Piercing the partner's skin influences sperm uptake in the earthworm Lumbricus terrestris

Sexual conflict between mating partners can give rise to strategies that are advantageous for one sex but harmful to the opposite sex. Usually, sperm donors develop (offensive) traits to enhance their chances in sperm competition, while sperm recipients evolve (defensive) traits that allow them to stay in control of who fathers their offspring. Here, we demonstrate that these processes are also at work in simultaneous hermaphrodites. The hermaphroditic earthworm Lumbricus terrestris uses 40 to 44 copulatory setae to pierce into its partner's skin, causing damage and injecting a substance from its setal glands. Experimental injection of the gland substance indicates that a refractory period may be induced. More importantly, removal of the copulatory setae shows that they influence the partner's sperm uptake. When the setae are present, more sperm are taken up and sperm are distributed more equally over the four spermathecae. We interpret this as a strategy that stacks the odds for the donor's sperm in fertilizing cocoons.     

Among- and within-population variation in sperm quality in the simultaneously hermaphroditic land snail Arianta arbustorum

Sperm competition models on the evolution of sperm size assume associations with another sperm quality trait, sperm longevity. Sperm length can also provide an indication of possible mechanisms affecting motility and thus fertilization success. Despite their importance, however, detailed mechanisms of sperm competition at the gamete level are poorly understood. In simultaneously hermaphroditic land snails, sperm traits and cryptic female choice are assumed to be crucial in determining fertilization success. We examined the variation in sperm length and number among individuals from four natural populations of the land snail Arianta arbustorum, a species with multiple mating and long-term sperm storage. We also assessed variation in velocity, motility and longevity of sperm in snails from two of the four populations. Independent of shell size, sperm length differed among populations and, to a minor extent, even among individuals within populations. Mean sperm length of a snail was not correlated with the number of sperm delivered in a spermatophore. The mean sperm velocity (=VCL) did not differ between snails from two populations. However, VCL varied among snails. Percentage motility and longevity of sperm differed between snails from the two populations. No correlations were found between length, velocity, percentage motility and longevity of sperm. To conclude, individual snails differed in sperm quality, and this variation may partly explain the differential fertilization success between A. arbustorum snails. Moreover, our findings did not support the positive association between sperm length and longevity assumed by sperm competition models for internally fertilizing species.     

Cloacal erection promotes vent apposition during forced copulation in the New Zealand stitchbird (hihi): implications for copulation efficiency in other species

Cloacal protuberances (CP) in male birds result from spermatic engorgement of storage tubules around the cloaca during the breeding season. We examined seasonal changes in the volume and orientation of the CP in the New Zealand stitchbird Notiomystis cincta. The male stitchbird has one of the largest recorded CPs for any species (max = 1,570 mm³), with CP volume increasing by almost 400% between the non-breeding and breeding seasons. While sperm competition has been positively correlated with the magnitude of CP storage in other species, no evidence previously existed for the CP improving copulation efficiency. By measuring the relative orientation of the CP throughout the year, we show that not only does the CP increase in size as males become sexually active, it also changes its orientation by approximately 60°. This results in it shifting from facing posteriorly to becoming almost perpendicular to the abdomen. This cloacal erection improves the apposition of the male and female cloacal openings during face-to-face forced copulation in this species. This provides the first reported evidence supporting the copulation efficiency hypothesis of the avian CP. While the magnitude of seasonal changes in female cloacal volume was similar to males, female cloacal orientation remained virtually unchanged across seasons. This difference between the sexes is likely to reflect differing selection pressures for optimizing sperm transfer. In females, a posterior-facing cloaca is ideal for both waste evacuation and sperm reception, whereas, for the male, a posterior-facing cloaca is well suited for waste evacuation, but possibly hinders sperm delivery. Changes in male cloacal orientation from the non-breeding to the breeding season are a likely reflection of conflict in this dual function. Evidence of changes in CP orientation in another passerine species suggests this phenomenon is widespread and also important for understanding related fields such as sperm competition, forced copulation and constraints on the evolution of the avian intromittent organ.     

