The function of prolonged copulations in Enchenopa treehoppers (Hemiptera: Membracidae).

Copulations are very brief in many species, sometimes taking only seconds, but in other species, they can be quite prolonged. Potential explanations for prolonged copulations include time requirements for the transfer of sperm and/or other ejaculate substances. Ejaculate substances could function to regulate female receptivity to subsequent matings, provide nutritional nuptial gifts, or hasten egg oviposition at a potential survival cost to the female. We investigated prolonged copulation in a member of the Enchenopa binotata complex of treehoppers (Hemiptera: Membracidae), in which females rarely remate and copulation can last several hours. We assigned females to treatments in which we interrupted copulation at different times. We also included a control where copulation was not interrupted. We found that females that experienced shorter copulations were more likely to be subsequently receptive to an attractive male. We also found that few females produced offspring when they engaged in short copulations compared to those with longer copulations. We did not find any differences in female survival. Our results support the sperm transfer and receptivity regulation hypotheses. We discuss potential reasons for why these processes should take so long in a species with low female remating.

Queen-Worker Conflict over Acceptance of Secondary Queens in Eusocial Insects.

AbstractThe coexistence of multiple reproductives in eusocial insects is widespread, yet the decisions leading to additional queen acceptance are not well understood. Unlike in vertebrates, acceptance decisions are likely controlled by the more numerous helper population rather than the parent reproductive. Yet there are likely to be queen-worker differences in acceptance criteria because workers and queens differ in their relatedness to a secondary queen. We develop a model that examines queen-worker conflict in two scenarios: accepting a queen's sister or daughter. We additionally ask how the mating frequency and split sex ratios affect the outcomes of these conflicts. Our results reveal that conflict over queen acceptance is highest in monandrous mating systems. We identify a "window of conflict" in which a queen is selected to accept her sister but her workers do not. Our result, that polyandry neutralizes conflict over acceptance thresholds, suggests that conflict suppression may be an additional contributor to the maintenance of polyandrous mating systems.

Female mating rates and their fitness consequences in the common house spider Parasteatoda tepidariorum.

Mating systems, with varying female mating rates occurring with the same partner (monandry) or with multiple mates (polyandry), can have far reaching consequences for population viability and the rate of gene flow. Here, we investigate the mating rates of the common house spider Parasteatoda tepidariorum (Theridiidae), an emerging model for genetic studies, with yet undescribed reproductive behavior. It is hypothesized that spiders belonging to this family have low re-mating rates. We paired females twice with the same male (monandry) or with different males (polyandry), and recorded behaviors, mating success and fitness resulting from single- and double-matings, either monandrous or polyandrous. Despite the study being explorative in nature, we predict successful matings to be more frequent during first encounters, to reduce female risk of remaining unmated. For re-mating to be adaptive, we expect higher fitness of double-mated females, and polyandrous females to experience highest mating success and fitness if reproductive gains are achieved by mating with multiple partners. We show that the majority of the females did not mate, and those that did mated only once, not necessarily on their first encounter. The likelihood of re-mating did not differ between monandrous and polyandrous encounters and female mating experience (mated once, twice monandrous, twice polyandrous) did not affect fitness, indicated by similar offspring production. Female twanging of the web leads to successful matings suggesting female behavioral receptivity. Cannibalism rates were low and mostly occurred pre-copulatory. We discuss how the species ecology, with potentially high mating costs for males and limited female receptivity, may shape a mating system with low mating rates.

Atypical functioning of female genitalia explains monandry in a butterfly.

Monandrous species are rare in nature, especially in animals where males transfer nutrients to females in the ejaculate. The proximate mechanisms responsible for monandry are poorly studied. In butterflies and moths, the male transfers a nutritious spermatophore into the corpus bursae (CB) of the female. The CB is a multifunctional organ that digests the spermatophore and has partial control of the post-mating sexual receptivity of the female. The spermatophore distends the CB and the post-mating sexual receptivity of the female is inversely proportional to the degree of distension. The CB of many butterfly species has a muscular sheath whose contractions mechanically contribute to digest the spermatophore. As the contents of the CB are absorbed, the degree of distension decreases and the female recovers receptivity. We studied the monandrous butterfly Leptophobia aripa (Boisduval, 1836) (Pieridae) and found that females do not digest the spermatophores. We investigated the structure of the CB and found that a muscular sheath is absent, indicating that in this butterfly females lack the necessary "apparatus" for the mechanical digestion of the spermatophore. We propose that female monandry in this species is result of its incapability to mechanically digest the spermatophore, which results in a constant degree of CB distension after mating and, thus, in the maintenance of the sexually unreceptive state of females. Hypotheses on the evolution of this mechanism are discussed.

