Epicuticular compounds and sexual recognition in the ball-roller scarab, Canthon cyanellus cyanellus

We analyzed the role of cuticular compounds in sexual recognition of the carrion ball‐roller scarab, Canthon cyanellus cyanellus (LeConte) (Coleoptera: Scarabaeidae). Males rolled food balls with freeze‐killed females on them as though the females were alive. Acceptance of female cadavers was low when they were washed with solvents and increased significantly when the solvent extract was reapplied to the cadavers. Acceptance of female cadavers by males was significantly higher than that of male cadavers by females. Male cadavers washed with solvent and impregnated with female cuticular compounds were accepted by males as though they were frozen or living females. Similarly, extracted female cadavers impregnated with male cuticular compounds were accepted by females. These results suggest that C. c. cyanellus males recognize females by their cuticular compounds, whereas females use a combination of cuticular compounds and other unknown cues to recognize males.     

Individual recognition through olfactory–auditory matching in lemurs

Individual recognition can be facilitated by creating representations of familiar individuals, whereby information from signals in multiple sensory modalities become linked. Many vertebrate species use auditory–visual matching to recognize familiar conspecifics and heterospecifics, but we currently do not know whether representations of familiar individuals incorporate information from other modalities. Ring-tailed lemurs (Lemur catta) are highly visual, but also communicate via scents and vocalizations. To investigate the role of olfactory signals in multisensory recognition, we tested whether lemurs can recognize familiar individuals through matching scents and vocalizations. We presented lemurs with female scents that were paired with the contact call either of the female whose scent was presented or of another familiar female from the same social group. When the scent and the vocalization came from the same individual versus from different individuals, females showed greater interest in the scents, and males showed greater interest in both the scents and the vocalizations, suggesting that lemurs can recognize familiar females via olfactory–auditory matching. Because identity signals in lemur scents and vocalizations are produced by different effectors and often encountered at different times (uncoupled in space and time), this matching suggests lemurs form multisensory representations through a newly recognized sensory integration underlying individual recognition.     

Tests for sex differences in fitness-linked traits in the spiderAgelenopsis aperta (Araneae: Agelenidae)

The idea that traits linked to individual fitness may differ between males and females was tested in the desert funnel-web spider,Agelenopsis aperta. The study entailed comparison of juvenile male and female behavior with respect to three traits previously shown to be linked to female body mass and ultimately to individual female fitness: habitat discrimination, territorial behavior, and agonistic behavior. As juveniles, male and female spiders behave similarly: they utilize the same habitat cues in locating web sites, maintain similar territory sizes, and exhibit the same behavior patterns in territorial disputes. Like females, males that obtain the highest-quality web sites achieve a greater body mass and are more likely to survive to maturity.     

Seeking a Sex‐Specific Coolidge Effect in a Simultaneous Hermaphrodite

In polygamous mating systems, a capability to discriminate against familiar mates may be beneficial to both sexes. Polyandrous females, for instance, may enhance the odds of finding sires with optimal genetic compatibility or high genetic quality by mating with multiple different males; polygynous males, in addition, may more efficiently invest their limited ejaculates across multiple (rather than single) females. The Coolidge effect facilitates this kind of mate discrimination, as sexual motivation declines across consecutive copulations with a familiar partner but resurrects with a novel mate. In simultaneous hermaphrodites, we expect the Coolidge effect to show sex role‐dependent patterns and vary with previous sex‐specific mating activity. Using the promiscuous hermaphroditic sea slug Chelidonura sandrana, we investigated (1) whether sexual motivation indeed declines when repeatedly exposed to familiar partners, (2) whether the Coolidge effect occurs in a sex‐specific manner and (3) whether ejaculation is strategic with respect to partner familiarity. We found neither mating latency, nor penis intromission duration, mating propensity or the frequency of sex role alternations to vary significantly with treatments. Furthermore, slugs did not donate larger ejaculates to novel than to familiar partners. Partner novelty thus elicited no detectable response in sexual motivation or mating effort in C. sandrana. We suggest that the sea slugs' promiscuous mating system in often large mating aggregations makes mate discrimination based on novelty obsolete in comparison with more relevant criteria such as partner body size or mating history.     

Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition

Across sexually reproducing species, males and females are in conflict over the control of reproduction. At the heart of this conflict in a number of taxa is male harassment of females for mating opportunities and female strategies to avoid this harassment. One neglected consequence that may result from sexual harassment is the disruption of important social associations. Here, we experimentally manipulate the degree of sexual harassment that wild female guppies (Poecilia reticulata) experience by establishing replicated, semi-natural pools with different population sex ratios. We quantify the effects of sexual harassment on female social structure and the development of social recognition among females. When exposed to sexual harassment, we found that females had more disparate social networks with limited repeated interactions when compared to females that did not experience male harassment. Furthermore, females that did not experience harassment developed social recognition with familiar individuals over an 8-day period, whereas females that experienced harassment did not, an effect we suggest is due to disruption of association patterns. These results show that social network structure and social recognition can be affected by sexual harassment, an effect that will be relevant across taxonomic groups and that we predict will have fitness consequences for females.     

Low mate encounter rate increases male risk taking in a sexually cannibalistic praying mantis

Male praying mantises are forced into the ultimate trade-off of mating versus complete loss of future reproduction if they fall prey to a female. The balance of this trade-off will depend both on (1) the level of predatory risk imposed by females and (2) the frequency of mating opportunities for males. We report the results of a set of experiments that examine the effects of these two variables on male risk-taking behavior and the frequency of sexual cannibalism in the praying mantis Tenodera sinensis. We experimentally altered the rate at which males encountered females and measured male approach and courtship behavior under conditions of high and low risk of being attacked by females. We show that male risk taking depends on prior access to females. Males with restricted access to females showed greater risk-taking behavior. When males were given daily female encounters, they responded to greater female-imposed risk by slowing their rate of approach and remained a greater distance from a potential mate. In contrast, males without recent access to mates were greater risk-takers; they approached females more rapidly and to closer proximity, regardless of risk. In a second experiment, we altered male encounter rate with females and measured rates of sexual cannibalism when paired with hungry or well-fed females. Greater risk-taking behavior by males with low mate encounter rates resulted in high rates of sexual cannibalism when these males were paired with hungry females.     

Male-derived cuticular hydrocarbons signal sperm competition intensity and affect ejaculate expenditure in crickets

Female sexual promiscuity can have significant effects on male mating decisions because it increases the intensity of competition between ejaculates for fertilization. Because sperm production is costly, males that can detect multiple matings by females and allocate sperm strategically will have an obvious fitness advantage. The presence of rival males is widely recognized as a cue used by males to assess sperm competition. However, for species in which males neither congregate around nor guard females, other more cryptic cues might be involved. Here, we demonstrate unprecedented levels of sperm competition assessment by males, which is mediated via the use of chemical cues. Using the cricket Teleogryllus oceanicus, we manipulated male perception of sperm competition by experimentally coating live unmated females with cuticular compounds extracted from males. We found that males adjusted their ejaculate allocation in response to these compounds: the viability of sperm contained within a male's ejaculate decreased as the number of male extracts applied to his virgin female partner was increased. We further show that males do not respond to the relative concentration of male compounds present on females, but rather to the number of distinct signature odours of individual males. Our results conform to sperm competition theory, and show for the first time, to our knowledge, that males can detect different intensities of sperm competition by using distinct chemical cues of individual males present on females.     

Social preferences based on sexual attractiveness: a female strategy to reduce male sexual attention

Male sexual harassment of females is common across sexually reproducing species and can result in fitness costs to females. We hypothesized that females can reduce unwanted male attention by constructing a social niche where their female associates are more sexually attractive than themselves, thus influencing the decision-making of males to their advantage. We tested this hypothesis in the Trinidadian guppy (Poecilia reticulata), a species with high levels of male sexual harassment. First, we confirmed that non-receptive females were harassed less when they were paired with a more sexually attractive (receptive) female than with another non-receptive female. We then found that, indeed, females exploit this as a strategy to reduce sexual harassment; non-receptive females actively preferred to associate with receptive over non-receptive females. Importantly, when given access only to chemosensory cues, non-receptive females still showed this preference, suggesting that they use information from chemical cues to assess the sexual attractiveness of potential female partners. Receptive females in contrast showed no such preferences. Our results demonstrate that females can decrease male harassment by associating with females that are more sexually attractive than themselves and that they perform active partner choices based on this relative attractiveness. We propose that this strategy is likely to represent an important pathway by which females can construct social niches that influence the decision-making of others to their advantage; in this case, to reduce the sexual harassment they experience.     

