Interpopulation variation in female remating is attributable to female and male effects in <Emphasis Type="Italic">Callosobruchus chinensis</Emphasis>

The evolution of female multiple mating is best understood by consideration of male and female reproductive perspectives. Females should usually be selected to remate at their optimum frequencies whereas males should be selected to manipulate female remating to their advantage. Female remating behavior may therefore be changed by variation of male and female traits. In this study, our objective was to separate the effects of female and male strains on female remating for the adzuki bean beetle, Callosobruchus chinensis, for which there is interstrain variation in female remating frequency. We found that interstrain variation in female remating is primarily attributable to female traits, suggesting genetic variation in female receptivity to remating in C. chinensis. Some interstrain variation in female remating propensity was attributable to an interaction between female and male strains, however, with the males of some strains being good at inducing nonreceptivity in females from one high-remating strain whereas others were good at inducing copulation in nonvirgin females from the high-remating strain. There is, therefore, interstrain variation in male ability to deter females from remating and in male ability to mate successfully with nonvirgin females. These results suggest that mating traits have evolved along different trajectories in different strains of C. chinensis.     

Mating speed differences between Australian populations ofDrosophila melanogaster

Mating speeds have been studied for two natural populations ofDrosophila melanogaster collected in Victoria, Australia, about 100 miles apart. Significant differences in mating speed were found and hybrids between the populations gave intermediate mating speeds. A small difference between strains in duration of copulation was found.The differences in mating speeds were controlled by the strain of the male, the strain of the female having little effect. No appreciable behavioural interactions were found between strains.     

Intraclonal variation in propensity for mixis in several rotifers: variation among females and with maternal age

In 14 previously published experiments with clones of three species of Brachionus, Epiphanes ukera, and Rhinoglena frontalis, females cultured singly in small volumes from birth through most of their reproductive period usually produced some mictic daughters. Here, these data are analyzed to test two hypotheses: the propensity of females in a clone to produce mictic daughters will vary significantly among females, and will decrease with female age. Significant heterogeneity (χ²) among replicate females was demonstrated in two clones of B. calyciflorus from Florida, in single clones of this species from Georgia, Texas, Spain, and Australia, in a clone of B. angularis from Argentina, and in single clones of E. ukera and R. frontalis from Germany. No significant heterogeneity was found in two other clones of B. calyciflorus from Florida and in a clone of B. variabilis from Spain. Significant heterogeneity among females of a clone could be caused by chance fluctuations during development, differences in birth order, or epimutations. This heterogeneity is an important component of a bet-hedging strategy to balance production of diapausing fertilized eggs and future population growth by female parthenogenesis. The propensity of a female to produce mictic daughters did not decrease with her age. In experiments with six different clones from four strains of B. calyciflorus, and with single clones of E. ukera and R. frontalis, only one clone of B. calyciflorus showed significant variation (ANOVA) in the proportion of mictic daughters produced in three or four successive one- or two-day periods. In this clone the proportion of mictic daughters increased with maternal age. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer research     

The evolution of male and female mating preferences in Drosophila speciation*

The relative importance of male and female mating preferences in causing sexual isolation between species remains a major unresolved question in speciation. Despite previous work showing that male courtship bias and/or female copulation bias for conspecifics occur in many taxa, the present study is one of the first large‐scale works to study their relative divergence. To achieve this, we used data from the literature and present experiments across 66 Drosophila species pairs. Our results revealed that male and female mate preferences are both ubiquitous in Drosophila but evolved largely independently, suggesting different underlying evolutionary and genetic mechanisms. Moreover, their relative divergence strongly depends on the geographical relationship of species. Between allopatric species, male courtship and female copulation preferences diverged at very similar rates, evolving approximately linearly with time of divergence. In sharp contrast, between sympatric species pairs, female preferences diverged much more rapidly than male preferences and were the only drivers of enhanced sexual isolation in sympatry and Reproductive Character Displacement (RCD). Not only does this result suggest that females are primarily responsible for such processes as reinforcement, but it also implies that evolved female preferences may reduce selection for further divergence of male courtship preferences in sympatry.     

