The role of cuticular hydrocarbons in male attraction and repulsion by female Dawson's burrowing bee, Amegilla dawsoni

Cuticular hydrocarbons (CHCs) are long-chain fatty acids and their derivatives that protect insects from desiccation. They can also be important semiochemicals in insect reproduction. We used behavioural and chemical assays to examine the potential role of CHCs in sexual communication in a solitary burrowing bee, Amegilla dawsoni. Washing CHC blends from the cuticle of emerging virgin females made them unattractive to mate-searching males. Returning the CHC blends restored their attractiveness. Nesting females were unattractive to mate-searching males, whether they were washed or not. Chemical analysis identified significant differences between male and female CHC blends and between virgin female and nesting female blends. Some of these differences were due to specific compounds. Loss of attractiveness is unlikely to be due to antiaphrodisiac compounds delivered by males, because male-specific compounds were not found on nesting females, and because recently mated females with intact CHC blends were attractive to searching males. Nesting females could not be made attractive to searching males by removing their CHC blends. Adding virgin female CHC blends tended to improve attractiveness but the effect was weak, suggesting that some form of volatile compound may also be involved in signalling unreceptivity.     

Indirect benefits for choosy female grasshoppers (Chorthippus biguttulus)?

Since direct benefits are likely to be absent in the grasshopper Chorthippus biguttulus, indirect genetic benefits are a potential explanation for costly female preference. Choosy females may improve their fitness in terms of enhanced attractiveness of sons alone or additionally by improved viability of offspring. We tested the predictions of these two hypotheses by comparing attractiveness-related song traits and viability in offspring of attractive and unattractive grasshopper males. The experiment was conducted with larvae reared under semi natural lab conditions in one year and under natural conditions in the field in the following year. If reared under natural conditions no significant differences in viability and song traits between offspring of attractive and unattractive males could be found. Offspring reared in the lab produced calling songs with a significantly more exact song rhythm when sired by attractive males than offspring of unattractive males. Offspring of attractive males should thus have a theoretical advantage in mate choice, which, however, did not translate into higher attractiveness values in acoustic female choice experiments. Therefore our experiments could not resolve whether female choice in C. biguttulus evolved according to the sexy son hypothesis. Since viability in offspring of attractive males did not differ from offspring of unattractive males, “good genes” seems unlikely to be the underlying mechanism of female choice.     

Mating is associated with immediate changes of the hydrocarbon profile of Leptothorax gredleri ant queens

While sexual communication is often characterized by attempted manipulation, both sexes agree about females reliably signalling their receptivity. Female sexuals of the ant Leptothorax gredleri quickly became unattractive to males after their first copulation. This loss of attractiveness coincided with almost immediate changes in their cuticular hydrocarbon (CHC) profiles. Already 30 min after mating, the CHC profiles of female sexuals had significantly lower relative amounts of branched alkanes and higher amounts of linear alkanes than those of unmated and freshly mated female sexuals. Discriminant analysis did not distinguish between the profiles of freshly mated and unmated female sexuals, suggesting that the extremely rapid modification of CHC profiles is not caused by males marking females with anti-aphrodisiac CHCs. Instead, the new profile is produced by the female sexuals themselves. In addition to making them unattractive to males, this change may also help mated female sexuals when seeking adoption into established colonies.     

The indirect benefits of mating with attractive males outweigh the direct costs

The fitness consequences of mate choice are a source of ongoing debate in evolutionary biology. Recent theory predicts that indirect benefits of female choice due to offspring inheriting superior genes are likely to be negated when there are direct costs associated with choice, including any costs of mating with attractive males. To estimate the fitness consequences of mating with males of varying attractiveness, we housed female house crickets, Acheta domesticus, with either attractive or unattractive males and measured a variety of direct and indirect fitness components. These fitness components were combined to give relative estimates of the number of grandchildren produced and the intrinsic rate of increase (relative net fitness). We found that females mated to attractive males incur a substantial survival cost. However, these costs are cancelled out and may be outweighed by the benefits of having offspring with elevated fitness. This benefit is due predominantly, but not exclusively, to the effect of an increase in sons' attractiveness. Our results suggest that the direct costs that females experience when mating with attractive males can be outweighed by indirect benefits. They also reveal the value of estimating the net fitness consequences of a mating strategy by including measures of offspring quality in estimates of fitness.     