An empirical test of the bet-hedging polyandry hypothesis: Female red flour beetles avoid extinction via multiple mating.

Bet-hedging via polyandry (spreading the extinction risk of the female's lineage over multiple males) may explain the evolution of female multiple mating, which is found in a wide range of animal and plant taxa. This hypothesis posits that females can increase their fitness via polyandrous mating when "unsuitable" males (i.e., males causing reproductive failure for various reasons) are frequent in the population and females cannot discriminate such unsuitable mates. Although recent theoretical studies have shown that polyandry can operate as a bet-hedging strategy, empirical tests are scarce. In the present study, we tested the bet-hedging polyandry hypothesis by using the red flour beetle Tribolium castaneum. We compared female reproductive success between monandry and polyandry treatments when females mated with males randomly collected from an experimental population, including 20% irradiated (infertile) males. In addition, we evaluated geometric mean fitness across multiple generations as the index of adaptability of bet-hedging traits. Polyandrous females showed a significantly higher egg hatching rate and higher geometric mean fitness than monandrous females. These results strongly support the bet-hedging polyandry hypothesis.

No selection for change in polyandry under experimental evolution.

What drives mating system variation is a major question in evolutionary biology. Female multiple mating (polyandry) has diverse evolutionary consequences, and there are many potential benefits and costs of polyandry. However, our understanding of its evolution is biased towards studies enforcing monandry in polyandrous species. What drives and maintains variation in polyandry between individuals, genotypes, populations and species remains poorly understood. Genetic variation in polyandry may be actively maintained by selection, or arise by chance if polyandry is selectively neutral. In Drosophila pseudoobscura, there is genetic variation in polyandry between and within populations. We used isofemale lines to found replicate populations with high or low initial levels of polyandry and tracked polyandry under experimental evolution over seven generations. Polyandry remained relatively stable, reflecting the starting frequencies of the experimental populations. There were no clear fitness differences between high versus low polyandry genotypes, and there was no signature of balancing selection. We confirmed these patterns in direct comparisons between evolved and ancestral females and found no consequences of polyandry for female fecundity. The absence of differential selection even when initiating populations with major differences in polyandry casts some doubt on the importance of polyandry for female fitness.

How Efficient Is Fertilization by Traumatic Insemination in Orius insidiosus (Hemiptera: Anthocoridae)?

Traumatic insemination (TI) can be injurious to females, and females have evolved various paragenital structures to mitigate these impacts. We examined the mating behavior of Orius insidiosus (Say) and the consequences of single and double matings for female fitness. A total of 100 virgin females (4-6-d old) were directly observed while they mated with virgin males. Some of these females were mated a second time with a different, nonvirgin male 3-5 d later, after they oviposited in sunflower stems. Females were held in isolation, fed eggs of Ephestia kuehniella Zeller, and reproductive success was tracked for 30 d. Six females died during their first copulation (6%), and another within 48 h, without laying eggs. Four percent of the females died during their second copulations. Copulations lasting less than 90 s usually did not result in successful fertilization, and duration of copula was positively correlated with egg fertility in singly-mated females. Duration of copula was more than halved in second matings, twice as variable, and negatively correlated with 30 d fecundity. Thirty-seven percent of singly-mated females and 31% of twice-mated females were infertile, with fewer than half of all females producing 88% of all eggs. We conclude that O. insidiosus females are likely monandrous in the wild, and that TI in this species is inefficient, contributing to high variation in female fitness. Thus, mating involves a significant mortality risk for females, despite their possession of complex paragenital structures that ostensibly mitigate copulatory injury.

Dependence of mating rate on variance of eclosion time of cicadas (cicadidae).

In many cicadas, a phenomenon is observed in which the peak of emergence of males precedes that of females (protandry). Generally, it is known that the ecology of emergence of insects is closely related to mating, and protandry is said to be the strategy of males to eclose early and to attempt mating with virgin females. However, as for the cicadas, males only call females with songs, and as far as the behavior of males is concerned, the necessity of protandry is not felt. Thus, as to cicadas, little is known about the relationship between emergence ecology and mating. Therefore, investigating the temporal distribution of cicada emergence is an important step in considering cicadas mating, and even the evolution of mating system. Unfortunately, it is known that cicadas have a very long lifetime and are known to be difficult to keep, so it is meaningful to predict results mathematically. In this paper, we investigate the influence of male and female emergence spreading in appearance period on the mating rate by using mathematical (stochastic) model. Furthermore, we propose a new mathematical virtual model and try to explain the results that mating rate depends on sexual difference and variance of eclosion date.

