Repeatability and heritability of sperm competition outcomes in males and females of <Emphasis Type="Italic">Tribolium castaneum</Emphasis>

Differences among males in their success in achieving fertilisations when females mate with more than one partner are now recognised as an important target of sexual selection. However, very few studies have attempted to determine whether particular males are consistently successful in sperm competition and whether success in sperm competition is a heritable trait. Additionally, the potential heritability of female traits that influence the outcome of sperm competition has received only limited attention. Using the polyandrous beetle Tribolium castaneum, we examined repeatability of male success in sperm competition by mating pairs of males carrying different visible genetic markers to a string of different females. Males showed consistency in their ability to successfully transfer sperm to females, but not in their success in sperm competition. Furthermore, when we independently compared success in sperm competition of fathers with their sons, we found no evidence for heritability of this trait. Similarly, females that exhibited high or low first male sperm precedence did not tend to have daughters that showed the same pattern. Our results suggest that we should be wary of assuming that success in sperm competition is heritable through either sex.     

Algal blooms decrease care but increase egg survival in a fish with paternal care

Human-induced eutrophication, resulting in algal blooms and increased water turbidity, is an alarming problem in aquatic systems. Here, we experimentally tested the impact of algal turbidity on parental care, egg fanning, and time in the nest, in the sand goby, Pomatoschistus minutus, a fish with uniparental male care. We allowed males to care for their eggs in either clear water or water made turbid by planktonic algae. In the early brood cycle, males fanned their eggs less in turbid than in clear water, but this difference disappeared later. Despite decreased care, egg survival was higher in turbid conditions, indicating that early fanning may partly be redundant for egg survival and perhaps used more as courtship. Males also spent more time out of their nest in turbid water, perhaps as a means to encounter additional females under conditions of low visibility.     

Sperm competition and evidence of sperm fertilization patterns in the carrion ball-roller beetle Canthon cyanellus cyanellus LeConte (Scarabaeidae: Scarabaeinae)

Field observation and laboratory experiments were conducted to investigate sperm competition and patterns of sperm fertilization under different experimental conditions in the carrion ball-roller beetle Canthon cyanellus cyanellus. Males in nature can mate with females whose spermathecae contains fertile sperm from other males. Sperm precedence was investigated using a visible genetic marker. The progeny of red (homozygous recessive) virgin females mated once with a red male and afterward, once with a green beetle (homozygous dominant) and vice versa, revealed that regardless of its color, the last male to mate fertilized c.a. 50% of the eggs, suggesting strong sperm competition. Males were able to achieve higher levels of paternity (more than 80%) when mated ad libitum with previously mated females, although large amount of variance in paternity does not exclude the possibility of first male sperm precedence or female cryptic choice. These results suggest that repeated mating and sperm replacement are the mechanisms by which last males achieve sperm precedence.     

Is male rhesus macaque red color ornamentation attractive to females?

Male sexually selected traits can evolve through different mechanisms: conspicuous and colorful ornaments usually evolve through intersexual selection, while weapons usually evolve through intra-sexual selection. Male ornaments are rare among mammals in comparison to birds, leading to the notion that female mate choice generally plays little role in trait evolution in this taxon. Supporting this view, when ornaments are present in mammals, they typically indicate social status and are products of male-male competition. This general mammalian pattern, however, may not apply to rhesus macaques (Macaca mulatta). Males of this species display conspicuous skin coloration, but this expression is not correlated to dominance rank and is therefore unlikely to have evolved due to male-male competition. Here, we investigate whether male color expression influences female proceptivity toward males in the Cayo Santiago free-ranging rhesus macaque population. We collected face images of 24 adult males varying in dominance rank and age at the peak of the mating season and modeled these to rhesus macaque visual perception. We also recorded female sociosexual behaviors toward these males. Results show that dark red males received more sexual solicitations, by more females, than pale pink ones. Together with previous results, our study suggests that male color ornaments are more likely to be a product of inter- rather than intra-sexual selection. This may especially be the case in rhesus macaques due to the particular characteristics of male-male competition in this species.     