Walking in Fourier’s space: algorithms for the computation of periodicities in song patterns by the cricket Gryllus bimaculatus

Is discrimination of the envelope of an acoustic signal based on spectral or temporal computations? To investigate this question for the cricket Gryllus bimaculatus, pattern envelopes were constructed by the addition of several sine waves and modified by systematic phase changes. The phonotactic response of female crickets towards such sinusoidal but also rectangular pulse patterns was quantified on a locomotion compensator. Envelope patterns that exhibited a modulation frequency of 25 Hz as the dominant frequency were attractive and although changes of phase modified the temporal pattern, the values of attractiveness remained unaffected. Removal of the 25-Hz component reduced the phonotactic scores. Patterns in which other frequency components exhibited a larger amplitude than the 25-Hz component were less attractive. However, the combination of an unattractive pulse period with the attractive modulation frequency of 25 Hz in a pattern revealed that such stimuli were unattractive despite the presence of the 25-Hz component. A comparison of the attractiveness of all patterns revealed that female crickets evaluated the duration of pulse period over a wide range of duty cycles. The combined evidence showed that pattern envelopes were processed in the time- and not in the spectral domain.     

Male Courtship Behavior and Weapon Trait as Indicators of Indirect Benefit in the Bean Bug,Riptortus pedestris

Females prefer male traits that are associated with direct and/or indirect benefits to themselves. Male–male competition also drives evolution of male traits that represent competitive ability. Because female choice and male–male competition rarely act independently, exploring how these two mechanisms interact is necessary for integrative understanding of the evolution of sexually selected traits. Here, we focused on direct and indirect benefits to females from male attractiveness, courtship, and weapon characters in the armed bug Riptortus pedestris. The males use their hind legs to fight other males over territory and perform courtship displays for successful copulation. Females of R. pedestris receive no direct benefit from mating with attractive males. On the other hand, we found that male attractiveness, courtship rate, and weapon size were significantly heritable and that male attractiveness had positive genetic covariances with both courtship rate and weapon traits. Thus, females obtain indirect benefits from mating with attractive males by producing sons with high courtship success rates and high competitive ability. Moreover, it is evident that courtship rate and hind leg length act as evaluative cues of female choice. Therefore, female mate choice and male–male competition may facilitate each other in R. pedestris. This is consistent with current basic concepts of sexual selection.     

Acquired resistance affects male sexual display and female choice in guppies

Is resistance to parasites related to the expression of male secondary sex characters? Handicap models predict a positive relationship, proposing that males displaying extravagant sex characters may be honestly signalling their resistance to females. However, no current evidence addresses whether individual changes in immunity (acquired resistance) are reflected in sexual traits. In this experiment I use guppies to compare male orange colour, sigmoid display and female preferences for individual males, before and after a primary challenge infection of males. Challenge infections were terminated chemically and fish were given ten days'' recovery time before proceeding with the second measurements. The degree of acquired resistance was quantified a posteriori, by exposing males to a secondary infection. Sigmoid display rates and female preference for males differed for males of different resistance groups after challenge infection only. This difference was due to resistant males displaying more than non-resistant ones. No differences were detected in male orange colour, but this may be because colour needs a longer time than ten days to be recovered and adjusted. The results show that the level of acquired resistance affects sexual display and attractiveness in guppies. They suggest that once an effective immunity is built up by a male, he can afford to incur higher costs for sexual characteristics, whereas a male that lacks the ability to build up effective resistance cannot. These costs probably consist of higher energy expenditure and/or higher circulating levels of testosterone, which may be needed to increase display. Priming and effective establishment of an individual''s resistance to parasitic infection could eventually result in a higher availability of resources for sexual functions.     

