The function of mate guarding in a field cricket (Orthoptera: Gryllidae;Teleogryllus natalensis otte and cade)

Three hypotheses relating to the function of postcopulatory mate guarding were tested for the cricketTeleogryllus natalensis. The hypothesis that guarding allows the male to remain with the female for repeated matings was rejected. This was because the mean intercopulatory interval for maleT. natalensis was found to be nearly twice as long as the mean duration of guarding. Nor do the results provide evidence to support the hypothesis that guarding functions to prevent copulation attempts by rival males (the rival exclusion hypothesis): the presence of a rival male was found to have no significant effect on the duration of spermatophore attachment for either guarded or unguarded females. The results do, however, support a third hypothesis, namely, that guarding functions to prevent the female from removing the spermatophore ampulla before complete sperm transfer. As predicted by this hypothesis, the presence of a guarding male was found to have a significant positive effect on the duration of spermatophore attachment. Further support for this hypothesis was provided by the fact that there was a significant positive correlation between the duration of mate guarding and the duration of spermatophore attachment.     

Competing for last place: Mating behaviour in a pill-box crab, Halicarcinus cookii (Brachyura: Hymenosomatidae)

The mating strategy of Halicarcinus cookii was investigated to ascertain how males maximised their fitness through mate choice. An intertidal population at Kaikoura, New Zealand, was dominated by mature crabs of both sexes in summer and by immature crabs in the colder months. More than 95% of mature females were ovigerous with early stage and late stage broods found in almost every month, indicating that egg production and larval release is continuous. The operational sex ratio was less than 1 male/female in summer, but often more than 1.0 in the colder months. The gonosomatic index increased along with brood development so that as soon as zoeae were released, the next clutch of eggs was ready to be fertilised. Males searched for receptive females and began pre-copulatory mate guarding without any courtship display. They mated preferentially with late stage or non-ovigerous females: copulation duration was longest for stage 5 females as was post-copulatory guarding (mean 18.3 h). Late stage females were up to 14% of the female population. Mate attraction seems to be the result of an ovarian signal rather than from the developing brood. Manipulation of the sex ratio had effects upon copulation duration and post-copulatory guarding: presence of a rival male increased duration of guarding. Females showed precocious mating in the penultimate instar and were able to lay fertilised eggs after their pubertal moult in the absence of males. H. cookii females have many mates, but males attempt to ensure paternity by preferentially pursuing mature females close to egg laying and by guarding these females after copulation. These behaviours are all elements of a competitive strategy to ensure that a male loses (not wins) the race to copulate because females have a ventral seminal receptacle, giving sperm precedence to the last male to mate. Male mating behaviour is a consequence and evolutionary response to female morphology.     

Rapid diversification of male genitalia and mating strategies in Ohomopterus ground beetles

We analysed evolutionary diversification and covariation in male genitalia and four mating traits related to sexual selection, i.e. testis size, spermatophore size, copulation duration and post-copulatory guarding duration, in Ohomopterus ground beetles using phylogenetically independent contrasts. Male genital size and mating duration have evolved more rapidly than body size and the other traits studied. Male genital size was negatively correlated with copulation duration, suggesting that elongated male genitalia may enable decreased time investment in a single copulation because it is more effective at facilitating spermatophore deposition. Male genital size was positively correlated with spermatophore size, suggesting coevolution between offensive and defensive male mating tactics because the elongated male genitalia may be advantageous in displacement of rivals' plug-like spermatophores, and decreased mating duration may intensify sperm competition. Thus, the remarkable diversity of male genitalia in Ohomopterus may have been facilitated by the interplay between inter- and intrasexual selection processes.     

Increased copulation duration before ejaculate transfer is associated with larger spermatophores, and male genital titillators, across bushcricket taxa

Copulation duration varies considerably across species, but few comparative studies have examined factors that might underlie such variation. We examined the relationship between copulation duration (prior to spermatophore transfer), the complexity of titillators (sclerotized male genital contact structures), spermatophore mass and male body mass across 54 species of bushcricket. Using phylogenetic comparative analyses, we found that copulation duration was much longer in species with titillators than those without, but it was not longer in species with complex compared with simple titillators. A positive relationship was found between spermatophore size and copulation duration prior to ejaculate transfer, which supports the hypothesis that this represents a period of mate assessment. The slope of this relationship was steeper in species with simple rather than complex titillators. Although the data suggest that the presence of titillators is necessary to maintain long copulation prior to ejaculate transfer, mechanisms underlying this association remain unclear.     

