Female-specific color is a signal of quality in the striped plateau lizard (Sceloporus virgatus)

Recent theoretical and empirical studies confirm that male matechoice and/or female–female mate competition can be expressedin the absence of sex-role reversal. Such reproductive patternsmay select for the evolution of female traits that indicatefemale phenotypic or genotypic quality among non–role-reversedspecies. Although attention to the evolution and function offemale ornaments is increasing, additional focus is needed onfemale-specific ornaments (those not expressed in conspecificmales) and on nonavian systems in order to gain a broad understandingof how selection acts directly on ornamentation of female animals.In the striped plateau lizard, Sceloporus virgatus, only femalesdevelop orange throat patches during the reproductive season.The color peaks in expression near the time of ovulation andappears to stimulate male courtship. Here, I examine whetherthis female-specific ornament can be used by males to reliablyassess female phenotypic quality. Using multivariate regressionanalyses, I show that the area of the orange color patch predictsbody condition and mite load, the chroma (i.e., saturation)of the color patch predicts body size, and both patch area andchroma reliably predict average egg mass. Thus, female reproductivecolor may function as a condition-dependent signal, indicatingphenotypic quality to potential mates.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

Male mate choice and the potential for complex mating dynamics in the tree lizard (Urosaurus ornatus)

A growing body of literature is recognizing that males may also play a role in the mating process by behaving non‐randomly toward potential female mates during courtship. In numerous species, discrete color polymorphisms in males are inferred to represent alternative mating tactics, which often correspond with concomitant asymmetries in ecology and behavior. In terms of their mating behavior, these ecological outcomes of a color polymorphism should affect a morph's likelihood and frequency of encountering females in a population, possibly favoring the evolution of morph‐specific mating preferences. Knowledge of how male morphs contribute to a species’ overall mating dynamics will improve our understanding of how sexual selection shapes phenotypic diversity in color polymorphic systems. We conducted a mate choice experiment to evaluate the extent and morph specificity of non‐random mating preferences by male ornate tree lizards, Urosaurus ornatus. We observed the behavior of blue and yellow males in an experimental arena in response to a choice between an orange or yellow female. We found that blue males preferred yellow females over orange females, and although yellow males visited females more often than blue males overall, their attention was not morph‐specific. Given male morph differences in choosiness, and their differences in social dominance, we conclude that female throat color may be partly under sexual selection in U. ornatus. However, a lack of concordance between male and female mating preferences (drawn from an earlier study) suggests that overall mating dynamics may serve to maintain, rather than enhance, color morph differences in this species.     

Field evidence challenges the often‐presumed relationship between early male maturation and female‐biased sexual size dimorphism

Female‐biased sexual size dimorphism (SSD) is often considered an epiphenomenon of selection for the increased mating opportunities provided by early male maturation (i.e., protandry). Empirical evidence of the adaptive significance of protandry remains nonetheless fairly scarce. We use field data collected throughout the reproductive season of an SSD crab spider, Mecaphesa celer, to test two hypotheses: Protandry provides fitness benefits to males, leading to female‐biased SSD, or protandry is an indirect consequence of selection for small male size/large female size. Using field‐collected data, we modeled the probability of mating success for females and males according to their timing of maturation. We found that males matured earlier than females and the proportion of virgin females decreased abruptly early in the season, but unexpectedly increased afterward. Timing of female maturation was not related to clutch size, but large females tended to have more offspring than small females. Timing of female and male maturation was inversely related to size at adulthood, as early‐maturing individuals were larger than late‐maturing ones, suggesting that both sexes exhibit some plasticity in their developmental trajectories. Such plasticity indicates that protandry could co‐occur with any degree and direction of SSD. Our calculation of the probability of mating success along the season shows multiple male maturation time points with similar predicted mating success. This suggests that males follow multiple strategies with equal success, trading‐off access to virgin females with intensity of male–male competition. Our results challenge classic hypotheses linking protandry and female‐biased SSD, and emphasize the importance of directly testing the often‐assumed relationships between co‐occurring animal traits.     