Sperm: seminal fluid interactions and the adjustment of sperm quality in relation to female attractiveness

An important predictor of male fitness is the fertilizing efficiency of their ejaculates. Ejaculates are costly to produce and males are predicted to devote greater resources to copulations with reproductively superior females. It is well established that males allocate different numbers of sperm to ejaculates. However, less is known about how males adjust their sperm quality, which has important implications for our understanding of fertilization and the evolution of sexual strategies. Here we test in the fowl, Gallus gallus, whether males adjust their sperm velocity by differentially allocating seminal fluid to copulations with attractive and unattractive females. To disentangle the contributions of sperm and seminal fluid to sperm velocity, we separated and remixed sperm and seminal fluid from ejaculates allocated to females of different attractiveness. We show that dominant males increase the velocity of the sperm they invest in more attractive females by allocating larger ejaculates that contain seminal fluid that increases sperm velocity. Furthermore, we find weak evidence that males also allocate sperm with higher velocity, irrespective of seminal fluid, to more attractive females.     

ABSOLUTE VERSUS RELATIVE MEASUREMENTS OF SEXUAL SELECTION: ASSESSING THE CONTRIBUTIONS OF ULTRASONIC SIGNAL CHARACTERS TO MATE ATTRACTION IN LESSER WAX MOTHS,ACHROIA GRISELLA (LEPIDOPTERA: PYRALIDAE)

When females choose a mate among a group of signaling males concentrated in a small area, a male's mating success is often determined not only by his absolute attractiveness but by the attractiveness of his neighbors as well. Multivariate analyses of sexual selection measurements based on absolute values of predictor variables are then misleading, because such analyses assume that the fitness of a given individual is not influenced by others. We addressed this problem of relative fitness in sexual selection by developing two adjustments of the predictor variables in the multivariate analyses by including group means in addition to absolute values and by using relative values, deviations from group means. In the lesser wax moth, Achroia grisella (Lepidoptera: Pyralidae), aggregated males produce ultrasonic signals attractive to females in the vicinity of honey bee colonies. Playback experiments showed that females prefer signals whose pulses are louder, longer, delivered at a faster rate, and include lengthy silent gaps within pulse pairs (long asynchrony intervals). To measure sexual selection on ultrasonic signals, attractiveness was assessed by observing the number of females orienting toward given males in four-choice trials. The various signal characters of the males were computed from ultrasound recordings. Multivariate regression and nonparametric analyses revealed that peak amplitude, asynchrony interval, and pulse rate were the direct targets of selection. Nonlinear stabilizing and correlational selection were also detected. Comparisons of results from multivariate analyses based on absolute values and on the two relative adjustments indicated that the quality of a four-male group did not affect the strength or direction of linear selection but changed the strength of nonlinear selection. Both relative adjustments improved the prediction of male attractiveness. Three-dimensional surface plots generated by the nonparametric regression analyses showed that attractiveness increased monotonically with any combination of the signal characters. This graphical technique also showed that female choice in A. grisella was based on a relative, as opposed to a threshold, decision rule. Overall, male attractiveness for the population remained consistent between testing days. However, attractiveness values for some individual males changed between successive testing days. In these cases, the change in attractiveness was related only to the change in peak amplitude.     

Sexual conflict and cryptic female choice in the black field cricket, Teleogryllus commodus

The prevalence and evolutionary consequences of cryptic female choice (CFC) remain highly controversial, not least because the processes underlying its expression are often concealed within the female reproductive tract. However, even when female discrimination is relatively easy to observe, as in numerous insect species with externally attached spermatophores, it is often difficult to demonstrate directional CFC for certain male phenotypes over others. Using a biological assay to separate male crickets into attractive or unattractive categories, we demonstrate that females strongly discriminate against unattractive males by removing their spermatophores before insemination can be completed. This results in significantly more sperm being transferred by attractive males than unattractive males. Males respond to CFC by mate guarding females after copulation, which increases the spermatophore retention of both attractive and unattractive males. Interestingly, unattractive males who suffered earlier interruption of sperm transfer benefited more from mate guarding, and they guarded females more vigilantly than attractive males. Our results suggest that postcopulatory mate guarding has evolved via sexual conflict over insemination times rather than through genetic benefits of biasing paternity toward vigorous males, as has been previously suggested.     

