The Cost of Being Bright

Male ornaments are predicted to be a signal of the genetic quality of a male but often the relationship between ornaments and the physiological performances of males is unclear. Males of Metichnogaster cillipennis, a hover-wasp, during their courtship flights send luminous signals to females by using two abdominal reflectors. Malpighian tubules, an essential organ in the physiology of insects, participate in the building of the larger of these reflectors but at a physiological cost. By evaluating the capacity of light reflection from males during their courtship, females can assess directly the physiological quality of males.     

Incidental Sanctions and the Evolution of Direct Benefits

Recent research has shown that a variety of traits that increase male success in mating and sperm competition can impose costs on females, resulting in antagonistic coevolution between the sexes. Yet, in many animals, females are known to receive direct benefit from their mates, including many in which female multiple mating results in intense sperm competition among males. The most common explanation for the evolution of male‐provided direct benefits is pre‐mating female choice based on benefit quality. This explanation is insufficient, however, for those direct benefits that females cannot directly assess prior to mating. Given that intrasexual selection will often favor male traits that increase female mating costs, many types of direct benefits can thus be difficult to explain. In this paper, we review four additional hypotheses for the evolution of male‐provided direct benefits, and present a fifth hypothesis that has received little attention. This latter hypothesis proposes that selection often favors female reproductive tactics that are conditional upon the past costs and benefits of mating. These conditional female reproductive tactics should evolve because the quality of the benefit provided by a previous mate can change the costs and benefits of alternative reproductive decisions. Furthermore, many of the conditional reproductive tactics we might expect females to express should incidentally penalize males which provide lower quality direct benefits. These conditional reproductive tactics may thus play an important role in determining whether females incur costs or receive benefits from their mates. In addition to favoring the evolution of direct benefits, we argue that conditional female reproductive tactics may also favor reliable signaling of benefit quality. The most common explanation for reliable signaling is the handicap mechanism, which proposes that differential costs of signaling prevent low quality males from deceptively producing attractive signals. For direct benefits, however, there is a second type of deception: males which produce attractive signals and can afford to provide high quality direct benefits may choose to cheat on the advertised benefit. The handicap mechanism does nothing to prevent cheating on direct benefits by males which can afford to produce attractive signals, and is thus insufficient for ensuring reliable signaling of benefit quality. In contrast, conditional female reproductive tactics that incidentally penalize low benefit males should also penalize males which cheat on the benefits advertised by their signals.     

A two-sex life history model of handicap signaling

The males of many species (such as peacocks) develop excessively large traits, which appear to interfere with their agility and hence survival probability. Moreover, it is also observed that females seem to prefer mating males with such clumsy traits. Zahavi (1975) proposed a handicap theory to explain this phenomenon, suggesting that this trait/preference interaction is a way in which strong males can signal their viability by yielding a handicap in terms of a clumsy trait size. This paper presents a two-sex model of selfish genes that generates this particular male-female interaction, and characterizes the conditions behind a handicap equilibrium. We first show the female dominance result of Bateman (1948) in this two-sex model, and then specify the relevant equilibrium conditions, including the incentive compatibility condition for females, the individual rationality condition for males, and the stability condition of population composition. Identifying these conditions helps us understand the various features of the searching/signaling of sex selection in evolution.     

Male chemical cues as reliable indicators of infection in the wolf spider Schizocosa ocreata

Sexual signals can convey important information about mate quality, such as critical information about a signaler's health status, helping an individual to avoid infected or immunocompromised conspecifics. Chemical signals are especially important in this context, because they represent an honest and dynamic signaling modality that receivers can use to make updated mate choice decisions to avoid compromising their own health. In this study, we investigated the viability of male chemical cues in the wolf spider Schizocosa ocreata as a reliable indicator of health status. Using video playback with images of an average male that is simultaneously paired with male cuticular compounds on filter paper, we show that females are more receptive to videos paired with cues from control males rather than infected males. We also show that these cuticular compounds can be isolated and retain similar female behavioral responses when extracted with a nonpolar solvent, suggesting that these cuticular compounds may not be just complex hydrocarbons, but a combination of cuticular compounds. This is the first evidence for female discrimination and recognition of male chemical cues in this species, which opens up important new avenues of research in a well‐studied species with complex multimodal signaling.     

Evolution of risk‐taking during conspicuous mating displays

In many species, males possess conspicuous characteristics to attract females. These traits often attract predators as well, and males thus may have to balance the conspicuousness of their signals in relation to the prevailing predation risk. Here we develop a theoretical model of optimal signaling and risk‐taking behavior for males differing in the attractiveness of their signals. All else being equal, more attractive males should behave more cautiously. Yet this prediction may drastically change if males differ in any additional characteristic, especially if basal mortality rate or signaling costs are higher or if the vulnerability to predators is lower for attractive males. A key insight from our model is that male competition will create a positive feedback so that selection on male risk‐taking strategies is acting in opposite directions. If selection acts on one male type to behave more cautiously, this will strengthen selection on males of the other types to take higher risks and vice versa. Our results further demonstrate that the asset‐protection principle, which states that individuals with higher future expectations should behave more cautiously, may often be violated. We also offer an alternative to the handicap principle explaining the often found positive association between male ornamentation and viability: attractive males may simply behave more cautiously.     