Conflict resolution in the Odonata: implications for understanding female mating patterns and female choice

Predictions of mating patterns in animals have focused on males and how they compete for fertilizations by controlling females. With reference to the Odonata, a taxon in which mating requires cooperation of the female, the active role that females play in mating decisions is often ignored, leading to the premature conclusion that male coercion of females is common. A critical review of the outcome of sexual conflict among odonates leads me to alternative explanations of female mating patterns that need to be refuted before concluding that males coerce matings. Because Anisoptera males have greater control over tandem formation, they have a greater potential for coercion than Zygoptera males. However, Anisoptera females may simply be willing to remate more often if they receive insufficient sperm to fertilize an entire egg clutch. Contrary to prior assumptions, in both suborders, male defence of oviposition sites does not preclude females from choosing among sites or among males. I find that the evolution of non-aggressive sexual signals by males is a reliable indication that sexual conflict has been resolved in favour of female interests. Although I predict that the benefits to females of choice of male phenotype should rarely exceed the cost of such discrimination in Odonata, female choice is most likely to evolve in territorial species whose males must endure high physiological stress in order to mate, and when site quality is not a reliable predictor of the genetic quality of a potential mate.     

Genetics of Drosophila simulans male mating discrimination in crosses with D. melanogaster

The genetic bases of sexual isolation between Drosophila melanogaster and D. simulans have been mainly studied in females, and there is little information about the role of the males in interspecific mating discrimination. Using D. simulans synthetic lines with compound chromosomes from a population of the Seychelles Islands (high frequency of interspecific mating) and a multimarker strain (low frequency), we show that D. simulans males play an important role in discriminating D. melanogaster females. The genetics of male discrimination fits well with the inheritance mode of a single locus, dominant for sexual isolation, located in chromosome II near the net mutation (2L-0.0). The heterospecific mating success of the male was not related to his sexual vigor. The specific load of male cuticular hydrocarbons was counted as a possible source of discrimination used by the D. melanogaster female.     

Female discrimination thresholds frequently exceed local male display variation: implications for mate choice dynamics and sexual selection

Among the factors that can influence female mate choice decisions is the degree to which females differentiate among similar displays: as differences decrease, females are expected to eventually stop discriminating. This discrimination threshold, in conjunction with the magnitude of male trait variation females regularly encounter while making mate choice decisions, may have important consequences for sexual selection. If local display variation is above the discrimination threshold, female preferences should translate into higher mating success for the more attractive male. But if display variation is frequently below the threshold, the resulting increased pattern of random mating may obscure the existence of female mate choice. I investigated the interplay between female discrimination and male display variation in green treefrogs (Hyla cinerea) and found that call trait differences between nearest neighbour males were frequently smaller than what females are expected to discriminate. This finding has two important consequences for our understanding of sexual selection in the wild: first, low display variation should weaken the strength of selection on male display traits, but the direction of selection should mirror the one predicted from females choice trials. Second, caution is needed when interpreting data on realized mating success in the wild: a pattern of random mating with respect to male display traits does not always mean that female preferences are weak or that conditions are too challenging for females to express their preferences. Rather, insufficient display variation can generate the same pattern.     

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.     

Sex-specific responses to sexual familiarity,and the role of olfaction in Drosophila

Studies of mating preferences have largely neglected the potential effects of individuals encountering their previous mates (‘directly sexually familiar’), or new mates that share similarities to previous mates, e.g. from the same family and/or environment (‘phenotypically sexually familiar’). Here, we show that male and female Drosophila melanogaster respond to the direct and phenotypic sexual familiarity of potential mates in fundamentally different ways. We exposed a single focal male or female to two potential partners. In the first experiment, one potential partner was novel (not previously encountered) and one was directly familiar (their previous mate); in the second experiment, one potential partner was novel (unrelated, and from a different environment from the previous mate) and one was phenotypically familiar (from the same family and rearing environment as the previous mate). We found that males preferentially courted novel females over directly or phenotypically familiar females. By contrast, females displayed a weak preference for directly and phenotypically familiar males over novel males. Sex-specific responses to the familiarity of potential mates were significantly weaker or absent in Orco¹ mutants, which lack a co-receptor essential for olfaction, indicating a role for olfactory cues in mate choice over novelty. Collectively, our results show that direct and phenotypic sexual familiarity is detected through olfactory cues and play an important role in sex-specific sexual behaviour.     