Sexual conflict over mating in Gnatocerus cornutus? Females prefer lovers not fighters

Female mate choice and male–male competition are the typical mechanisms of sexual selection. However, these two mechanisms do not always favour the same males. Furthermore, it has recently become clear that female choice can sometimes benefit males that reduce female fitness. So whether male–male competition and female choice favour the same or different males, and whether or not females benefit from mate choice, remain open questions. In the horned beetle, Gnatocerus cornutus, males have enlarged mandibles used to fight rivals, and larger mandibles provide a mating advantage when there is direct male–male competition for mates. However, it is not clear whether females prefer these highly competitive males. Here, we show that female choice targets male courtship rather than mandible size, and these two characters are not phenotypically or genetically correlated. Mating with attractive, highly courting males provided indirect benefits to females but only via the heritability of male attractiveness. However, mating with attractive males avoids the indirect costs to daughters that are generated by mating with competitive males. Our results suggest that male–male competition may constrain female mate choice, possibly reducing female fitness and generating sexual conflict over mating.     

Sexes of a monomorphic species differ in preference for mates with a novel trait

We investigated the different roles of the sexes in the originationof novel traits in the sexually monomorphic Javanese mannikinLonchura leucogastroides. We introduced a red feather as anevolutionarily novel trait in both sexes and tested their preferencesfor prospective mates with this trait. Males rejected femalesbearing the red feather and preferred to court unadorned females.In contrast, females partly preferred adorned males. Specifically,previously unattractive males gained in attractiveness and could increasetheir reproductive success when bearing the ornament, whereas previouslyattractive males lost in attractiveness, but this did not affect theirreproductive success. We introduced two other novel traits inmales and investigated the females' response to these in matechoice tests. Each of the three new traits interacted with thenatural attractiveness of males. The more attractive a malewas before ornamentation, the more it lost in attractiveness afterornamentation and vice versa. Thus, the position of the traitdid not affect the interaction. Because males rejected adornedfemales and females partly preferred adorned males, novel traitsmight evolve by intersexual selection in males rather than infemales. This can lead to a sexual dimorphism with conspicuoustraits in males. Our study reveals a new insight into the mechanismof the evolution from monomorphism to dimorphism with ornamentaltraits in males.     

Hot or Not: The Effects of Exogenous Testosterone on Female Attractiveness to Male Conspecifics in the Budgerigar

An increasing number of studies indicate that not only females but also males can be selective when choosing a mate. In species exhibiting male or mutual mate choice, females may benefit from being attractive. While male attractiveness is often positively influenced by higher plasma levels of the androgenic hormone testosterone, it has been shown that testosterone can masculinise female behavior and morphology in several bird species, potentially rendering them less attractive. In this study, we investigated whether female budgerigars, Melopsittacus undulatus , suffer from increased plasma testosterone levels through a negative effect on their attractiveness to males. We experimentally increased plasma testosterone levels in testosterone-treated females (T-females) compared to controls (C-females) and allowed males to choose between a T- and a C-female in a two-way choice situation. Although testosterone treatment significantly affected female behavioral and morphological characteristics, males did not show a significant difference in preference between T- and C-females. These results suggest that experimentally increasing testosterone levels in females does not appear to influence male preference during initial mate choice. Our findings indicate that selection for higher levels of testosterone in male budgerigars is probably not constrained by a correlated response to selection causing negative effects on female attractiveness during initial mate choice. Evaluating whether or not a potential constraint may arise from negative testosterone-induced effects on other fitness related traits in females requires further work.     

Scototaxis and shape discrimination in the female cricket Acheta domesticus in an arena and on a compensatory treadmill

ABSTRACT. The scototactic responses of adult female Acheta domesticus L. were tested toward various shaped targets in an orientation arena and on a compensatory treadmill. In an arena, crickets oriented toward dark targets (positive scototaxis) if they had horizontal visual angles ≥30, and if the vertical dimensions of the target was ≤ its horizontal dimension. Unattractive targets did not result in negative scototaxis but caused crickets to orient randomly with no net directionality. When complex targets were composed of two or more simple rectangular targets which had been previously defined as attractive or unattractive, intermediate responses were obtained. Crickets oriented less toward complex targets than toward simple attractive targets, but oriented more than toward simple unattractive targets. The responses of a female cricket toward an attractive target can be modified by the presence of a chemical signal emitted by previously tested females. When the signal is present females are less likely to orient toward a target that would otherwise be very attractive. This effect was not as great when the females being tested were taken directly from densely populated colonies.
When running on a compensatory treadmill, female crickets exhibit scototactic tendencies similar to those displayed in the arena. When tested on the treadmill over long periods, the amount of time spent orienting toward an attractive target increased.     