Fitness cost of pheromone production in signaling female moths

A secondary sexual character may act as an honest signal of the quality of the individual if the trait bears a cost and if its expression is phenotypically condition dependent. The cost of increasing the trait should be tolerable for individuals in good condition but not for those in a poor condition. The trait thus provides an honest signal of quality that enables the receiver to choose higher quality mates. Evidence for sex pheromones, which play a major role in shaping sexual evolution, inflicting a signaling cost is scarce. Here, we demonstrate that the amount of the major component of the pheromone in glands of Lobesia botrana (Lepidoptera) females at signaling time was significantly greater in large than in small females, that male moths preferred larger females as mates when responding to volatile signals, and small virgin females, but not large ones, exposed to conspecific pheromone, produced, when mated, significantly fewer eggs than nonexposed females. The latter indicates a condition-dependent cost of signaling. These results are in accordance with the predictions of condition-dependent honest signals. We therefore suggest that female signaling for males using sex pheromones bears a cost and thus calling may serve as honest advertisement for female quality.     

SEXUAL SELECTION IN A WOLF SPIDER: MALE DRUMMING ACTIVITY,BODY SIZE,AND VIABILITY

Females are often believed to actively choose highly ornamented males (males with extravagant morphological signals or intense sexual display), and ornaments should be honest signals of male viability. However, this belief is relying only on some pieces of empirical evidence from birds. Our study reports active female choice on sexual display that indicates male viability in spiders. We established trials in which we studied female choice in relation to male courtship drumming activity and body size. Females chose the most actively drumming males as mating partners, but the body size of the males did not seem to be selected. Male drumming activity turned out to be a good predictor of male viability, whereas male viability was independent of male body mass. Our results suggest that by actively choosing mates according to male drumming performance, but independently of male body mass, females are preferring viable males as mates. Because Hygrolycosa rubrofasciata males do not provide obvious direct benefits to their offspring, females may gain some indirect benefits; offspring may have higher chance of survival, or the offspring may inherit the attractiveness of their father.     

Sizing‐up the Competition: Factors Modulating Male Display Behavior During Mate Competition

When competing for mates, males of many species use cues from their rivals to evaluate their chances of success. Signaling behavior is a vital component of male–male contests and courtship, and may inform males of a rival's quality or intentions. We used eastern fence lizards (Sceloporus undulatus) to investigate how the time a male spent signaling during mate competition is influenced by his quality, his rival's quality relative to his own, and the value of a contested female. Furthermore, we examined how a male's behavioral response to a competitor's signals would be mediated by his relative quality. We simulated natural encounters by allowing two males to compete over a single female in the laboratory. We measured the time males spent performing two types of displays (pushups and shudders) and categorized male behavioral responses to rival pushup and shudder displays. Time spent signaling was not related to a male's absolute quality (body and head size, condition, and badge sizes), or his quality relative to that of his rival, although males did spend more time performing pushups when competing over females in better condition. Male behavior was also influenced by his rival's signals, such that males of relatively lower quality than their opponents were more likely to aggressively respond to rival pushups and shudders. We discuss these results with respect to the evolution and function of signaling behavior in courtship and male–male contests.     

False promises: females spurn cheating males in a field cricket

Females commonly prefer to mate with males that provide greater material benefits, which they often select using correlated male signals. When females select higher-benefit males based on correlated signals, however, males can potentially deceive females by producing exaggerated signals of benefit quality. The handicap mechanism can prevent lower-quality males from producing exaggerated signals, but cannot prevent cheating by higher-quality males that choose to withhold the benefit, and this poses a major problem for the evolution of female choice based on direct benefits. In a field cricket, Gryllus lineaticeps, females receive seminal fluid products from males with preferred songs that increase their fecundity and lifespan. We tested the hypothesis that female behaviour penalizes males that provide lower-quality benefits. When females were paired with males that varied in benefit quality but had experimentally imposed average songs, they were less likely to re-mate with males that provided lower-quality benefits in the initial mating. This type of conditional female re-mating may be a widespread mechanism that penalizes males that cheat on direct benefits.     