Remating, sperm transfer, and sperm displacement in the cactophilic species Drosophila buzzatii Patterson & Wheeler (Diptera: Drosophilidae)

The mating system of Drosophila buzzatii is characterized by short copulation duration, frequent remating in both males and females, and male ejaculate partitioning. Additional features of the system are strong sperm displacement and a high frequency of sterile matings. Remating frequencies and the effects of remating on various mating parameters were studied. In order to characterize variation, five isofemale lines from geographically distant localities in Australia (three localities), Brazil and the Canary Islands were used. Mating parameters studied were: premating time, copulation duration, interval between successive matings, and progeny number as a measure of sperm transfer. Variation for sperm displacement was studied in crosses between laboratory stocks and a number of isofemale lines from Australia. There were significant between‐line differences in female remating frequencies, premating time, copulation duration, interval between successive matings, and progeny numbers, indicating genetic variation for these traits. Females from the five lines mated on average 1.6 to 3.1 times in 4 h, with a maximum of eight matings for one female. The males were given a maximum of ten virgin females in sequence and more than one‐third of the males mated all ten females in the 2 h observation period. Copulation duration decreased and interval between matings increased with copulation number in multiply mated males. Mean copulation duration was c. 2 min. Sperm transfer, measured as the average number of progeny from a single mating, was low (c. 25) and multiply mated females gave more progeny than single mated females, although with much lower progeny numbers than observed in wild‐caught non‐virgin females. A surprisingly high proportion of observed matings gave no progeny, i.e. they were sterile matings. Sperm displacement was strong in most crosses and remained strong in multiply mated females. The results are discussed in relation to the evolution of mating patterns in Drosophila.     

Time‐shifted reproductive behaviours among fall armyworm (Noctuidae: Spodoptera frugiperda) host strains: evidence for differing modes of inheritance

The noctuid moth Spodoptera frugiperda consists of two strains associated with different larval host plants (most notably corn and rice). These strains exhibit differential temporal patterns of female calling and copulation during scotophase, with the corn strain more active earlier in the night. We investigated strain‐specific constraints in reproductive timing, mating interactions between the two strains, and the mode of inheritance of timing of female calling, male calling, copulation and oviposition. We observed an allochronic shift of all reproductive behaviours by approximately 3 h and a parallel shift of nonreproductive locomotor activity, suggesting involvement of the circadian clock. The corn strain was more variable in the timing of calling and copulation than the rice strain. Rice strain females were more restricted in the timing of copulation than rice strain males, while such differences between the sexes were not apparent in the corn strain. There were significant interactions between the strains affecting onset times of copulation and male calling. The four investigated reproductive traits differed in their modes of inheritance: timing of female and male calling exhibited strong maternal effects, timing of copulation was controlled by a combination of maternal effects and corn strain dominant autosomal factors, and timing of oviposition was inherited in a corn strain dominant fashion. We conclude that the allochronic separation of reproduction between fall armyworm strains is asymmetric, less pronounced than previously thought, and under complex genetic control.     

Reproductive compatibility among latitudinally separated Scottolana canadensis (Willey) (Harpacticoida)

Scottolana canadensis is distributed widely along the east coast of North America. Although males exhibited mating behavior with females copepodites, copulation occurred following the female terminal molt. Crossing experiments revealed that in some cases, male mating behavior decreased with increasing latitudinal distance of male and female source locales. However, latitudinally-related differences in copepod body size most likely did not contribute to variation in male mating behavior. Survival patterns of fertilized eggs and fertility of offspring revealed postzygotic reproductive isolation of Florida populations from all others collected north to Maine.     

Females gain survival benefits from immune‐boosting ejaculates

Females in many animal taxa incur significant costs from mating in the form of injury or infection, which can drastically reduce survival. Therefore, immune function during reproduction can be important in determining lifetime fitness. Trade‐offs between reproduction and immunity have been extensively studied, yet a growing number of studies demonstrate that mated females have a stronger immune response than virgins. Here, we use the Texas field cricket, Gryllus texensis, to test multiple hypotheses proposed to explain this postmating increase in immune function. Using host‐resistance tests, we found that courtship, copulation, and accessory fluids alone do not affect female immunity; rather, only females that acquire intact ejaculates containing testes‐derived components exhibit significant increases in survival after exposure to bacterial pathogens. Our data suggest that male‐derived components originating from an intact ejaculate and transferred to females during sex are required for the increased immune function characteristic of mated female crickets to occur.     