Variation in oxygen consumption of the western diamondback rattlesnake (Crotalus atrox): implications for sexual size dimorphism

Variation in metabolism affects energy budgets of individuals and may serve as a mechanism that influences variation at whole organism or population levels. For example, sex differences in metabolic expenditure may contribute to bioenergetic sources of sexual size dimorphism. We measured oxygen consumption rates of 48 western diamondback rattlesnakes (Crotalus atrox) from a sexually dimorphic population and tested the effects of body mass, body temperature and time of day, in three groups of snakes: males, non-reproductive females, and vitellogenic females. Metabolic rates of male and non-reproductive female C. atrox were similar to rates reported for other rattlesnakes (mass exponents ranging from 0.645–0.670). Oxygen consumption was affected by body mass, body temperature and time of day, and was approximately 1.4 times greater in vitellogenic females than in non-reproductive females. No differences were found between males and non-reproductive females. Accordingly, differences in metabolic rate apparently do not contribute directly to sexual dimorphism in this population. Nevertheless, estimates of size-dependent maintenance expenditure lead us to hypothesize that adult female body size may represent a compromise between selection for increased litter size (accomplished by increasing body size), and selection for increased reproductive frequency (accomplished by decreasing body size, and, therefore inactive maintenance expenditure); this is a mechanistic scenario suggested previously for some endotherms. Accepted: 20 May 1998     

Changes in male sexual responses from silk cues of females at different reproductive states in the wolf spider Schizocosa malitiosa

Male wolf spiders are capable of recognising sexual signals associated with female silk threads. In the wolf spider Schizocosa malitiosa variations in female receptivity have been studied, but changes in female silk attractiveness remain unknown. We analysed the sexual responses of adult males (leg shaking, papal drumming and searching) exposed to silk cues from subadult, virgin and mated females of different ages, and females that were or were not carrying an egg-sac. Penultimate and recently moulted adult females elicited low levels of male sexual behaviour, while those of virgin females (21–40 days old) were the most attractive. Silk threads slowly became less attractive after mating. Cues from females carrying an egg-sac as well as females in the inter egg-sac period were fairly attractive. The low attractiveness of recently moulted females disagrees with their high sexual receptivity. In contrast, females continued to elicit strong male responses during a 10-day period after mating, despite the fact that they immediately become sexually reluctant, suggesting strong selection for male searching ability. Low attractiveness during the egg-carrying period could reflect the fact that females do not require any further sperm. Concordances and discordances between attractiveness and sexual receptivity suggest that they respond to different physiological mechanisms.     

Fluctuating asymmetries and advertisement call variation in the cricket frog, Acris crepitans

We used an anuran acoustic communication system to test a predictionof the "fluctuating asymmetries/good genes" hypothesis thatfemales prefer more symmetric mates because symmetry indicatesgenetic quality. Mate preferences of female cricket frogs (Acriscrepitans) can be influenced by three call characters: dominantfrequency, numbers of pulses per call, and number of pulse groupsper call. We tested the hypothesis that these preferences resultin females preferring more symmetric males. We measured fluctuatingasymmetries of characters not involved with the communicationsystem (head and tibia), and those involved in signal production(laryngeal characters) and signal reception (aural characters).We determined whether the asymmetries in these characters wererelated to the three variables that enhance call attractiveness.Most of the multiple regression models showed no significantassociation between the fluctuating asymmetries of charactersand any of the calls. The regression of head and tibia fluctuatingasymmetry on pulse number was significant, but partial regressioncoefficients revealed that more pulses were associated witha more symmetric head length and a less symmetric tibia length.Our findings provide little or no support for the fluctuatingasymmetries/good genes hypothesis. We emphasize, however, thatthis hypothesis should not be abandoned based on negative resultsof a single study, but deserves further scrutiny.     