Heritable variation in the timing of spermatophore removal, a mechanism of post-copulatory female choice in crickets

Female crickets can exert post-copulatory mating preferences by prematurely removing a male's spermatophore after copulation, which terminates sperm transfer. Although most models of sexual selection assume that female mating preferences are heritable, there has been little work addressing genetic variation underlying post-copulatory mate choice. We used a paternal half-sib design, in which different males were randomly assigned as mates to several females to create half-sib families, to determine the heritability of spermatophore retention time in female house crickets, Acheta domesticus. There was significant additive genetic variance in the timing of spermatophore removal by females [h(2) = 0.50 +/- 0.19 (+/- SE)], suggesting that the timing of spermatophore removal is determined, in part, by the female's own genotype independent of the quality of her mate. The relatively high heritability of spermatophore retention time may be reflective of the absence of strong selection on this trait, consistent with previous work showing no difference in the fitness of females permitted to freely remove the spermatophore of their mates and those forced to accept complete ejaculates.     

Mating behavior and genital damage during copulation in the leaf beetle Lema coronata (Chrysomelidae: Criocerinae)

The criocerine leaf beetle Lema coronata Baly has extremely long genitalia that reach more than twice the body length in both sexes. We observed mating behavior of this species in the laboratory and inspected the male genital morphology using a scanning electron microscope. The males did not perform pre-copulatory courtship and post-copulatory guarding of the mates, and copulation lasted only for about 30 min. The surface of male genitalia is smooth without any special structures at the tip. A fragment of broken male genitalia was detected in the spermathecal duct of one female. We discuss the adaptive significance of male genital damage and the selective factor of elongated genitalia.     

Sperm traits negatively covary with size and asymmetry of a secondary sexual trait in a freshwater crayfish

In species where females mate promiscuously, the reproductive success of males depends both on their ability to acquire mates (pre-copulatory sexual selection) and ability of their ejaculates to outcompete those of other males (post-copulatory sexual selection). Sperm competition theory predicts a negative relationship between investment in body traits favouring mate acquisition (secondary sexual characters, SSCs) and investment in ejaculate size or quality, due to the inherent costs of sperm production. In contrast, the phenotype-linked fertility hypothesis posits that male fertilizing efficiency is reliably reflected by the phenotypic expression of male SSCs, allowing females to obtain direct benefits by selecting more ornamented males as copulation partners. In this study, we investigated the relationships between male SSCs and size and quality (viability and longevity) of ejaculates allocated to females in mating trials of the freshwater crayfish Austropotamobius italicus. We showed that the relative size of male weapons, the chelae, was negatively related to ejaculate size, and that chelae asymmetry, resulting from regeneration of lost chelipeds, negatively covaried with sperm longevity. Moreover, males allocated more viable sperm to mates from their own rather than different stream of origin. Our findings thus suggest that, according to sperm competition theory, pre-copulatory sexual selection for large weapons used in male fighting may counteract post-copulatory sperm competition in this crayfish species, and that investment in cheliped regeneration may impair ejaculate quality.     

Genetic association between male attractiveness and female differential allocation

Differential allocation of reproductive effort towards offspring of attractive mates is a form of post-copulatory mate choice. Although differential allocation has been demonstrated in many taxa, its evolutionary implications have received little attention. Theory predicts that mate choice will lead to a positive genetic correlation between female preference and male attractiveness. This prediction has been upheld for pre-copulatory mate choice, but whether such a relationship between male attractiveness and female differential allocation exists has never been tested. Here, we show that both female pre-copulatory mate choice and post-copulatory differential allocation are genetically associated with male attractiveness in house crickets, Acheta domesticus. Daughters of attractive males mated sooner and laid more eggs when paired with larger males. These forms of mate choice are strongest in large females, suggesting that costs decrease with increasing female size. The genetic association between attractiveness and differential allocation suggests potential for differential allocation to become exaggerated by coevolutionary runaway processes in an analogous manner to pre-copulatory choice. Sexual selection is thus likely to be stronger than predicted by pre-copulatory choice alone.     