Sex‐specific repeatabilities and effects of relatedness and mating status on copulation duration in an acridid grasshopper

In species with direct sperm transfer, copulation duration is a crucial trait that may affect male and female reproductive success and that may vary with the quality of the mating partner. Furthermore, traits such as copulation duration represent the outcome of behavioral interactions between the sexes, for which it is important—but often difficult—to determine which sex is in phenotypic control. Using a double‐mating protocol, we compared copulation durations between (1) virgin and nonvirgin and (2) sibling and nonsibling mating pairs in rufous grasshoppers Gomphocerippus rufus. Nonvirgin copulations took on average approximately 30% longer than virgin copulations, whereas relatedness of mating partners was not a significant predictor of copulation duration. Longer nonvirgin copulations may represent a male adaptation to sperm competition if longer copulations allow more sperm to be transferred or function as postinsemination mate guarding. The absence of differences between pairs with different degrees of relatedness suggests no precopulatory or preinsemination inbreeding avoidance mechanism has evolved in this species, perhaps because there is no inbreeding depression in this species, or because inbreeding avoidance occurs after copulation. Controlling for the effects of male and female mating status (virgin vs. nonvirgin) and relatedness (sibling vs. nonsibling), we found significant repeatabilities (R) in copulation duration for males (R = 0.33; 95% CI: 0.09–0.55) but not for females (R = 0.09; 95% CI: 0.00–0.30). Thus, copulation durations of males more strongly represent a nontransient trait expressed in a consistent manner with different mating partners, suggesting that some aspect of the male phenotype may determine copulation duration in this species. However, overlapping confidence intervals for our sex‐specific repeatability estimates indicate that higher sampling effort is required for conclusive evidence.     

Mating duration and sperm precedence in the spider <Emphasis Type="Italic">Linyphia triangularis</Emphasis>

In many animal species, mating behaviour is highly ritualised, which may allow us to relate some of its consequences, e.g. male paternity and female receptivity, to the progression of phases in the mating sequence; at the same time, ritualisation raises the question of to what extent the partners, especially the males, are able to influence the outcome of mating for their own benefit. We studied the linyphiid spider Linyphia triangularis in which mating follows a strict sequence during which the male inducts two droplets of sperm and transfers them to the female. We performed sperm competition experiments (sterile-male technique) including four treatments, in which the copulation of the first male was interrupted at prescribed phases of the mating sequence, while the second male was allowed a complete mating. Second males spent a shorter time than first males on the behaviours prior to sperm transfer, but the amount of sperm (2 droplets) and the time spent in sperm transfer were independent of the females’ mating status. The proportion of females accepting the second male depended on the mating duration of the first male, i.e. whether the first male had transferred one or two sperm droplets. After a complete first mating, most females accepted no further males. A last-male sperm precedence was apparent if only half of the first sperm droplet had been transferred by the first male, but this switched to a first male precedence if one full sperm droplet had been transferred. Thus, even in the face of sperm competition, it is sufficient for the first male to transfer one sperm droplet. The second sperm droplet and the extended copulatory courtship associated with its transfer may serve to induce a lack of receptivity in the female, but the males seem unable to enhance their reproductive success through variable copulatory tactics.     

Opposite Latitudinal Clines for First Mating and Second Mating (Remating) in Males of Drosophila melanogaster

Although female remating has been studied extensively in insects, few studies have been carried out for male remating (second mating). In this study, we analyzed Drosophila melanogaster males for their remating potential, using iso‐female line culture initiated with wild flies collected from eight Indian geographic localities. We examined the association of latitude and percent melanization with first and second male mating (including mating‐related traits). Our results indicated that second male mating has a negative latitudinal cline opposite to that of first mating. Body melanization is negatively correlated with second mating by male and positively with first mating (measured in terms of percent mated pairs). Mating latency during first (ML1) and second (ML2) male mating has a negative latitudinal cline, but slope values differ significantly as ML2 is great at higher latitudes as compared to ML1. The difference between ML1 and ML2 is non‐significant at lower latitudes. However, copulation period of second mating (CP2) has a negative latitudinal cline, whereas copulation period of first mating (CP1) has positive latitudinal cline. Next, the latency and copulation period differ significantly between first and second male mating treatments in within‐population analyses as well as in melanic strains. Furthermore, male remating ability (number of maximum successful remating attempts continuously by a male in 12 h) also follows negative latitudinal cline. The lower latitudinal lighter males have more remating ability as compared to darker males from higher latitudes.     

