum size, an indicator that large body size provides a reproductive advantage at the expense of a shorter reproductive time period. In this study, 286 surveys were conducted between 2005 and 2010 using photo-identification and photogrammetry to study the reproductive ecology of a resident population of manta rays off Maui, Hawaii, and investigate the reproductive benefits of large body size in each sex. Although reproductive activities occurred year-round, mating trains and late-term pregnant females were significantly more likely to be observed during the winter months. Some females were pursued by males during both winter and summer of the same year, suggesting multiple ovulations may be possible in a single year. Males likely detect a female’s reproductive state by positioning directly behind her, or passing through her bodily excretions. The mean pregnancy rate was estimated at 0.56 pregnancies/adult female/year with larger females pregnant more often, and more likely in consecutive years. The operational sex ratio was heavily skewed with 2.68 adult males per reproductively available female. Although males appear to compete with one another for females within a mating train, no direct physical competition was ever observed between males. Evidence of highly dynamic mating trains lasting more than one day suggests endurance rivalry may be the primary mating strategy among males, during which larger males may benefit from greater energy reserves. The study area appears to be an important staging area for mating individuals in this population.     

Inter-Mount Social Interactions During Heterosexual and Homosexual Consortships in Japanese Macaques

In this study, we examined whether female Japanese macaques ( Macaca fuscata ) exhibited male-typical patterns of inter-mount social behaviour during homosexual consortships. Data were collected on heterosexual and homosexual consortships from a population of free-ranging Japanese macaques in Arashiyama, Japan. Inter-mount intervals were defined as the period between two consecutive mounts. A total of 3374 heterosexual inter-mount intervals and 1412 homosexual inter-mount intervals were analysed. We examined nine different categories of inter-mount behaviour. Sex differences between heterosexually consorting males and females existed for three of the inter-mount behaviours we examined (i.e. ventral-to-dorsal orientation, aggression directed toward third party individuals, tree-shaking displays). The subordinate female partners in homosexually consorting pairs exhibited some sex-atypical inter-mount behaviours. This sex-atypicality did not appear to reflect generalized developmental canalization in the direction of a male-typical suite of behaviours. Rather, the observed behavioural sex-atypicality could be best explained in terms of the unique alliance-related dynamics that characterized homosexual consortships in Japanese macaques.     

Mating system in the tarantula spider Eupalaestrus weijenberghi (Thorell, 1894): evidences of monandry and polygyny

The road tarantula Eupalaestrus weijenberghi shows a strongly female-biased sex ratio since adult females live several years while adult males live only for 2 months. In this scenario selective males could be expected. However, several factors such as the rates of reproduction of each sex, degree of sexual selectivity and synchronicity of female receptiveness determine the operational sexual ratio and mating system of the species. Our objective was to determine the mating rates and mating tactics for females and males of E. weijenberghi and their variation throughout the reproductive period. Four hundred sexual encounters among 20 females and 20 males in all possible pair-wise combinations were carried out during 29 days, a brief but intense experimental period, as it also occurs in the field. Mating success differed strongly between sexes. Females mated once: five females mated at the first attempt, eight initially rejected males and copulated in subsequent attempts. Half of the males did not copulate and the others copulated 1-3 times. Mated females actively rejected males. Results indicate a mating system with monogamous females and polygamous males. Not all the females were receptive in every reproductive season. We suggest that female monogamy drastically affects the operational sex ratio, since several females were unreceptive after a single copulation, directly diminishing the male potential reproductive rate. This is the first experimental approach to estimate tarantula mating systems, their determinants and the consequences of the strategies shown by each sex.     