Sperm competition generates evolution of increased paternal investment in a sex role‐reversed seed beetle

When males provide females with resources at mating, they can become the limiting sex in reproduction, in extreme cases leading to the reversal of typical courtship roles. The evolution of male provisioning is thought to be driven by male reproductive competition and selection for female fecundity enhancement. We used experimental evolution under male‐ or female‐biased sex ratios and limited or unlimited food regimes to investigate the relative roles of these routes to male provisioning in a sex role‐reversed beetle, Megabruchidius tonkineus, where males provide females with nutritious ejaculates. Males evolving under male‐biased sex ratios transferred larger ejaculates than did males from female‐biased populations, demonstrating a sizeable role for reproductive competition in the evolution of male provisioning. Although larger ejaculates elevated female lifetime offspring production, we found little evidence of selection for larger ejaculates via fecundity enhancement: males evolving under resource‐limited and unlimited conditions did not differ in mean ejaculate size. Resource limitation did, however, affect the evolution of conditional ejaculate allocation. Our results suggest that the resource provisioning that underpins sex role reversal in this system is the result of male–male reproductive competition rather than of direct selection for males to enhance female fecundity.     

The Haremic Mating System and Mate Choice in the Wide-Eyed Flounder, Bothus podas

The social structure and reproductive behaviour of the wide-eyed flounder, Bothus podas, was studied in the coastal waters around the Azorean Islands. Both sexes are territorial throughout the year. Adult males defend large territories, which include several smaller female territories. Intraspecific agonistic behaviour was frequent and differed between sexes: males were more aggressive towards other males, while females were only aggressive towards each other and juveniles. During the reproductive season and only at dawn, territorial males court and mate successively with females in their territories, and females seem to show mating fidelity to their dominant male. Such territoriality and mating patterns indicate a haremic social system in the wide-eyed flounder. In order to identify potential factors influencing female mate choice acting on this haremic system, we examined male mating success and some of its potential correlates. We found no evidence for female preference for any of the males' physical or territory characteristics. However, courtship effort was strongly correlated with the total number of attempted and successful spawnings, indicating that females seem to mate preferentially with males that court them more vigorously. Thus, our data suggest that courtship plays an important role in determining male mating success in the wide-eyed flounder and, that it may possibly serve as an honest indicator of male `quality' for female choice.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Non‐Conceptive Sexual Behavior in Spiders: A Form of Play Associated with Body Condition,Personality Type,and Male Intrasexual Selection

In the socially polymorphic spider Anelosimus studiosus, males mature early in the reproductive season and recruit to the webs of juvenile females and guard them until they mature. During the period before females mature, males and females engage in repeated bouts of non‐conceptive (play) sexual behavior, where the pair courts and engages in mock copulation; both males and females gain performance‐enhancing experience via these encounters. In this study, we examined the factors that underlie individual variation in the tendency to engage in non‐conceptive mating and determine whether it impacts male–male competition for females. We found that docile females, being less resistant to mating in general, are more likely to accept male courtship and non‐conceptive copulation as juveniles. Personality type influenced the exhibition of non‐conceptive sexual behavior in males as well. High body condition males of the aggressive phenotype were more likely to engage in non‐conceptive sexual behavior than males with lower body condition. Body condition did not influence docile males’ propensity to engage in non‐conceptive sexual behavior, but female size did. Docile males engaged in more non‐conceptive sexual displays with larger females. Engaging in non‐conceptive sexual displays negatively impacted male performance in staged male–male contests for access to females. This cost was greatest for males of the aggressive phenotype, which are otherwise favored in male–male contests. Our findings indicate expression of non‐conceptive sexual displays is linked to personality and results in reproductive performance trade‐offs for male A. studiosus.     

Physiological approach to the onset of receptivity in female Acheta domesticus : I. Role of the corpora allata and ovaries

During the 32 hr following the imaginal moult, all female Acheta domesticus actively or passively refuse male courtship; they are unreceptive. As of 32 hr, the most precocious females become receptive and accept mating. At this time, juvenile hormone (JH III) synthesized by corpora allata (CA) is already detectable in hemolymph, while ecdysteroids (synthesized by ovaries) begin increasing at 48 hr. JH III and ecdysteroid levels in hemolymph were measured by RIA. After allatectomy and/or ovariectomy, all females became receptive, thus showing that CA and/or ovaries are not essential to the onset of receptivity. However, male courtship is longer for allatectomized females; in ovariectomized females, mating is delayed.     

The mating strategy and reproductive performance of Agriphila aeneociliella (Lepidoptera: Crambidae), a new insect pest of wheat in China

1. The Eastern Grass‐veneer Agriphila aeneociliella (Eversmann) (Lepidoptera: Crambidae) is a serious stem‐feeding pest of wheat crops that has become established in China in recent years. A better understanding of the mating strategy and reproductive performance of A. aeneociliella could improve integrated pest management programmes against this newly established species by disturbing its behaviour and reducing its reproduction potential. Based on ethological and reproductive biological approaches, the mating and reproductive performances of A. aeneociliella were investigated.
2. Unlike the common nocturnal lepidopterans, the mating rhythm of A. aeneociliella moths showed a marked diurnal pattern. The female courtship rhythm and the mating rhythm reached peaks within the first 2 h after the onset of photocycle.
3. The mating success rate of monogamous pairs was 55.6%, whereas the male‐biased sex ratio (2♂:1♀) increased the mating rate (72.2%) and the female‐biased ratio (1♂:3♀) led to the lowest mating rate (27.8%).
4. Both females and males were able to mate twice. The duration of copulation decreased substantially with male mating frequency, whereas, when a previously mated female was paired with a virgin male, fecundity significantly increased. Monogamous couples who mated only once in their lives have the highest hatchability (97.13 ± 0.49%).