Extreme cost of rivalry in a monandrous species: male-male interactions result in failure to acquire mates and reduced longevity.

Mating system variation is profound in animals. In insects, female willingness to remate varies from mating with hundreds of males (extreme polyandry) to never remating (monandry). This variation in female behaviour is predicted to affect the pattern of selection on males, with intense pre-copulatory sexual selection under monandry compared to a mix of pre- and post-copulatory forces affecting fitness under polyandry. We tested the hypothesis that differences in female mating biology would be reflected in different costs of pre-copulatory competition between males. We observed that exposure to rival males early in life was highly costly for males of a monandrous species, but had lower costs in the polyandrous species. Males from the monandrous species housed with competitors showed reduced ability to obtain a mate and decreased longevity. These effects were specific to exposure to rivals compared with other types of social interactions (heterospecific male and mated female) and were either absent or weaker in males of the polyandrous species. We conclude that males in monandrous species suffer severe physiological costs from interactions with rivals and note the significance of male-male interactions as a source of stress in laboratory culture.

Polyandry in nature: a global analysis.

A popular notion in sexual selection is that females are polyandrous and their offspring are commonly sired by more than a single male. We now have large-scale evidence from natural populations to be able to verify this assumption. Although we concur that polyandry is a generally common and ubiquitous phenomenon, we emphasise that it remains variable. In particular, the persistence of single paternity, both within and between populations, requires more careful consideration. We also explore an intriguing relation of polyandry with latitude. Several recent large-scale analyses of the relations between key population fitness variables, such as heterozygosity, effective population size (Ne), and inbreeding coefficients, make it possible to examine the global effects of polyandry on population fitness for the first time.

Female teneral mating in a monandrous species.

Schultesia nitor is a gregarious species living in Cacicus and Psarocolius ssp. pouch-like nests. Due to gregariousness, opportunities for multiple copulations in both sexes are not supposed to be restricted. Females produce only one brood during their life and die within a few days following the birth of their nymphs, but this unique brood could be the result of either single or multiple mating events (i.e., monandry vs. polyandry). In this study, we first determined the age of sexual receptivity of both males and females. Larval development in this species is shorter in males than in females and thus, this species is protandric. Males were not able to copulate the day after emergence. Contrary to males, teneral females (i.e., females achieving their imaginal molt but not yet fully sclerotised and colored) were attractive and were able to mate with males. In the second experiment, we tested the existence of multiple matings in both sexes. Our results showed that females were monandrous whereas males were polygynous. Since we had observed that females were monoandrous, we expected them to be choosy and we determined their ability to discriminate between virgin and nonvirgin males. When given the choice, females preferred virgin males and overall, they were more successful at mating than experienced ones. Our results suggest that monandry may be primarily driven by the female's short life-span fecundity. The occurrence of teneral mating in this species calls into question the existence of a male strategy for monopolizing females, and as well as the implication of female choice. Although further work is required, this species provides an interesting model for understanding sexual conflicts.

The role of male harassment on female fitness for the dengue vector mosquito Aedes aegypti.

Sexual harassment studies in insects suggest that females can incur several kinds of costs from male harassment and mating. Here, we examined direct and indirect costs of male harassment on components of female fitness in the predominantly monandrous mosquito Aedes aegypti. To disentangle the costs of harassment versus the costs of mating, we held females at a low or high density with males whose claspers were modified to prevent insemination, and compared these to females held with normal males and to those held with females or alone. A reduced longevity was observed when females were held under high density conditions with males or females, regardless if male claspers had been modified. There was no consistent effect of harassment on female fecundity. Net reproductive rate (R0) was higher in females held at low density with normal males compared to females held with males in the other treatments, even though only a small number of females showed direct evidence of remating. Indirect costs and benefits that were not due to harassment alone were observed. Daughters of females held with normal males at high density had reduced longevity compared to daughters from females held without conspecifics. However, their fitness (R0) was higher compared to females in all other treatments. Overall, our results indicate that A. aegypti females do not suffer a fitness cost from harassment of males when kept at moderate densities, and they suggest the potential for benefits obtained from ejaculate components.