The scent of senescence: sexual signalling and female preference in house mice

Sexual signals are expected to be costly to produce and maintain, thus ensuring that only males in good condition can sustain their expression at high levels. When males reach senescence they lose physiological function and condition, which could constrain their ability to invest in costly sexual signals, decreasing their attractiveness to mates. Furthermore, females may have evolved mating preferences that cause avoidance of senesced males to enhance fertilization success and viability of offspring. Among mammals, the size of antlers and other weapons can decrease with senescence, but changes in olfactory sexual signals have been largely unexplored. We examined changes in olfactory signals with senescence in house mice (Mus musculus domesticus), where males excrete volatile and involatile molecules in scent marks that elicit behavioural and priming responses in females. Compared to middle-aged males, the urine of senesced males contained a lower concentration of involatile signalling proteins (major urinary proteins or MUPs), and associated volatiles that bind to these proteins. The reduced intensity of male scent will affect the longevity of scent signals deposited in the environment and, accordingly, females were less attracted to urine from senesced males deposited 12 h previously. Females also discriminated against senesced males encountered behind a mesh barrier. These results reveal that investment in olfactory signalling is reduced during senescence and suggest that senesced males and their scent may be less attractive to females.     

Contrasting patterns of selection on the size and coloration of a female plumage ornament in common yellowthroats

Females often possess ornaments that appear smaller and duller than homologous traits in males. These ornaments may arise as nonfunctional by‐products of sexual selection in males and cause negative viability or fecundity selection in females in proportion to the cost of their production and maintenance. Alternatively, female ornaments may function as signals of quality that are maintained by sexual or social selection. In a 4‐year study of 83 female common yellowthroats (Geothlypis trichas) and their 222 young, we found strong viability and fecundity selection on the yellow bib, a carotenoid‐based plumage ornament that is a target of sexual selection in males. Females with larger bibs were older, larger and more fecund than females with smaller bibs. However, bib size positively covaried with bib total brightness and carotenoid chroma, aspects of bib coloration that were under negative viability and fecundity selection. Females with more colourful bibs laid fewer eggs in their first clutch, were more likely to suffer total brood loss due to predation and were less likely to return to the study area. Selection against bib coloration limits the value of bib size as a quality indicator in females and may constrain the elaboration of bib attributes in males.     

Responses to Femoral Gland Secretions by Visually Adept Male and Female Collared Lizards

Although visually transmitted social signals are well documented in many diurnal iguanians, including collared lizards, secretory femoral glands also suggest a role of chemical signals in intraspecific communication. We conducted laboratory trials to test the extent to which male and female collared lizards responded by tongue‐flicking femoral gland secretions, neutral (water), and odoriferous (cologne) control substances, males distinguished self‐secretions from those produced by unfamiliar rival males, and females distinguish secretions from unfamiliar vs. familiar males. Both males and females spent similar amounts of time in four arena quadrants each with a Petri dish treated with one of the four test compounds. Males gave more tongue flicks/trial to secretions produced by unfamiliar rivals and cologne than they tongue‐flicked their own secretions and water. By contrast, the number of tongue flicks by females on control substances and familiar and unfamiliar males was similar. Results support the hypothesis that femoral gland secretions function in intrasexual signaling among male collared lizards, perhaps allowing them to distinguish unfamiliar rivals. Females tongue‐flicked secretions from familiar and unfamiliar males with similar frequency that was high relative to that of males, suggesting a possible role of secretions in assessment of males. Ours is the first evidence of a signaling role of femoral gland secretions in collared lizards and adds to a growing body of evidence that chemical signaling has evolved in diurnal lizards that also have highly developed visual‐based signaling.     

Offspring viability benefits but no apparent costs of mating with high quality males

Traditional models of sexual selection posit that male courtship signals evolve as indicators of underlying male genetic quality. An alternative hypothesis is that sexual conflict over mating generates antagonistic coevolution between male courtship persistence and female resistance. In the scarabaeine dung beetle Onthophagus taurus, females are more likely to mate with males that have high courtship rates. Here, we examine the effects of exposing females to males with either high or low courtship rates on female lifetime productivity and offspring viability. Females exposed to males with high courtship rates mated more often and produced offspring with greater egg-adult viability. Female productivity and lifespan were unaffected by exposure to males with high courtship rates. The data are consistent with models of sexual selection based on indirect genetic benefits, and provide little evidence for sexual conflict in this system.     

The evolution of postpairing male mate choice

An increasing number of empirical studies in animals have demonstrated male mate choice. However, little is known about the evolution of postpairing male choice, specifically which occurs by differential allocation of male parental care in response to female signals. We use a population genetic model to examine whether such postpairing male mate choice can evolve when males face a trade‐off between parental care and extra‐pair copulations (EPCs). Specifically, we assume that males allocate more effort to providing parental care when mated to preferred (signaling) females, but they are then unable to allocate additional effort to seek EPCs. We find that both male preference and female signaling can evolve in this situation, under certain conditions. First, this evolution requires a relatively large difference in parental investment between males mated to preferred versus nonpreferred females. Second, whether male choice and female signaling alleles become fixed in a population versus cycle in their frequencies depends on the additional fecundity benefits from EPCs that are gained by choosy males. Third, less costly female signals enable both signaling and choice alleles to evolve under more relaxed conditions. Our results also provide a new insight into the evolution of sexual conflict over parental care.     