Discrimination of conspecific individuals via cuticular pheromones by males of the cricket Gryllus bimaculatus

Cuticular substances on the body surface of crickets serve as pheromones that elicit a variety of different behaviors in male crickets. Antennal contact between males and females resulted in courtship behavior, and that between two males resulted in aggressive displays. As a first step in elucidating how crickets recognize and discriminate individuals, behavioral responses of male individuals to cuticular substances of conspecific males or females were investigated. The behavioral responses of males to antennal or palpal stimulation with an isolated antenna from a male or a female were recorded. To both antennal and palpal stimulation with female antennae, the majority of males responded with courtship behavior; to stimulation with male antennae, males responded with aggressive displays. To gain insight into the chemical nature of the behaviorally relevant components, isolated antennae were washed in either n-hexane, acetone or ethanol before behavior assays. Washed antennae no longer elicited courtship or aggressive responses in males. Next, polypropylene fibers were smeared with substances from the body surface of females and used for antennal stimulation. This experiment showed that the quality and quantity of cuticular substances appear to be highly age-dependent. Significantly more males responded with courtship behavior to cuticular substances from younger females. Isolated males generally showed higher levels of aggression than males reared in groups. Grouped males also were more likely to display courtship behavior towards antennae from younger females, and aggressive behavior towards antennae from older females. These results suggest that male discrimination of mating partners depends on the nature of female cuticular substances.     

Effects of social experience on subsequent sexual performance in naïve male Japanese quail (Coturnix japonica)

On their first sexual encounter, naïve male Japanese quail will attend to and approach a female; they sometimes mount but they do not always copulate. During the second encounter, most males successfully copulate. Although sexual experience facilitates subsequent sexual interactions, sensory cues provided by females, independent of any sexual encounter, may also enhance sexual performance. To investigate whether previous exposure to a conspecific affects subsequent sexual behavior, we allowed inexperienced males to observe an empty box, or a conspecific consisting of either an experienced female or male for 2.5 min/day on 7 days. Measures of appetitive sexual behavior were recorded during these tests. On day 8, subjects were allowed to copulate with a novel female for 5 min. On the following days, all subjects were repeatedly provided with visual access to a female and allowed to mate. In the pre-copulatory trials males initially exhibited a high frequency of appetitive responses that dissipated with repetition. Pre-copulatory experience also significantly affected motivation to mate with subjects exposed to females copulating more quickly than other subjects. Post-copulatory appetitive behavior also differed between groups: control subjects showed the highest behavioral frequency followed by males exposed to females and finally males exposed to males. These data indicate that pre-copulatory social experience profoundly influences subsequent sexual behavior and probably reproductive success. This experience effect is independent of any hormonal effect (such as one resulting from changes in secretion following different social interactions) given that the subjects were castrates chronically treated with testosterone.     

Females prefer to associate with males with longer intromittent organs in mosquitofish

Sexual selection is a major force behind the rapid evolution of male genital morphology among species. Most within-species studies have focused on sexual selection on male genital traits owing to events during or after copulation that increase a male''s share of paternity. Very little attention has been given to whether genitalia are visual signals that cause males to vary in their attractiveness to females and are therefore under pre-copulatory sexual selection. Here we show that, on average, female eastern mosquitofish Gambusia holbrooki spent more time in association with males who received only a slight reduction in the length of the intromittent organ (‘gonopodium’) than males that received a greater reduction. This preference was, however, only expressed when females chose between two large males; for small males, there was no effect of genital size on female association time.     

A phylogenetic analysis of the evolution of sexual dimorphism and mating systems in water striders (Hemiptera: Gerridae)

Conflicts over mating decisions characterize the sexual behaviour of many insects, in particular when males encounter females that already carry enough sperm to fertilize their eggs, since a mating often will inflict greater costs than benefits upon females. Therefore, coevolutionary models predict adaptation and counter-adaptation by the sexes in a battle to control the outcome of sexual encounters. A phylogenetic analysis was performed on patterns of sexual dimorphism and mating systems within water striders (Hemiptera, Gerridae). Phylogenetic effects or 'constraints' have significantly shaped patterns of sexual dimorphism in female/ male size ratios, legs and genitalia of males, and the structure of the female abdomen. Males of ancestral gerrids were probably slightly smaller than conspecific females, had powerful fore legs adapted to grasp the female's thorax during mating, and had clasping genitalic structures suited to grasp or pinch the female posteriorly. Most gerrids have a female/male size ratio between 1.05 and 1.14, but more pronounced sexual size ratios (above 1.25) have independently evolved several times in the family, usually in association with extended post-copulatory mate guarding. The comparative, phylogenetic analysis suggests coevolution of female anticlasper and male clasping devices for the clade comprising the subfamilies Cylindrostethinae, Ptilomerinae, and Halobatinae while female anticlasper devices have evolved in the absence of male clasping genitalia in the Gerrinae. The ancestral and most common mating system in gerrids is 'scramble competition polygyny' from which has evolved 'resource defence polygyny' at least four times independently of each other. The phylogenetic effects on patterns of mating behaviour are much less obvious, as exemplified by the large amount of interspecific variation in some genera.     