     

RAPID LOSS OF BEHAVIORAL PLASTICITY AND IMMUNOCOMPETENCE UNDER INTENSE SEXUAL SELECTION

Phenotypic plasticity allows animals to maximize fitness by conditionally expressing the phenotype best adapted to their environment. Although evidence for such adjustment in reproductive tactics is common, little is known about how phenotypic plasticity evolves in response to sexual selection. We examined the effect of sexual selection intensity on phenotypic plasticity in mating behavior using the beetle Callosobruchus maculatus. Male genital spines harm females during mating and females exhibit copulatory kicking, an apparent resistance trait aimed to dislodge mating males. After exposing individuals from male‐ and female‐biased experimental evolution lines to male‐ and female‐biased sociosexual environments, we examined behavioral plasticity in matings with standard partners. While females from female‐biased lines kicked sooner after exposure to male‐biased sociosexual contexts, in male‐biased lines this plasticity was lost. Ejaculate size did not diverge in response to selection history, but males from both treatments exhibited plasticity consistent with sperm competition intensity models, reducing size as the number of competitors increased. Analysis of immunocompetence revealed reduced immunity in both sexes in male‐biased lines, pointing to increased reproductive costs under high sexual selection. These results highlight how male and female reproductive strategies are shaped by interactions between phenotypically plastic and genetic mechanisms of sexual trait expression.     

The Influence of Male Body Size and Social Environment on the Mating Behavior of Phallichthys quadripunctatus (Pisces: Poeciliidae)

In some poeciliid fishes, variation in male size is accompanied by differences in mating behavior. Large males are preferred by females and perform courtship displays followed by copulatory thrusts, whereas small males perform copulatory thrusts with few or no displays. This phenomenon has been described in only a few genera and little is known about mating behavior in other poeciliids. Although Phallichthys quadripunctatus males display size dimorphism that has a genetic component, mating behavior of this species has not been documented. We conducted experiments using socially experienced and socially naive males to characterize the mating behavior of this species and to evaluate potential size-dependent differences in behavior. Males were tested with postpartum (presumably receptive) and midcycle (presumably unreceptive) females in different social environments. Whereas neither size class of P. quadripunctatus males performed courtship displays or altered behavior based on female receptivity, large males performed several reproductive behaviors more frequently than small males. This trend was repeatable and occurred in all social environments examined. Some males also attempted to mate with other males, with small males showing a greater tendency to perform this behavior than large males. The manner in which differences in reproductive activity translate into differences in reproductive success must be examined before inferring sexual selection favoring large males in P. quadripunctatus.     

Mate Preference for Novel Phenotypes: A Fresh Face Matters

Conspicuous polymorphism in sexually selected traits is usually attributed to processes such as frequency‐dependent selection that can maintain genetic variation. Recent evidence indicates that dramatic variation of male coloration in guppies (Poecilia reticulata) is promoted by a form of frequency‐dependent selection in which males bearing rare or novel color patterns achieve higher mating success than males bearing common patterns. Active female preference for unfamiliar or rare color patterns has been implicated in generating this rare‐phenotype advantage, but the behavioral processes responsible for the preference remain unclear. To determine whether familiarity that is developed over a very short timescale can lead to a rare‐male mating advantage, we measured female response to courtship by males with color patterns that were the same as or different from that of the previous male to court. Females showed two types of short‐term preference variation in this experiment. On the first trial day, females shifted their preferences on a timescale of minutes, showing strong preference for males bearing a color pattern different from that of the immediately previous male to court. Twenty‐four hours later, females were less responsive to male courtship overall, and there was no difference in females’ response to different‐ and same‐morph males. Females also preferred males with more orange coloration on both trial days, but this color preference was independent of the preference for ‘different’ color patterns. These data suggest that the behavioral process underlying rare‐male advantage in guppies is that females prefer males bearing unfamiliar color patterns and that familiarity is determined over a very short timescale.     