The Effects of Age and Previous Mating Experience on Pre- and Post-copulatory Mate Choice in Female House Crickets (<Emphasis Type="Italic">Acheta domesticus</Emphasis> L.)

Although females’ mating preferences are influenced by male characteristics, there are a number of factors intrinsic to females and unrelated to male phenotype that can modulate female choice. We assessed the effects of age and mating experience on mechanisms of pre- and post-copulatory mate choice in female house crickets, Acheta domesticus L., by randomly assigning males to females, but independently varying the age and number of previous matings of females at the time of experimental matings. Latency to mating, a measure of a female’s pre-copulatory preference, was influenced by female age at the time of mating, with older females mating sooner than younger females. The reduced selectivity of older females appears consistent with life-history theory, which predicts that the reproductive value of females should decline with age. The length of time that females retained the spermatophore after mating, a measure of a female’s post-copulatory mating preference, was not influenced by female age at the time of mating, the number of previous matings, or any interaction between the two main effects. Contrary to previous reports, male mass had no effect on either the latency to mating or female retention of the spermatophore in A. domesticus. We conclude that female age and mating experience can moderate female selectivity, but that their impact varies according to the mechanism by which females favor particular sires.     

Male Gryllus bimaculatus Guard Females to Delay Them from Mating with Rival Males and to Obtain Repeated Copulations

Three hypotheses for the function of postcopulatory mate guarding were tested in the field cricket Gryllus bimaculatus De Geer. The duration of spermatophore attachment was greater in the absence than in the presence of a guarding male. The ejaculate protection hypothesis was, therefore, rejected. The duration of mate guarding was found to be equal to the interval between copulations, supporting the spermatophore renewal hypothesis. In support of the rival exclusion hypothesis, the presence of a guarding male did increase the duration of spermatophore attachment when a rival male was also present. The presence of a guarding male also delayed the female from mating with the rival male. Female mating status had a significant effect on the duration of spermatophore attachment. Females mating for the first time retained the spermatophore for a significantly longer period of time than females that had mated previously.     

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

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

Delayed copulation as a means of female choice by the hermit crab Pagurus filholi

Male hermit crabs perform precopulatory mate-guarding behavior during their reproductive season. As females generally cannot reject guarding attempts by males, male guarding prevents females from inspecting and choosing other male mates. However, as guarding males are often replaced by other males through competition for females during the guarding phase, females may be able to select males by delaying their copulation. To examine the possibility of female choice by hermit crabs, we investigated whether female Pagurus filholi that were being guarded in the field were ready to copulate and spawn. We found that about 30% of females guarded in the field were ready to spawn, indicating that guarded females delayed copulation with their current male. Our results suggest that by delaying copulation females may exploit male–male competition to choose dominant males. However, delaying copulation reduced the spawning potential of females. Hence, there is a trade-off between waiting for the opportunity to mate with a dominant male and decreased spawning success if females exploit male–male competition.     

Age-Related Variation in Mate-Guarding Intensity in the Bluethroat (Luscinia s. svecica)

Female extra‐pair copulations (EPCs) have selected for male paternity guarding strategies in many bird species. In the bluethroat, Luscinia s. svecica, males guard their mates closely during the last 2 d before the start of egg laying, but there is great individual variation in the intensity of mate guarding. Here we show that some of this variation is related to male age. Old males guarded their mates with much lower intensity and sang more than young males, although the latter difference was not statistically significant. Controlling for male age, male and female coloration and size were not significantly related to the intensity of mate guarding. We have previously shown that young and old males had a similar paternity loss in their own broods. On the other hand, old males were far more successful than young males in achieving extra‐pair fertilizations. These patterns suggest that young and old males have different trade‐offs between preventing paternity loss in own nest and gaining paternity in others, because male skills in obtaining EPCs improve with experience and/or because of female preferences for old males as copulation partners. There were no significant relationships between paternity and male mate‐guarding behaviour during the fertile period, indicating that mate guarding is not a very effective paternity‐assurance strategy in the bluethroat.     