Eat or Not to Eat: Reversed Sexual Cannibalism as a Male Foraging Strategy in the Spider Micaria sociabilis (Araneae: Gnaphosidae)

Reversed sexual cannibalism represents an unusual situation in which a male kills and consumes a female. We examined this rare phenomenon in the spider Micaria sociabilis, whose males were observed to regularly cannibalise old females. In this study, we investigated male motivation for such behaviour in the light of ecological conditions such as mate availability and prey availability. We found that male cannibalism is not affected by short‐term starvation but rather by male feeding history during the ontogenetic development in combination with prey availability during the adult stage. Males from the summer generation reached bigger sizes than males from the spring generation and females from both generations. They developed in the period with exceptionally high prey occurrence, but when they reached the adult stage, the prey availability decreased. In this period, we observed the highest frequency of cannibalism, however, only when the sex ratio was female biased. Reversed sexual cannibalism in M. sociabilis seems to represent an advantageous male foraging strategy, which is affected by prey availability and male feeding history, tuned by sex ratio and directed towards females of inferior quality.     

Social relationships between adult male and female rhesus macaques: 1. Sexual consortships

The sexual relationships of 15 adult male rhesus macaques (Macaca mulatta), of one social group in the Cayo Santiago colony, Puerto Rico, were studied during the 1981 mating season. Two criteria were used to determine whether or not a focal male was in consort in a given 20-min observational sample. One hundred and thirty-two consortships were recorded. The distribution and duration of all consortships, and the distribution of those consortships that coincided with the estimated time of conception, were positively correlated with male dominance rank and length of tenure. Correlations with dominance were stronger than those with tenure. Older females had more consortships with focal males than younger females. There was no relationship between female rank and the distribution of consortships. Consortships did not give exclusive access to receptive females although they may do in feral situations. Males were primarily responsible for maintaining proximity in about two thirds of consortships. The four top-ranking males were primarily responsible in all but one of their consortships. Almost all of those in which the female was primarily responsible involved younger, slightly lower-ranking males. This pattern may have resulted from the females being attracted to the latter males although other interpretations are possible. The partner who was primarily responsible for maintaining proximity also tended to be the predominant groomer, supporting the view that grooming plays a role in the maintenance of sexual consortships. Almost a third of consortships were with females who had conceived, suggesting that males could not accurately assess female reproductive state. Consortships maintained by the male were longer than those maintained by the female. Males may try to increase their chances of fertilising a female by prolonging the consortship. High-ranking males may have been more successful at this, resulting in the positive correlation between male dominance rank and consortship duration.     

Nonconceptive Sexual Behavior in Bonobos and Capuchins

Sexual behavior by infecundable females, and by same-sex and adult-immature dyads, occurs in wild and captive bonobos (Pan paniscus). Proposed functions of these behaviors, in social primates generally, include practice, paternity confusion, exchange, and communication as well as appeasement. We used this framework to interpret and to compare observations of sexual behavior in a captive bonobo group and a wild white-faced capuchin (Cebus capucinus) group. In both species, (a) sexual behavior was no more frequent in cycling females than in pregnant or lactating females and (b) same-sex and adult-immature dyads engaged in as much mounting or genitogenital contact as adult heterosexual dyads did. The species differed in that (a) bonobos engaged in sexual behavior 65 times as frequently as capuchins, (b) only bonobos engaged in sexual contact other than ventrodorsal mounting during focal observation, and (c) bonobo sexual contact was concentrated most heavily in socially tense situations in adult female–female dyads, whereas capuchin sexual contact was concentrated most heavily in socially tense situations in adult male–male dyads. These data and published literature indicate that (a) practice sex occurs in both species, (b) paternity confusion may be a current function of C. capucinus nonconceptive sex, (c) exchange sex remains undemonstrated in capuchins, and (d) communication sex is more important to members of the transferring sex—female bonobos and male capuchins—than to members of the philopatric sex.     