Seismic Signaling is Crucial for Female Mate Choice in a Multimodal Signaling Wolf Spider

Complex courtship signals can be dissected into distinct components that can either function independently or via interactions with one another. Male Rabidosa rabida wolf spiders use courtship displays that couple a seismic signal with the waving of an ornamented foreleg. While previous studies suggest that female R. rabida exhibit mate choice and that both the seismic and visual modalities are important in mating interactions, it remains unclear how variation in each component influences female mating decisions. To investigate this, we ran two separate experiments in which we manipulated (1) male diets, to induce variation in the seismic courtship signal, and (2) male foreleg color, to artificially induce variation in visual foreleg ornamentation. To determine the influence of variation in each component independently, females were paired with males in environments that allowed the detection of only the manipulated signal component (e.g. seismic signal only and visual signal only). Variability in the seismic signal alone influenced female mate choice, but variability in visual ornamentation alone did not. In a third experiment, we manipulated foreleg color and allowed it to interact with the seismic signal to determine whether inter‐signal interactions influence female mating decisions. When females were able to detect both signal components, variation in visual ornamentation did influence mate choice – females preferred ornamented males. Together, these results suggest that the seismic signal of male R. rabida is integral for female mate choice and that the components of the courtship display interact to influence female mating decisions.     

Mating plugs and sexual cannibalism in the Colombian orb‐web spider Leucauge mariana

In Leucauge orb‐web spiders, females form mating plugs, which play a part in cryptic female choice after they evaluate male performance during courtship and mating. Our aim was to assess sexual behavior and mating plug formation of Leucauge mariana from Colombia (CO). We carried out mating trials to describe in detail courtship and mating behavior of sexual pairs from the CO population, and then compared the results to previously collected data from a Costa Rican population (CR). In addition, we assessed the amount of sperm transferred during mating predict plug formation in CO pairs. All CO virgin females mated (n = 23), and mating plug formation occurred in only 11 cases (48%). In contrast, all CR virgin females mated (n = 43) and 74% formed mating plugs. None of the male courtship or mating behaviors that we measured in CO pairs predicted mating plug formation, in contrast to previous reports from CR. Mating plugs of CO consisted of a matrix of unknown composition (77.2%) and encapsulated and decapsulated sperm (22.8%). In CR, plug composition varied in color and consistency, but also comprised a matrix with encapsulated and decapsulated sperm. We observed female cannibalism of males in CO pairs, which had never been reported in CR pairs. Some female displays during courtship and mating were unique to each population. Different female preferences could explain the observed geographic differences between L. mariana in male traits and behaviors that could, in turn, promote reproductive isolation. More studies are needed to test this hypothesis.     

Short‐Term Nutritional Limitation Affects Mating Behaviour and Reproductive Output in Dwarf Spiders

Mating decisions can vary considerably depending on individual experience, mate availability and nutritional status. Here, we applied short‐term dietary restrictions to adult female spiders that were well fed during the juvenile stage in an effort to understand whether and how brief periods of food shortage can influence male and female mating decisions and mating behaviour. To assess whether responses vary between closely related species, we conducted the same experiment on the dwarf spiders Oedothorax retusus and O. apicatus. During courtship and mating, males of both species offer secretion to females from glandular tissue in their prosoma. Females were subject to food shortage over a period of 3 wks (‘low‐diet’ treatment, LD) or fed regularly (‘high‐diet’ treatment, HD). We compared courtship probability, mating probability/behaviour, and reproductive output between dietary groups and species. In both Oedothorax species, females in the LD treatment were less likely to mate and more aggressive towards males. Furthermore, LD females produced egg sacs that were significantly lighter than were those of the HD females. Effects of food deprivation on copulation duration, gustatory behaviour and oviposition latency differed between species. Our study shows that short periods of dietary restriction during the adult stage can strongly affect mating behaviour and reproductive output with differences between closely related species.     