Chemosensory assessment of sperm competition levels and the evolution of internal spermatophore guarding

Males of many species adjust their reproductive behaviour according to the perceived risk of sperm competition. Although this phenomenon is widespread in insects and other animals, the mechanisms that allow mates to assess sperm competition levels remain largely unexplored. In this study, we analysed the mating behaviour of pairs of Tenebrio molitor beetles under three odour treatments representing increasing levels of sperm competition risk (SCR) and sperm competition intensity (SCI). Copula duration and male and female post-copulatory behaviour varied significantly with odour treatment. Both copula duration and post-copulatory associations (PCAs) increased significantly in odour treatments reflecting high male density. To our knowledge, this is the first study to report that insects may assess the actual density of potential competitors at the time of mating, a cue to SCR and SCI, on the basis of chemical cues. In T. molitor, males inhibit sperm release from the spermatophore of a rival male when remating takes place at short intervals. We show that, when sperm competition levels are high, PCAs increase female remating interval just above that necessary to prevent spermatophore inhibition by rival males. This finding strongly suggests that strategic male behaviour plays a 'spermatophore guarding' role in this species. Although common in insects with external spermatophore transfer, spermatophore guarding is not expected in species with rapid ejaculate transfer and internal spermatophore delivery. Our results reveal that spermatophore guarding may evolve, even under these circumstances, as an evolutionary response to short-term spermatophore inhibition or displacement mechanisms.     

Intensive resistance by females before copulation induces insemination failure in the West Indian sweet potato weevil <Emphasis Type="Italic">Euscepes postfasciatus</Emphasis>

Persistent mating attempts by males (sexual harassment) are frequently observed among animals. For females, resisting persistent males can be costly because vigorous resistance increases both energy expenditure and the possibility of injury. Although one tactic for coping with male harassment is to cease resistance and mate with the persistent partner, the females of several species are able to prevent the fertilization of their egg(s) despite copulation. In this study, we used three different sex ratios to investigate whether a male’s mating persistence affects his mating success in the West Indian sweet potato weevil Euscepes postfasciatus, in which males mount females both before and after copulation. Consistent with our predictions, we found that female weevils resist and manipulate sperm transfer either before or during copulation according to their preferences. Female weevils were able to reject the sperm of persistent males despite having copulated with them. However, neither copulation and/or post-copulatory mounting affected insemination success. We speculate that the intensive resistance shown by females before copulation may induce mechanical sterility in E. postfasciatus.     

Hang on or run? Copepod mating versus predation risk in contrasting environments

Mating durations of copepods were found to differ significantly between fishless high-altitude waters and lowland lakes containing fish. In lowland species the whole mating process was completed within a few minutes, but it averaged over an hour in high-altitude species. Alpine copepods showed a prolonged post-copulatory association between mates, during which the male clasped the female for an extended period after spermatophore transfer, while in lowland species males abandoned their partner immediately after copulation. Prolonged associations also occurred after transfer of spermatophores to heterospecific females with shorter conspecific mating duration, suggesting that male interests largely dictate the time spent in tandem. The differences observed may be adaptations to environments with different predation pressure, as pairs in tandem are more conspicuous and less reactive than single animals. We argue that differences in mating behavior and mating duration evolved under sexual versus natural selection, reflecting trade-offs between enhancement of fertilization success and reduction of vulnerability to visual predation. In fishless mountain lakes with high intrasexual competition, guarding males can reduce the risk of spermatophore displacement or the risk that the female will accept sperm from rival males without increased risk of being eaten, thereby maximizing paternity. Populations from fishless alpine lakes further differed from lowland species by exhibiting higher female/male size dimorphism and more intense pigmentation. While these traits vary between populations according to predation pressure, mating duration appears to be more species-specific.     