Is there a role for amplifiers in sexual selection?

The amplifier hypothesis states that selection could favour the evolution of traits in signallers that improve the ability of receivers to extract honest information from other signals or cues. We provide a formal definition of amplifiers based on the receiver's mechanisms of signal perception and we present a game-theoretical model in which males advertise their quality and females use sequential-sampling tactics to choose among prospective mates. The main effect of an amplifier on the female mating strategy is to increase her mating threshold, making the female more selective as the effectiveness of the amplifier increases. The effects of the amplifier on male advertising strategy depends both on the context and on the types of the amplifier involved. We consider two different contexts for the evolution of amplifiers (when the effect of amplifiers is on signals and when it is on cues) and two types of amplifiers (the ‘neutral amplifier’, when it improves quality assessment without altering male attractiveness, and the ‘attractive amplifier’, when it improves both quality assessment and male attractiveness). The game-theoretical model provides two main results. First, neutral and attractive amplifiers represent, respectively, a conditional and an unconditional signalling strategy. In fact, at the equilibrium, neutral amplifiers are displayed only by males whose advertising level lays above the female acceptance threshold, whereas attractive amplifiers are displayed by all signalling males, independent of their quality. Second, amplifiers of signals increase the differences in advertising levels between amplifying and not-amplifying males, but they decrease the differences within each group, so that the system converges towards an ‘all-or-nothing’ signalling strategy. By applying concepts from information theory, we show that the increase in information transfer at the perception level due to the amplifier of signals is contrasted by a decrease in information transfer at the emitter level due to the increased stereotypy of male advertising strategy.     

Fisherian flies: benefits of female choice in a lekking sandfly

We experimentally investigated the fitness consequences of female mate choice in order to test the relative importance of three competing but non-exclusive hypotheses for the maintenance of pronounced female mating preferences on leks: that females benefit directly; that they gain indirect Fisherian benefits by producing more attractive sons; or that they benefit indirectly because preferred males possess ''good genes'' that confer increased viability on their sons and daughters. We allowed lekking female sandflies, Lutzomyia longipalpis, to choose between males of varying attractiveness to females, and monitored the consequences for their own survival and reproductive success as well as for their offspring. In contrast to the predictions of the direct-benefits model, we found no clear sire effect on the fecundity or survival of the females themselves; females mating with more attractive males did survive longer after oviposition, but never long enough to undertake a second batch of egg laying. We also found no evidence that females gained good-genes benefits in terms of enhanced offspring survival. However, we did find that generally attractive males fathered sons who were then chosen when they in turn formed leks. Although not completely precluding other benefits, our results indicate that Fisherian benefits are at least partly responsible for maintaining female choice at L. longipalpis leks. These findings indicate the importance of testing all putative benefits concurrently in exploring the maintenance of female mate choice.     

Energetic costs of size and sexual signalling in a wolf spider

A prerequisite for honest handicaps is that there are significant condition-dependent costs in the expression of sexual traits. In the wolf spider Hygrolycosa rubrofasciata (Ohlert), sexual signalling (drumming) is costly in terms of increased mortality. Here we investigated whether this mortality may be caused by increased energy expenditure. During sexual signalling, metabolic rate was 22 times higher than at rest and four times higher than when males were actively moving. Metabolic rate per unit mass was positively related to absolute body mass during sexual signalling but not during other activities. This positive relationship is novel to any studies of metabolic rates. Indeed, it seems that the largest males can drum only 12 times per minute before reaching the maximum sustainable metabolic rate, whereas the smallest males may drum up to 39 times per minute. However, there is no relationship between body mass and drumming rate, indicating that larger males are able to compensate for the higher cost of drumming. There was a quadratic relationship between relative abdomen mass and overall body mass, which may provide a partial explanation for the increased energy expenditure of largest males while drumming. Altogether, our results indicate that sexual signalling is highly energetically demanding, which may be the main reason for the honesty of signalling in this species. In addition, the energetic costs are surprisingly strongly size dependent, which may compensate any disadvantage of small male size.