Homosexual behavior in primates: A review of evidence and theory

Homosexual behavior is defined as genital contact, genital manipulation or both between same-sex individuals. Available data indicate that this behavior is phylogenetically widespread among the anthropoid primates, but totally absent among prosimians. The majority of the 33 species that demonstrate homosexual behavior do so rarefy, but for a substantial number (N =12) it appears to be a more common pattern under free-ranging conditions. I summarize data on homosexual behavior as it relates to form, living condition, age, sex, social organization, and ecological context, and discuss hormonal, demographic, and sociosexual theories for primate homosexual behavior. Among adult primates, the behavior is not the product of abnormal excesses or deficiencies in androgens. Prenatal excesses of androgens may have some effect on the expression of female homosexual behavior, but these effects might vary over the life span, and data are equivocal at present. Demographic processes that result in skewed sex ratios can favor the expression of homosexual behavior in a population, which causes intraspecific variation. I examine several sociosexual explanations, including (a) proceptivity enhancement, (b) receptivity reduction, (c) dominance assertion, (d) practice for heterosexual copulation, (e) tension regulation, (f) reconciliation, and (g) alliance formation. An evolutionary scenario highlights the transformations this behavior underwent during the evolution of the anthropoid primates. I suggest exaptation as a theoretical framework for interpreting homosexual behavior and conclude that future consideration of sexual selection among primates should address homosexual components of this process.     

Avoidance of post-coital cannibalism in the brachyurid paddle crab Ovalipes catharus

Sexual cannibalism was examined experimentally in the New Zealand paddle crab Ovalipes catharus, where it is the female that risks cannibalism during and after sexual activity. In this species copulation only occurs when the female is soft-shelled after moulting and she is most vulnerable to cannibalism. Male paddle crabs protect and copulate with the females until the females are no longer sexually receptive. Males appear able to identify a female with whom they have recently copulated. After a brief separation, significantly fewer females were cannibalized in cases where a female was returned to her original sexual partner than in those that had exchanged partners. The asymmetry, where females gain some protection during moulting but males receive no such advantage, may, at least partially, explain the skewed sex ratios which are sometimes found in samples of O. catharus from nature.     

Female-biased operational sex ratio of sexual host fish in a gynogenetic complex of Carassius auratus

To study the coexistence of sexual and gynogenetic forms, we examined the population structure of a gynogenetic complex of the Japanese crucian carp, Carassius auratus Temminck et Schlegel, during the April–June reproductive season by collecting 1225 mature fish that migrated from Lake Suwa to a tributary river for spawning. There were more sexual fish (about 80%) than gynogenetic fish in this complex, and the operational sex ratio in the sexual form was female biased (males were about 20%). Mean standard length and body weight of sexual females were larger than those of sexual males. Sex ratio was male biased in smaller fish (standard length, <8.5 cm) but female biased in larger fish (standard length, ≥8.5 cm). We determined age by scale ring marks; the average age of sexual females was higher than that of males, but there was no significant difference in the average age between sexual and gynogenetic females. Sex ratio in the sexual form was more female biased for old than for young fish, and the mean size of sexual females was larger than that of males of the same age. The clear female-biased sex ratio and age difference between sexual females and males can be explained either by (1) higher mortality of males or by (2) female-biased sex allocation. The latter process reduces the disadvantage of sex and contributes to the coexistence of sexual and gynogenetic forms. Received: November 24, 2000 / Accepted: March 6, 2001     

Female behavioral strategies during consortship in Tibetan macaques (Macaca thibetana)

Consortship has been defined as a temporary association between an adult male and an estrous/receptive female. It has been considered as male mating strategies to improve male mating success and potential reproductive success. However, the female roles have been more or less neglected, and thus, less is known about female behavioral strategies during the consortship periods. In this study, during the two consecutive mating seasons, we collected behavioral data of free‐ranging Tibetan macaques (Macaca thibetana) habituated in Mt. Huangshan, China, to investigate female behaviors when she was consorted by an adult male. The results showed that (a) females were more likely to approach and exhibit sexual solicitation to their consorting males during the consorted period, and females also exhibited less approach to their nonconsorting males; (b) females exhibited strong responses (either departed distantly or formed affiliative relationships with their consorting male partner) when their consorting males mated with rival females or showed sexual motivation toward rival females; (c) female preferences were positively correlated to the duration of consortships and the frequencies of ejaculation copulations, independent of the social ranks of their consorting male partners. Our results suggested that female strategies played much more important roles in forming and maintaining consortship than previously assumed. It provides new insight into understanding female adaptive strategies to male strategies by forming consortships in multimale–multifemale primate species when males could not identify female''s fertile phase accurately.     