The role of functional constraints in nonrandom mating patterns for a dance fly with female ornaments

Most hypotheses to explain nonrandom mating patterns invoke mate choice, particularly in species that display elaborate ornaments. However, conflicting selection pressures on traits can result in functional constraints that can also cause nonrandom mating patterns. We tested for functional load‐lifting constraints during aerial copulation in Rhamphomyia longicauda, a species of dance fly that displays multiple extravagant female‐specific ornaments that are unusual among sexual traits because they are under stabilizing selection. R. longicauda males provide females with a nuptial gift before engaging in aerial mating, and the male bears the entire weight of the female and nuptial gift for the duration of copulation. In theory, a male's ability to carry females and nuptial gifts could constrain pairing opportunities for the heaviest females, as reported for nonornamented dance flies. In concert with directional preferences for large females with mature eggs, such a load‐lifting constraint could produce the stabilizing selection on female size previously observed in this species. We therefore tested whether wild‐caught male R. longicauda collected during copulation were experiencing load‐lift limitations by comparing the mass carried by males during copulation with the male's wing loading traits. We also performed permutation tests to determine whether the loads carried by males during copulation were lighter than expected. We found that heavier males are more often found mating with heavier females suggesting that whereas R. longicauda males do not experience a load‐lift constraint, there is a strong relationship of assortative mating by mass. We suggest that active male mate choice for intermediately adorned females is more likely to be causing the nonrandom mating patterns observed in R. longicauda.     

No evidence for external genital morphology affecting cryptic female choice and reproductive isolation in Drosophila

Genitalia are one of the most rapidly diverging morphological features in animals. The evolution of genital morphology is proposed to be driven by sexual selection via cryptic female choice, whereby a female selectively uptakes and uses a particular male's sperm on the basis of male genital morphology. The resulting shifts in genital morphology within a species can lead to divergence in genitalia between species, and consequently to reproductive isolation and speciation. Although this conceptual framework is supported by correlative data, there is little direct empirical evidence. Here, we used a microdissection laser to alter the morphology of the external male genitalia in Drosophila, a widely used genetic model for both genital shape and cryptic female choice. We evaluate the effect of precision alterations to lobe morphology on both interspecific and intraspecific mating, and demonstrate experimentally that the male genital lobes do not affect copulation duration or cryptic female choice, contrary to long‐standing assumptions regarding the role of the lobes in this model system. Rather, we demonstrate that the lobes are essential for copulation to occur. Moreover, slight alterations to the lobes significantly reduced copulatory success only in competitive environments, identifying precopulatory sexual selection as a potential contributing force behind genital diversification.     

Co-Evolution of the Mating Position and Male Genitalia in Insects: A Case Study of a Hangingfly

Hangingflies are unique for the male providing a nuptial gift to the female during mating and taking a face-to-face hanging copulation with the female. Their male genitalia are peculiar for an extremely elongated penisfilum, a pair of well-developed epandrial lobes (9th tergum), and a pair of degenerated gonostyli. However, the co-evolution of their face-to-face copulation behavior and the male genitalia has rarely been studied hitherto. In this paper the mating behavior of the hangingfly Bittacus planus Cheng, 1949 was observed under laboratory conditions, and the morphology of the male and female external genitalia was investigated using light and scanning electron microscopy. The male provides an insect prey as a nuptial gift to the female in courtship and mating process, and commits a face-to-face copulation. During copulation, the male abdomen twists temporarily about 180° to accommodate their face-to-face mating position. The aedeagal complex has an extremely elongated penisfilum, corresponding to the elongated spermathecal duct of the female. The well-developed epandrial lobes serve as claspers to grasp the female subgenital plate during copulation, replacing the function of gonostyli, which are greatly reduced in Bittacidae. The modified proctiger assists the penisfilum to stretch and to enter into the female spermathecal duct. The possible reasons why this species might mate face-to-face are briefly discussed.     

An enigmatic lineage of mites from Baltic amber shows a unique,possibly female‐controlled,mating