Effects of female mating status on copulation behaviour and sperm expenditure in the freshwater crayfish <Emphasis Type="Italic">Austropotamobius italicus</Emphasis>

Sperm competition models predict that males should adjust their sperm expenditure according to the risk and/or intensity of sperm competition. In this paper, we analysed copulatory behaviour of both sexes and sperm expenditure in relation to female mating status (virgin or mated) in the freshwater crayfish Austropotamobius italicus, a species where males have been reported to feed on and remove sperm laid by other males. The same females were allowed to be inseminated sequentially by two males, and we compared the sexual behaviours of partners between the first (virgin females) and the second mating (mated females). We found that female resistance did not differ between the first and the second mating, nor males refused or took more time to mount a mated female. However, when mating with a mated female, males reached an effective copulation position significantly later. This occurred because second-mating males removed, by eating, all or most spermatophores previously deposited by first males. As removal was often incomplete, this resulted in a larger amount of sperm being deposited on female ventral parts after the second mating, although second males did not allocate more sperm to mated females than first males did. Thus, the peculiar mode of sperm competition, where males remove previously deposited sperm, and the consequent predictable strong last male prevalence in paternity likely led to the observed lack of adjustment of sperm expenditure to female mating status in this species.     

The relative roles of predation and sperm competition on the duration of the post-copulatory association between the sexes in the blue crab, Callinectes sapidus

In many species, post-copulatory mate guarding prevents other males from mating with the guarded female. In crabs, males stay with their mates to protect the female from predators because, in some species, mating occurs when she is soft and vulnerable after molting. I tested the relative roles of sperm competition and predation on the duration of the post-copulatory association in the blue crab, Callinectes sapidus. Unpaired females suffered greater predation mortality than paired females and males stayed with the female longer in the presence of predators than in their absence, suggesting that the post-copulatory association protects females during their vulnerable period. However, the association may also occur in blue crabs because of sperm competition since spermathecal contents of females in the field indicate that 12.4% mated twice. Females experimentally mated with two males contained both males ejaculates and each ejaculate had access to the unfertilized eggs, suggesting that the size of a male's ejaculate influences his fertilization rate in a multiply-mated female. Males stayed longest in response to a high risk of sperm competition. Longer post-copulatory associations allowed the first male's ejaculate to harden into a type of sperm plug, which limited the size of a second inseminator's ejaculate in a non-virgin female as compared with a virgin. Males passed larger ejaculates in the presence of rivals and when previous ejaculates were in the female spermathecae, another response to sperm competition. Larger ejaculates may need longer post-copulatory associations before a more effective sperm plug forms. Large males stayed with the female longer, which is consistent with their ability to pass larger ejaculates than small males and suggests that there may be costs to minimizing the duration of the post-copulatory association. In the field, associations last long enough to protect the female during her vulnerable phase and may ensure that the guarding male fertilizes the most eggs in the female, even if she remates. Thus, the post-copulatory association protects female blue crabs from additional inseminators as well as from predators. Received: 23 January 1996 / Accepted after revision: 9 November 1996     

Alternative mating strategies in the water strider Gerris elongatus (Heteroptera,Gerridae)

Summary Males of the water strider Gerris elongatus established territories which included copulation and oviposition sites (small pieces of fallen bamboo). Males were aggressive and competition for territory and females was observed frequently. Male midlegs were more developed than female midlegs and were used as weapons. Reproductive behaviour changed as the breeding season advanced. Early in the season immature females were attracted by male surface wave courtship signals, then copulated white floating on the water surface without ovipositing (type 1). In midseason, males established territories, produced calling signals and attracted females which copulated and oviposited there with male postcopulatory guarding (type 2). In late season, many females oviposited without postcopulatory guarding on pondweed mats near fallen bamboo. Non-territorial males waiter for the arrival of these females and copulated without courtship, but mating success was low (type 3). These alternative mating strategies appeared to depend on differences in male size. Larger males were superior to smaller males in many ways (establishing territory, fighting, mating etc.). The largest males defended territories and had higher mating success than small non-territorial males. Medium sized males used all three strategies according to the number of empty territories and seasonal femald distribution.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Male preference for colourful females affected by male size in a marine fish

It is becoming increasingly clear that mate preferences are not static, but can vary as a function of ecological conditions and the state of the choosing individual. This applies not only to females, the sex that has usually been the subject in research on mate preferences, but also to males. Under certain conditions, males should be selective in their choice of breeding partner. In the two-spotted goby, Gobiusculus flavescens, a small marine fish, breeding females develop conspicuous yellow-orange bellies, which they actively display to males during courtship. We have recently shown that males prefer more colourful females as mates. In this study, we test if the size of a male affects his preference for colourful females. Using three-compartment mate-choice aquaria, we recorded the interest shown by a male in two females differing in coloration but similar in size. Large and small males were equally eager to court females, but only large males showed a greater interest in the more colourful females. We suggest that small males are unselective because they usually obtain few mating opportunities, as a result of being unsuccessful in mate attraction or male contest competition. This study provides the first demonstration that the size of a male affects his preference for female colour.     