Opposing effects of mortality factors on progeny operational sex ratio may thwart adaptive manipulation of primary sex ratio

Despite extensive research on mechanisms generating biases in sex ratios, the capacity of natural enemies to shift or further skew operational sex ratios following sex allocation and parental care remains largely unstudied in natural populations. Male cocoons of the sawfly Neodiprion abietis (Hymenoptera: Diprionidae) are consistently smaller than those of females, with very little overlap, and thus, we were able to use cocoon size to sex cocoons. We studied three consecutive cohorts of N. abietis in six forest stands to detect cocoon volume‐associated biases in the attack of predators, pathogens, and parasitoids and examine how the combined effect of natural enemies shapes the realized operational sex ratio. Neodiprion abietis mortality during the cocoon stage was sex‐biased, being 1.6 times greater for males than females. Greater net mortality in males occurred because male‐biased mortality caused by a pteromalid parasitic wasp and a baculovirus was greater and more skewed than female‐biased mortality caused by ichneumonid parasitic wasps. Variation in the susceptibility of each sex to each family of parasitoids was associated with differences in size and life histories of male and female hosts. A simulation based on the data indicated that shifts in the nature of differential mortality have different effects on the sex ratio and fitness of survivors. Because previous work has indicated that reduced host plant foliage quality induces female‐biased mortality in this species, bottom‐up and top‐down factors acting on populations can affect operational sex ratios in similar or opposite ways. Shifts in ecological conditions therefore have the potential to alter progeny fitness and produce extreme sex ratio skews, even in the absence of unbalanced sex allocation. This would limit the capacity of females to anticipate the operational sex ratio and reliably predict the reproductive success of each gender at sex allocation.     

Variable Female Mating Positions and Offspring Sex Ratio in the Spider Pityohyphantes phrygianus (Araneae: Linyphiidae)

Chromosomal sex determination and male heterogamety have been thought to seriously impede direct sex ratio control. However, in Pityohyphantes phrygianus, a solitary sheetweb spider with a skewed sex ratio, earlier experimental studies suggested that there are options for female control of offspring sex ratio, if females change their position during the normal mating sequence. Here we show that under natural conditions there is considerable between-female variation in positions, especially after termination of mating. Computer simulations of the orientation of female inner genitalia suggest that sperm are placed in different storage sites depending on the positions adopted. This means that a specific position after mating might potentially influence offspring sex ratio. The variance in offspring sex ratio among females in earlier experiments was binomially distributed, which leads us to conclude that females control the mean sex ratio but do not exercise direct control of the sex of individual offspring.     

Concealed fertility and extended female sexuality in a non-human primate (Macaca assamensis)