Male house mice evolving with post-copulatory sexual selection sire embryos with increased viability

Although mating is costly, multiple mating by females is a taxonomically widespread phenomenon. Theory has suggested that polyandry may allow females to gain genetic benefits for their offspring, and thus offset the costs associated with this mating strategy. For example, the good sperm hypothesis posits that females benefit from mating multiply when genetically superior males have increased success in sperm competition and produce high quality offspring. We applied the powerful approach of experimental evolution to explore the potential for polyandry to drive evolutionary increases in female fitness in house mice, Mus domesticus. We maintained polygamously mated and monogamously mated selection lines of house mice for 14 generations, before determining whether selection history could account for divergence in embryo viability. We found that males from lineages evolving with post-copulatory sexual selection sire offspring with increased viability, suggesting that polyandry results in the production of higher quality offspring and thus provides long-term fitness benefits to females.     

Female preference for complex/novel signals in a spider

Identifying the various factors that influence complex signalevolution is a difficult task, yet it is fundamental to understandingthe evolution of animal communication. Here we explore the evolutionof complex courtship signaling by taking advantage of a systemin which sexual selection on male courtship traits has driventhe diversification of geographically isolated populations ofthe jumping spider Habronattus pugillis Griswold. Using 2 populations(Santa Rita [SR] and Atascosa [AT]) in which SR females showxenophilic mating preferences for foreign (AT) over local males(SR), we examine the mechanisms driving this preference. BothAT and SR males produce multimodal signals (visual + seismic),and while SR and AT signals share certain seismic components,AT seismic signals are more complex and contain novel components.We conducted mate choice trials where SR females were presentedwith AT or SR males that were either muted or nonmuted. SR femalespreferred to mate and mated more quickly with foreign AT malesover local SR males only if AT males could produce seismic signals(nonmuted treatment). In addition, we found that SR femalesspent a higher proportion of time attentive to foreign AT malesonly if they could produce seismic signals. This evidence suggeststhat SR females have a bias for complex and/or novel forms ofseismic signals.     

Chemoreception, symmetry and mate choice in lizards

Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual's developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males' scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.     

Using Protein Motion to Read,Write, and Erase Ubiquitin Signals

Eukaryotes use a tiny protein called ubiquitin to send a variety of signals, most often by post-translationally attaching ubiquitins to substrate proteins and to each other, thereby forming polyubiquitin chains. A combination of biophysical, biochemical, and biological studies has shown that complex macromolecular dynamics are central to many aspects of ubiquitin signaling. This review focuses on how equilibrium fluctuations and coordinated motions of ubiquitin itself, the ubiquitin conjugation machinery, and deubiquitinating enzymes enable activity and regulation on many levels, with implications for how such a tiny protein can send so many signals.     

Sexual selection and fertility

Genetic models are analyzed in which sexual selection is combined with fertility selection. In these models, the sexual selection acts on males, the fertility selection on either males, females or both sexes. The phenotypes thus selected may be determined either by dominant and recessive alleles or by each homozygous and heterozygous genotype. Polymorphisms of dominant and recessive phenotypes can be maintained in equilibrium by a balance between sexual and fertility selection. Generally fertility selection has a greater effect than viability selection in determining the point of equilibrium. The dominant phenotype is maintained at a lower frequency when at a fertility disadvantage than when at a viability disadvantage. When about 20% or more of the females mate preferentially, the models show that equilibria will be established at very different frequencies depending on whether fertility selection acts on males, females or both sexes. These results, applied to data of preferential mating of melanic two-spot ladybirds, predict differences in fertility which can be use to test the models. Symmetric models of preferences for each genotype also give rise to polymorphisms if the heterozygotes obtain an overall advantage.     

Tactic-dependent plasticity in ejaculate traits in the swordtail Xiphophorus nigrensis

In species with alternative reproductive tactics, males that sneak copulations often have larger, higher quality ejaculates relative to males that defend females or nest sites. Ejaculate traits can, however, exhibit substantial phenotypic plasticity depending on a male's mating role in sperm competition, which may depend on the tactic of his competitor. We tested whether exposure to males of different tactics affected sperm number and quality in the swordtail Xipophorus nigrensis, a species with small males that sneak copulations and large males that court females. Sperm swimming speed was higher when the perceived competitor was small than when the competitor was large. Plasticity, however, was only exhibited by small males. Sperm number and viability were invariant between social environments. Our results suggest sperm quality is role-dependent and that plastic responses to the social environment can differ between male reproductive tactics.