Evidence for Inbreeding Depression and Pre-Copulatory,but Not Post Copulatory Inbreeding Avoidance in the Cabbage Beetle Colaphellus bowringi

Inbreeding is known to have adverse effects on fitness-related traits in a range of insect species. A series of theoretical and experimental studies have suggested that polyandrous insects could avoid the cost of inbreeding via pre-copulatory mate choice and/or post-copulatory mechanisms. We looked for evidence of pre-copulatory inbreeding avoidance using female mate preference trials, in which females were given the choice of mating with either of two males, a sibling and a non-sibling. We also tested for evidence of post-copulatory inbreeding avoidance by conducting double mating experiments, in which four sibling females were mated with two males sequentially, either two siblings, two non-siblings or a sibling and a non-sibling in either order. We identified substantial inbreeding depression: offspring of females mated to full siblings had lower hatching success, slower development time from egg to adult, lower survival of larval and pupal stages, and lower adult body mass than the offspring of females mated to non-sibling males. We also found evidence of pre-copulatory inbreeding avoidance, as females preferred to mate with non-sibling males. However, we did not find any evidence of post-copulatory inbreeding avoidance: egg hatching success of females mating to both sibling and non-sibling males were consistent with sperm being used without bias in relation to mate relatedness. Our results suggest that this cabbage beetle has evolved a pre-copulatory mechanism to avoid matings between close relative, but that polyandry is apparently not an inbreeding avoidance mechanism in C. bowringi.     

Pairing and insemination patterns in a giant weta (Deinacrida rugosa: Orthoptera; Anostostomatidae)

Positive size assortative mating can arise if either one or both sexes prefer bigger mates or if the success of larger males in contests for larger females leaves smaller males to mate with smaller females. Moreover, body size could not only influence pairing patterns before copulation but also the covariance between female size and size of ejaculate (number of spermatophores) transferred to a mate. In this field study, we examine the pre-copulatory mate choice, as well as insemination, patterns in the Cook Strait giant weta (Deinacrida rugosa). D. rugosa is a large orthopteran insect that exhibits strong female-biased sexual dimorphism, with females being nearly twice as heavy as males. Contrary to the general expectation of male preference for large females in insects with female-biased size dimorphism, we found only weak support for positive size assortative mating based on size (tibia length). Interestingly, although there was no correlation between male body size and the number of spermatophores transferred, we did find that males pass more spermatophores to lighter females. This pattern of sperm transfer does not appear to be a consequence of those males that mate heavier females being sperm depleted. Instead, males may provide lighter females with more spermatophores perhaps because these females pose less of a sperm competition risk to mates.     

Male Drosophila melanogaster show adaptive mating bias in response to female infection status

Given the non-trivial cost of reproduction for males and substantial variation in female quality, males have been predicted to show mating bias as an evolved strategy. Using a large outbred population of Drosophila melanogaster, we test this prediction and show that males may adaptively bias their mating effort in response to the infection status of females. Given a simultaneous choice between females infected with pathogenic bacteria and sham infected females, males preferentially mated with the latter, who had a higher reproductive output compared to infected females. This may provide evidence for pre-copulatory male mate choice. Assessment of the reproductive behaviour ensured that the observed pattern of mating bias was not due to differences in receptivity between females infected with pathogenic bacteria and sham infected females. Further, there was no evidence for post-copulatory male mate choice measured in terms of copulation duration.     

Extra-pair mate choice in the female great tit <Emphasis Type="Italic">Parus major</Emphasis>: good males or compatible males

The good genes hypothesis and the genetic compatibility hypothesis are the two main hypotheses that focus on the genetic benefit that a female can gain through her choice of a mate. We tested the two hypotheses on extra-pair mating in the great tit, Parus major. We found that female great tits choose males on the basis of breast stripe width, which is in accordance with the good genes hypothesis. Although females chose less related extra-pair males, the evidence for female choice for compatible males was overall weak. However, our data suggest a post-copulatory mechanism of inbreeding avoidance. The observed individual inbreeding coefficient, F, was similar for within-pair offspring (WPO) and extra-pair offspring (EPO). The observed individual F of WPO was lower than the expected individual F, whereas the observed F of EPO was similar to what was expected. These results highlight the importance of processes after copulation for the outcome of female mate choice. Our study shows that in a system with apparent pre-copulatory female choice for good genes, a post-copulatory mechanism may still promote the production of offspring that carry compatible genes.