In numerous primates living in mixed-sex groups, females display probabilistic cues of fertility to simultaneously concentrate paternity to dominant males while diluting it amongst others as a means to reduce the risk of infanticide and to increase male care for offspring. A few species, however, lack these cues and potentially conceal fertility from males; yet, to date, little is known about mating patterns and their underlying proximate mechanisms in such species. Here, we investigated mating activity and sexual consortships relative to female reproductive state in wild Assamese macaques (Macaca assamensis), a species where females lack prominent anogenital swellings and copulation calls. During two mating seasons (2837 contact hours) we recorded sexual and social behaviors, sexual consortships, and collected 1178 fecal samples (n = 15 females) which were analyzed for progestogen concentrations to assess female reproductive state and to determine the timing of ovulation and conception. Although mostly conceiving in their first ovarian cycle, females were sexually receptive throughout the entire 4-month mating season, and within-cycle mating frequencies were not increased during fertile phases. Dominant males did not monopolize fertile matings, and consortships by high-ranking males lasted for long periods, which were not exclusively linked to female fertile phases. Furthermore, females copulated promiscuously but not randomly, i.e. for almost every female, matings were concentrated to a certain male, irrespective of male rank. Collectively, we demonstrate that fertility is undisclosed to males. The extreme extended female sexuality facilitated by concealed fertility may allow females to create differentiated mating relationships within a promiscuous mating system. Our study provides important new insight into the plasticity of female sexuality in non-human primates.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Functional significance of behavioral,morphological, and endocrine correlates across the ovarian cycle in semifree ranging female Tonkean macaques

The role of sexual displays in mating strategies and their reliability in indicating the time of ovulation has given rise to multiple explanations in nonhuman primates. In order to discriminate among hypotheses, socio-sexual behaviors were recorded in a semifree ranging group of Tonkean macaques (Macaca tonkeana), together with sexual skin swelling volumes and measurements of urinary concentrations of estrone conjugates and pregnanediol glucuronide. A clear preovulatory peak of urinary estrogen levels occurred 2 days before a defined rise in pregnanediol glucuronide concentrations, indicating that both hormones pointed out the female's ovulatory period. The concept of estrus rightly could be applied to female Tonkean macaques since fluctuations in estrogen levels correlated with cyclic changes in genital swelling sizes and rates of female behavioral attractivity and proceptivity. Males proved to be capable of recognizing the optimal conception period as judged from the occurrence of maximal rates of following behavior, serial matings, and ejaculations during the peri-ovulatory phase. During this time, males succeeded in maintaining exclusive and enduring associations with females. However, consortships occurred precociously, with males starting to affiliate with females, follow, and mount them 1 week before the presumed time of ovulation. These long-lasting consortships appear to be a consequence of the female extended follicular phase. This presumably sexually selected character allowed females to extend conspicuous sexual displays: genital swelling and utterance of an estrous call, which might attract males' attention and arouse them. With regard to female mating tactics, the combination of reliably indicating the time of ovulation to the male and durable periods of competitor exclusion led to reject explanations assuming manipulation about paternity or long-lasting intermale competition incitement in Tonkean macaques. Competition for mates between females also turned out to be an irrelevant factor as it was very low in the species. We conclude that the main function of sexual displays is to herald the approach of ovulation toward available mates. Am. J. Primatol. 46:285–309, 1998. © 1998 Wiley-Liss, Inc.     

Female pheromonal chorusing in an arctiid moth, Utetheisa ornatrix

We report an unusual case of communal sexual display in thearctiid moth Utetheisa ornatrix that we designate "female pheromonalchorusing." As in most moths, female U. ornatrix release a long-distancesexual advertisement pheromone during a nightly activity period.We arranged U. ornatrix females in 2 types of signaling conditions:grouped and solitary. When the females were grouped with neighboringsignaling females (grouped), they initiated pheromone releasesooner, continued release with less interruption and over alonger total period, and performed the release with faster abdominalpumping than observed in isolated females (solitary). This differsfrom the usual form of sexual communication in moths: female(chemical) signalers, male receivers, and a general lack ofinteraction among females. At mating, male U. ornatrix transfera large spermatophore that may enhance female reproductive successand which represents either mating effort or paternal investment.This action results in an extended postmating male refractoryperiod leading to a female-biased operational sex ratio. Weargue that this biased sex ratio generates intrasexual competitionamong females, to which they respond by elevating signalingeffort such that the likelihood of at least matching their neighbors'signals is increased. In the field, U. ornatrix are clusteredaround patches of host plants, and we also explore the possibilitythat pheromonal chorusing is driven by cooperation among groupsof related—or nonrelated—females.