The Allure of the Distinctive: Reduced Sexual Responsiveness of Female Guppies to 'Redundant' Male Colour Patterns

Frequency-dependent mating behaviour has the potential to maintain genetic variation in characteristics of organisms. The colour patterns of guppies ( Poecilia reticulata ) provide an example of one of the most extreme genetically based polymorphisms known in nature, for which frequency-dependent mate choice could be a mechanism. Numerous studies have shown that female guppies base mating preferences on male colour patterns and there is evidence that females prefer to mate with males displaying novel or unfamiliar colour patterns. This preference could lead to frequency-dependent mating success in males. Nevertheless, the possibility that female sexual responsiveness itself may depend on the frequency of male types has not been tested systematically in guppies or any other species. This study examined the sexual responses of female guppies in experimental groups consisting of two males with similar (redundant) and two males with different (unique) colour patterns. We found that female guppies were much more likely to respond sexually to the displays of unique males than to those of redundant males. Further, there was no effect of orange colouration on female responsiveness as has been documented for this population in several previous studies, thus, discrimination against redundant male types appears to have overridden directional selection based on colour pattern characteristics. This discrimination against redundant male types could in turn lead to frequency-dependent mating success in males and maintenance of colour pattern polymorphism.     

Self-grooming induced by sexual chemical signals in male root voles (Microtus oeconomus Pallas)

Sniffing is one-way animals collect chemical signals, and many males self-groom when they encounter the odor of opposite-sex conspecifics. We tested the hypothesis that sexual chemical signals from females can induce self-grooming behavior in male root voles (Microtus oeconomus Pallas). Specifically, we investigated the sniffing pattern of male root voles in response to odors from the head, trunk, and tail areas of lactating and non-lactating females. The self-grooming behavior of males in response to female individual odorant stimuli was documented, and the relationship between self-grooming and sniffing of odors from the head, trunk, and tails areas were analyzed. Sniffing pattern results showed that males are most interested in odors from the head area, and more interested in odors from the tail as compared to the trunk area. Males displayed different sniffing and self-grooming behaviors when they were exposed to odors from lactating females as compared to non-lactating females. Males also spent more time sniffing and engaged in more sniffing behaviors in response to odors from the lactating females’ tail area as compared to the same odors from non-lactating females. Similarly, males spent more time self-grooming and engaged in more self-grooming behaviors in the presence of individual odors from lactating females as compared to individual odors from non-lactating females. Partial correlation analyses revealed that the frequency of self-grooming was significantly correlated with the frequency of tail area sniffs. Results from this experiment suggest that sexual attractiveness of lactating females is stronger than that of non-lactating females. Furthermore, the partial correlation analysis demonstrated that self-grooming in males is induced by odors from the tail area of females. Collectively, these results support the hypothesis that sexual chemical signals from females can induce self-grooming behavior in male root voles. Self-grooming may also reflect the groomer's sexual motivation and facilitate sexual interactions.     

Male-Biased Sex Ratios in Offspring of Attractive Males in the Guppy

The attractiveness hypothesis predicts that females produce offspring with male-biased sex ratios when they mate with attractive males because their male offspring will inherit the paternal sexual attractiveness and may have high reproductive success. In this study, we examined the effect of the attractiveness of the male guppy Poecilia reticulata in terms of the conspicuousness of its orange spot patterns, important criteria affecting female choice in this species, on the offspring sex ratios. We found that food-manipulation treatment altered the conspicuousness of the orange spot patterns in a full-sibling male pair. When females were presented to these males, they showed a greater mate preference for males having brighter orange spots than for those having duller orange spots. Subsequently, half of the females were mated with the preferred males and the remaining females were mated with the less preferred males. When the females exhibited a greater preference for their mates, their offspring sex ratios were more male biased. These results appear to be consistent with the prediction of the attractiveness hypothesis. In the guppy, as male sexual attractiveness is heritable, the male-biased sex ratios of the broods of attractive males may be adaptive.