Assessment of sperm competition by European bitterling,<Emphasis Type="Italic"> Rhodeus sericeus</Emphasis>

We investigated male assessment of sperm competition in the bitterling, Rhodeus sericeus, a freshwater fish that spawns on the gills of living unionid mussels. Field experiments showed males increased their inspection rate of mussels into which a testis solution containing sperm had been experimentally released. Males avoided leading females to mussels that contained high numbers of embryos, but did not alter their leading behavior in response to the presence of sperm. In laboratory experiments males also increased their inspection rate of mussels into which a testis solution had been released and also failed to alter their leading behavior in response to the presence of sperm in mussels. However, males avoided leading females to mussels in close proximity to other males, and thereby may avoid sperm competition. In a second field study, territorial males were shown to ejaculate into mussels at a low rate in the absence of competitors, increase the frequency of ejaculations in competition with a rival, then decrease relative ejaculate expenditure as the number of competing males increased. Observed data were shown to be significantly correlated with predicted estimates of ejaculate expenditure for a model of sperm competition intensity. We discuss our results in the context of adaptive responses of males to sperm competition.     

Determinants of mating success in the golden hamster (Mesocricetus auratus) IV. Sperm competition

Summary This study was designed to clarify several aspects of sperm competition in the golden hamster (Mesocricetus auratus). By mating females homozygous for a recessive coat color gene with two males, one homozygous for the dominat gene (agouti) and one homozygous for the recessive (cream), paternity of the resulting young could be determined by day 2 postpartum. When females received 5 consecutive ejaculations from a male of one strain followed by 5 ejaculations from a male of the other strain, there was a significant advantage with respect to litter composition for the male ejaculating first. A mechanism for the first male mating advantage was suggested by the results of an experiment in which the time interval between first and second males was varied. Delays of 30 min between first and second males significantly increased the proportion of young sired by the first male. With a delay of 8 h, the first male sired all of the young. Delays between matings by males of the two strains had greater effects when mating tests began around the time of ovulation, rather than several hours before. Ejaculations in excess of those required for pregnancy initiation were shown to have a two-fold effect. They afforded the first male a measure of protection against displacement by a subsequent male. Conversely, prolonged copulation by the second male resulted in significant displacement of the sperm of a predecessor. The protective effects of multiple ejaculations by the first male were mediated by two mechanisms: the delay in introduction of competing sperm and an increase in the relative concentration of the first male's sperm. Prolonged copulation by the second male also appeared to function through its effect on relative sperm concentrations. Males of these genotypes differed with respect to the relative fertilizing capacity of their cjaculates. When ejaculations from males of the two strains were alternated (thus minimizing mating order effects), agouti males sired over 90% of the young. The time during which mating tests were conducted (relative to the occurrence of ovulation) had no effect on the relative performance of the strains. The effects of prolonged copulation and first male mating advantage demonstrated in this study are consistent with observations of mating behavior under semi-natural conditions. Thus, it appears that patterns of sperm allocation and manipulation, male and female mating strategies, and mating systems have coevolved.     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

Effects of risks of sperm competition on the numbers of eupyrene and apyrene sperm ejaculated by the moth Plodia interpunctella (Lepidoptera: Pyralidae)

This study examines the effects of different risks of sperm competition upon ejaculate characteristics in the moth Plodia interpunctella. In this short-lived species, females will remate and thus generate sperm competition, while males have a limited sperm supply. We therefore predict males to have evolved prudence in ejaculate allocation and investigate the effects of (1) rival male presence, (2) female mating history and (3) female age, upon the ejaculation of eupyrene (fertilizing) and apyrene (non-fertile) sperm numbers. We found no effect of the presence of rival males upon ejaculate characteristics, and conclude, due to the mating system of P. interpunctella, that rival males do not represent a proximate risk of sperm competition. We controlled female mating history by allowing females to receive different and predictable numbers of sperm which they then store for at least 7 days. In subsequent matings (7 days later) we found that new males ejaculated significantly more eupyrene sperm to females that had previously received larger numbers of sperm. We conclude that males increase numbers of eupyrene sperm to maintain success in sperm competition with rival sperm already in storage in the female. We found no effect of female mating history upon the ejaculation of apyrene sperm. Female age, however, had a significant negative effect upon both sperm types. We discuss these results in relation to sperm competition theory and apyrene sperm function.     

Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio

We studied the effect of relative parental investment on potential reproductive rates (PRRs) to explain sex differences in selectivity and competition in the dart-poison frog Dendrobates pumilio. We recorded the reproductive behavior of this species in a Costa Rican lowland rainforest for almost 6 months. Females spent more time on parental care than males, and `time out' estimates suggest that PRRs of males are much higher than than those of females, rendering females the limiting sex in the mating process. Males defended territories that provide suitable calling sites, space for courtship and oviposition, and prevent interference by competitors. Male mating success was highly variable, from 0 to 12 matings, and was significantly correlated with calling activity and average perch height, but was independent of body size and weight. Estimates of opportunity for sexual selection and variation in male mating success are given. The mating system is polygamous: males and females mated several times with different mates. Females were more selective than males and may sample males between matings. The discrepancy in PRRs between the sexes due to differences in parental investment and the prolonged breeding season is sufficient to explain the observed mating pattern i.e., selective females, high variance in male mating success, and the considerable opportunity for sexual selection. Received: 9 June 1998 / Received in revised form: 27 March 1999 / Accepted: 3 April 1999     

Female choice in a promiscuous wild guinea pig,the yellow-toothed cavy (<Emphasis Type="Italic">Galea musteloides</Emphasis>)

Promiscuity is traditionally considered to increase only male reproductive success but, more recently, female benefits are also assumed to be the driving force for promiscuous mating. The yellow-toothed cavy (Galea musteloides) is characterised by an extremely high degree of multiple paternity (>80%), and females have greater offspring viability after mating multiple males. However, so far it is not clear whether or not it is the female's decision that leads to mating with more than one male. To elucidate the female's role in bringing about promiscuity, female yellow-toothed cavies were given the choice between four different males each, in a mate-choice apparatus that simultaneously prevented monopolisation and harassment of the females by the males. In 10 of the 12 choice tests, mating occurred. Nine of these ten females actively sought copulations with more than one male, and their mating behaviour was displayed in a way that might have favoured the mixing of sperm. At the same time, they significantly preferred heavier and more frequently courting males. These results show that female yellow-toothed cavies are actively involved in mating with more than one male. Thus, the present study is the first to show that in a species in which females produce more viable offspring as a consequence of polyandrous mating, these females are indeed motivated to actively bring about promiscuity. Nevertheless, females are selective in the choice of their mates. Thus, both sperm competition and direct female choice seem to be important for the increased offspring viability due to promiscuous mating.     

Effect of female molt stage and sex ratio on courtship behavior of the blue crab Callinectes sapidus

In many species, males and females actively participate in courtship, and the outcome of pre-mating interactions influences the mating success of both sexes. Female blue crabs, Callinectes sapidus, mate soon after their final molt to maturity; thus female molt stage dictates the timing of mating. In a field experiment, we manipulated female molt stage and sex ratio to test their effects on the courtship behavior of both sexes, if female behavior influences the behavior and pairing success of males, and if male courtship influences male pairing-success. Early-molt-stage females avoided males during courtship, whereas late-molt-stage females sought out males. As a result, males had to pursue and capture early-molt-stage females whereas males displayed to late-molt-stage females and more easily physically controlled them. Males sometimes abandoned late-molt-stage females, but this occurred more often when females were abundant. The rate at which females avoided males was positively correlated with that of males abandoning females, and males that were unsuccessful at pairing met with higher rates of female resistance than successful males, suggesting that female behavior influences male pairing-success. Unlike unsuccessful males, successful males more often made the transition between display and maintaining physical control of the female. At high male sex ratios, males initiated courtship more readily; thus both sexual competition and female behavior influence male courtship in this species. Received: 7 July 1996 / Accepted: 10 January 1998     

Male dimorphism, sperm traits and immunology in the corkwing wrasse (Symphodus melops L.)