Hunting and Nesting Phenology Influence Gobbling of Wild Turkeys

Wild turkeys (Meleagris gallopavo) use a polygynous mating system whereby males engage in multiple courtship behaviors, including vocalizations (gobbling) to attract females and compete with other males for breeding opportunities. Males must balance the risk of courtship behaviors with the reproductive potential of each courtship behavior. Male turkeys are primarily hunted during the reproductive period, so the associated risk of courtship behaviors is increased. Many state agencies attempt to set hunting season frameworks that maximize hunter satisfaction by allowing hunting when gobbling activity is greatest and most females are theoretically incubating nests, but the relationship between gobbling activity and nesting phenology is unclear. We used autonomous recording units and global positioning system transmitters to monitor gobbling activity by male turkeys and reproductive behaviors of female turkeys in the Piedmont region of Georgia, USA. We used 13,177 gobbles, behavioral data from 82 females during the reproductive season, and daily estimates of harvest of males by hunters to examine relationships between daily gobbling activity, cumulative removal of males, and reproductive behaviors (laying, incubating) of females during 2017–2018. We observed a weak negative relationship between daily gobbling activity and gobbling activity the following day. As the reproductive season progressed, gobbling activity decreased. As the proportion of females engaged in laying or incubating behaviors increased, expected daily gobbling activity increased. Conversely, we observed that hunting and removal of males had a negative effect on daily gobbling activity, and this effect was disproportionately greater than the positive effect of female reproductive behaviors. Our findings suggest that hunting and removal of males are important determinants of gobbling activity, and that corresponding reductions in gobbling activity may have mediating effects on the mating system of wild turkeys. © 2019 The Wildlife Society.     

Female mate preference in an invasive phytopathogen vector: how learning may influence mate choice and fecundity in Diaphorina citri

In some arthropods, females learn in the context of reproduction to refine their mate choices and avoid males displaying traits associated with impotency. Previous studies have shown that males of the Asian citrus psyllid, Diaphorina citriKuwayama (Hemiptera: Psyllidae), associate female odor with copulatory rewards. However, it is unclear whether females similarly learn about male traits. We compared mate choice in females previously mated to either blue or orange males and found that females may associate male color with reproductive success, and avoided blue males after previous experience. Orange males mated more frequently than blue males and appeared to be more sexually aggressive in their mating attempts. In addition, females mated to orange males laid approximately twice as many eggs as those mated to blue males. We dissected male psyllids and measured the size of their reproductive organs to determine whether abdominal color was associated with reproductive development. Our morphometric analysis indicated that blue males may not be physiologically immature compared with orange males. Rather, blue males displayed larger testes and seminal vesicles than orange males, suggesting that differences in reproductive output may be behavioral in nature. Based on our data, we suggest that females learn about the appearance and mating behaviors of blue and orange males from early mating experiences, and subsequently avoid blue males in future matings. Although the roles of blue males within a population are unclear, it is possible that blue morphs may specialize in other functions such as dispersal.     

Melodic males and flashy females: Geographic variation in male and female reproductive behavior in red‐eyed treefrogs (Agalychnis callidryas)

Geographic variation in courtship behavior can affect reproductive success of divergent phenotypes via mate choice. Over time, this can lead to reproductive isolation and ultimately to speciation. The Neotropical red‐eyed treefrog (Agalychnis callidryas) exhibits high levels of phenotypic variation among populations in Costa Rica and Panama, including differences in color pattern, body size, and skin peptides. To test the extent of behavioral premating isolation among differentiated populations, we quantified male advertisement calls from six sites and female responses to male stimuli (acoustic and visual signals) from four sites. Our results show that both male advertisement calls and female behavior vary among populations: Discriminant function analyses can predict the population of origin for 99.3% ± 0.7 of males based on male call (dominant frequency and bandwidth) and 76.1% ± 6.6 of females based on female response behavior (frequency and duration of visual displays). Further, female mate choice trials (n = 69) showed that population divergence in male signals is coupled with female preference for local male stimuli. Combined, these results suggest that evolved differences among populations in male call properties and female response signals could have consequences for reproductive isolation. Finally, population variation in male and female behavior was not well explained by geographic or genetic distance, indicating a role for localized selection and/or drift. The interplay between male courtship and female responses may facilitate the evolution of local variants in courtship style, thus accelerating premating isolation via assortative mating.     

Aggressive Behavior and Courtship Rejection in Brightly and Plainly Colored Female Keeled Earless Lizards (Holbrookia propinqua)

Relationships between bright secondary sexual coloration and behavior were studied in female Holbrookia propinqua, which develop striking orange and yellow colors during the breeding season. In tethered introduction studies, brightly colored females performed aggressive courtship rejection behaviors toward conspecific males; plainly colored females were not aggressive toward males, but attempted to avoid them. Responses of females of the two color patterns toward conspecific females of both color phases were not detectably different. Experimental introductions of lizards with coloration modified by paint showed that females of both color patterns recognize any other lizard bearing the bright female colors as female, regardless of actual sex. Both the orange and yellow components were shown to contribute to sex recognition. The yellow component alone allowed accurate sex identification, but only half the females responded to males painted with only the orange female component as if they were females. Because females did not behave differently toward other females on the basis of coloration, the hypothesis that bright coloration evolved as an adaptive signal between females is rejected. The dark ventrolateral stripes of male and plainly colored females did not appear to affect intraspecific social responses by females.