It is generally assumed that male control over mating and a lack of precopulatory female choice are prevalent in many animals and in astigmatan mites in particular. We show that several morphological structures of females of some astigmatan mites are indicative of precopulatory female choice: (1) copulatory tubes acting like intromittent organs; (2) specialized structures assisting male–female attachment and possibly allowing indirect mate selection in immature females; and (3) a unique, pad‐like terminal opisthosomal organ used to cling to the male during copulation in Glaesacarus rhombeus (= Acarus rhombeus Koch et Berendt, 1854) belonging to an extinct family, Glaesacaridae, from the Upper Eocene Baltic amber. An exceptionally well‐preserved copulating pair from amber provides insight into the function of this organ and reproductive behaviour in this mite. Female control over mating may reduce the timing of insemination, harassment by males, and damage caused by copulation. As a consequence, this can lessen male–male aggression, select against precopulatory guarding, and reduce the risk of predation. By contrast to extant taxa, males of G. rhombeus do not have any apparent specialized structures aiding clinging to the female during copulation, suggesting that this mating system is either an earlier step in the evolution of the female‐dominated mating system and/or a remarkable example of imbalanced female counteradaptations against the male's reproductive interest that may occur during an arms race between the two sexes. We offer an approach that can falsify the hypothesis assuming precopulatory female choice and discuss an alternative hypothesis suggesting that these female structures evolved in response to the need to reduce damage associated with mating or precopulatory guarding. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 661–668.     

Female Mating History Influences Copulation Behavior but Not Sperm Release in the Orb-Weaving Spider <Emphasis Type="Italic">Tetragnatha versicolor</Emphasis> (Araneae,Tetragnathidae)

We examined the influence of female mating history on copulation behavior and sperm release in the haplogyne spider Tetragnatha versicolor. Despite significant behavioral differences during mating, males released equivalent amounts of sperm to virgin and non-virgin females. When mating with non-virgin females, males showed twice as many pedipalp insertions and half the copulation duration as compared to virgin females; however, males were as likely to mate with non-virgin as virgin females. Even with these overt behavioral differences, males released half of the sperm contained within their pedipalps during mating, regardless of female mating history. With respect to male mating order, first or second, we suggest the numbers of sperm released would lead to an expectation of unbiased paternity. In this species, sperm release is not directly proportional to total copulationduration.     

Genotypes and their interaction effects on reproduction and mating‐induced immune activation in Drosophila melanogaster

Mating causes considerable alterations in female physiology and behaviour, and immune gene expression, partly due to proteins transferred from males to females during copulation. The magnitude of these phenotypic changes could be driven by the genotypes of males and females, as well as their interaction. To test this, we carried out a series of genotype‐by‐genotype (G × G) experiments using Drosophila melanogaster populations from two distant geographical locations. We expected lines to have diverged in male reproductive traits and females to differ in their responses to these traits. We examined female physiological and behavioural post‐mating responses to male mating traits, that is behaviour and ejaculate composition, in the short to mid‐term (48 hr) following mating. We then explored whether a sexually transferred molecule, sex peptide (SP), is the mechanism behind our observed female post‐mating responses. Our results show that the genotypes of both sexes as well as the interaction between male and female genotypes affect mating and post‐mating reproductive traits. Immune gene expression of three candidate genes increased in response to mating and was genotype‐dependent but did not show a G × G signature. Males showed genotype‐dependent SP expression in the 7 days following eclosion, but female genotypes showed no differential sensitivity to the receipt of SP. The two genotypes demonstrated clear divergence in physiological traits in short‐ to mid‐term responses to mating, but the longer‐term consequences of these initial dynamics remain to be uncovered.     

No Evidence for Heritability of Male Mating Latency or Copulation Duration across Social Environments in Drosophila melanogaster

A key assumption underpinning major models of sexual selection is the expectation that male sexual attractiveness is heritable. Surprisingly, however, empirical tests of this assumption are relatively scarce. Here we use a paternal full-sib/half-sib breeding design to examine genetic and environmental variation in male mating latency (a proxy for sexual attractiveness) and copulation duration in a natural population of Drosophila melanogaster. As our experimental design also involved the manipulation of the social environment within each full-sibling family, we were able to further test for the presence of genotype-by-environment interactions (GEIs) in these traits, which have the potential to compromise mate choice for genetic benefits. Our experimental manipulation of the social environment revealed plastic expression of both traits; males exposed to a rival male during the sensitive period of adult sexual maturation exhibited shorter mating latencies and longer copulation durations than those who matured in isolation. However, we found no evidence for GEIs, and no significant additive genetic variation underlying these traits in either environment. These results undermine the notion that the evolution of female choice rests on covariance between female preference and male displays, an expectation that underpins indirect benefit models such as the good genes and sexy sons hypotheses. However, our results may also indicate depletion of genetic variance in these traits in the natural population studied, thus supporting the expectation that traits closely aligned with reproductive fitness can exhibit low levels of additive genetic variance.