The evolution of alternative male mating phenotypes inevitably involves variation in risk or intensity of sperm competition. Males that typically mate in disfavoured roles (i.e. sneakers/parasitic males) are predicted to experience higher magnitudes of sperm competition than males mating in favoured roles (preferred by females/exhibiting mate monopolisation). Here, we investigated whether two distinct male phenotypes in corkwing wrasse (Symphodus melops) differ with respect to adaptations for sperm competition in terms of sperm quantity, sperm motility and fertilising capacity. We found that males exhibiting female mimicry (disfavoured role) had both absolutely and relatively larger gonads than males with typical secondary sexual characters (favoured role). Moreover, a higher proportion of sperm from female mimics was motile 5 min after activation compared to sperm of territorial males. There were, however, no differences in sperm concentration or in in vitro fertilisation ability between the two male morphs. We also examined whether the male phenotypes differed in some immune parameters and if any such variation is associated with sperm traits. Current theories predict that ejaculate quality should be negatively correlated with the level of immunocompetence. Territorial males had relatively heavier spleens than female mimics, but whether this reflects differing immunocompetence is uncertain, since the spleen is also an important organ for erythrocyte storage. Furthermore, there were no differences in the densities of circulating lymphocytes or granulocytes, and the immune parameters did not covary with the sperm traits. Differences in immunocompetence are therefore unlikely to provide a major proximate explanation for variations in sperm traits in corkwing wrasse.     

Male remating and female fitness in the wolf spider Pardosa astrigera: the role of male mating history

Although the effects of male mating history on female reproductive output and longevity have been studied in insects, few such studies have been carried out in spiders. In a mating system in which females are monandrous while males are polygynous, females may incur the risk by mating with successful males that have experienced consecutive matings and suffer from the possible depletion of sperm and/or associated ejaculates. Here, we examine the effects of male mating history on male courtship and copulation duration, female reproductive fitness, and female adult longevity of the wolf spider, Pardosa astrigera. Results indicated that male mating frequency had little effect on their subsequent copulation success, and of 35 males tested, about half of the males were able to copulate with five virgin females successively at an interval of 24 h. Male mating history had little effect on their courtship duration. However, male mating history significantly affected male copulation duration, female adult longevity, and reproductive output. Males that mated more frequently copulated longer and more likely failed to cause their mates to produce a clutch, although there was no significant difference in the number of eggs laid and the number of eggs hatched regardless of the first clutch or the second one. Multiple mating of male P. astrigera resulted in significant reduction in female adult longevity. Our results indicate that monandrous females mating with multiple-mated males may incur substantial fitness costs.     

Factors predicting male fertilization success in an external fertilizer

In postcopulatory sexual selection both sperm competition and cryptic female choice are considered to be important selective agents, but their relative importance for male fertilization success has received little attention. We tested whether sperm quality, male spawning coloration, male heterozygosity, and genetic overlap with the female explained a male’s fertilization success in controlled in vitro fertilization competition trials between equal numbers of sperm from pairs of male Arctic charr (Salvelinus alpinus), an external fertilizer. Offspring were genotyped to determine each males’ share of paternity. The velocity of a male’s sperm relative to the velocity of the competing male’s sperm was the best predictor of male fertilization success. Yet, sperm velocity was not related to spawning coloration or male heterozygosity. In fact, the most brightly colored male in a pair had the lowest fertilization probability. This could result from cryptic female choice for pale males, but might rather be a result of paler males producing more competitive sperm than more colored males. Furthermore, the more microsatellite alleles a male shared with the female relative to the competing male, the higher fertilization success he had. We argue that this latter may be an effect of assortative cryptic female choice, which might prevent hybridization with sympatric Arctic charr morphs or one form of kin selection.     

Male genital length and mating status differentially affect mating behaviour in an earwig

Phenotypic variation in male genitalia may affect copulation behaviour, which can have important fitness consequences for males. Male genitalia commonly possess traits that increase male control over copulation, but in species where females control mating, a poor functional understanding often prevents insight into the processes responsible for such effects. Here, I investigate the effect of male genital length on copulation behaviour in the earwig Euborellia brunneri, where both sexes exhibit extremely elongated genitalia that correspond in shape. This model system is particularly suitable because pairs mate repeatedly and females can limit both the number and duration of copulations. I used both virgin and mated males and females in a double-mating design because longer male genitalia confer benefits in sperm competition. Consistent with a greater predicted male mating effort in mated females, the duration of individual copulations increased, but this traded off against mating frequency as cumulative mating duration remained unchanged. In contrast, male genital length increased both individual and cumulative mating duration, regardless of mating status. This difference suggests that, while males may modify copulation duration in response to mating status, females facultatively adjust mating frequency to prevent mating excessively or express preferences for increased male genital length. Notably, this study demonstrates that male genital phenotypes that are successful in sperm competition also enjoy female-mediated mating benefits.