Hermaphrodite sex role preferences: the role of partner body size, mating history and female fitness in the sea slug Chelidonura sandrana

Costs and benefits associated with matings and the effects of mating frequency on fitness commonly differ between the sexes. As a result, outcrossing simultaneous hermaphrodites may prefer to copulate in the more rewarding sex role, generating conflicts over sperm donation and sperm receipt between mates. Because recent sex role preference models remain controversial, we contrast here some of their assumptions and predictions in the sea slug Chelidonura sandrana. For this hermaphrodite with sperm storage and internal fertilisation, risk-averse models assume that fitness pay-offs are constantly higher in the female than in the male function in any single mating. Moreover, excluding mutual partner assessment, these models predict male mating behaviour to be independent of receiver traits. The competing gender ratio hypothesis assumes that relative fitness pay-offs, and thus the preferred mating roles, vary and may reverse between matings and predicts that ejaculation strategies co-vary with receiver quality. We found that field mating rates of C. sandrana substantially exceeded what is required to maintain female fertility and fecundity, indicating large variation in direct female benefits between matings. We further demonstrate that male copulation duration adaptively increased with partner body size (i.e. fecundity) but decreased with recent partner promiscuity. These findings are compatible with the gender ratio hypothesis but contradict risk-averse models.     

Copulatory plug displacement and prolonged copulation in the Iberian rock lizard (Lacerta monticola)

Male Iberian rock lizards (Lacerta monticola) produce copulatory plugs that adhere firmly inside the female cloaca and occlude both oviducts. These plugs do not prevent rival male insemination, as they neither reduce female attractiveness or receptivity to rival males nor do they function as chastity belts. Prior to copulation, males bite various regions of the female body, including the cloacal region. A previous plug in the female cloaca is expelled only after rival male intromission. We hypothesized that: (1) such male pre-copulatory behaviours function to loosen plug adherence to the female cloaca, thereby facilitating intromission, and that (2) the hemipenis plays a role in displacing a previous plug prior to the delivery of a new plug. Neither of these hypotheses was supported. Instead, our results indicate that rival males can intromit the hemipenis past a previous plug in the female cloaca and deliver their own plug underneath it. Consequently, previous plugs are pushed away from the oviductal openings and even dislodged from the female cloaca. Copulation duration was determined both by the time used by males to deliver a plug and by the fact that males prolonged copulation beyond plug delivery. There seemed to be sexual conflict over prolonged copulation, which was resolved by the male/female head-length ratio. The adaptive value of Iberian rock lizard copulatory plugs and prolonged copulation in Iberian rock lizards is discussed in the context of sperm competition.Communicated by S. Downes     

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.     

Determinants of paternity success in the spider Pholcus phalangioides (Pholcidae: Araneae): the role of male and female mating behaviour

In double mating experiments, we examined whether and to what extent various male and female behavioural traits influence the course of mating and fertilization success in the cellar spider. In males, we focussed on pre-copulatory behaviour and on the rhythmic twisting movements that the male performs with his pedipalps during copulation. In females, we investigated remating decisions and the effect of female termination of copulation. Second males fertilized a high proportion of the eggs (P ₂: median 89%) despite much shorter second matings, with high variation in relative paternity success. The number of pedipalp movements (PPMs) of either male was a better predictor of paternity than copulation duration. Our results suggest that in second matings, PPMs help to remove sperm from previous males, whereas in first matings a high number of PPMs enhances fertilization success, either due to numerical sperm competition or cryptic female choice. Furthermore, we found a negative male age effect on paternity in second matings, implying that age-related deterioration of spermatozoa may promote variation in fertilization success. Female receptivity decreased significantly in second matings; only 70% of the females remated. Females that accepted a second copulation were found to terminate these much earlier and with higher probability than first matings. This suggests that the intensity of conflict between the sexes is higher in second matings. Increased intensity of sexual conflict may be responsible for stronger selection on male traits, as pre-copulatory behaviour and age only affected male copulatory performance and paternity in second matings. Electronic Publication     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

Sex allocation is an important reproductive decision for parents. However, it is often assumed that females have substantial control over sex allocation decisions, and this is particularly true in haplodiploid insects, in which females apparently determine sex by deciding whether to fertilise an egg (and produce a diploid daughter) or not (and produce a haploid son). Mechanisms by which males may influence sex allocation are not so straightforward, and their potential influence on sex ratios has been somewhat neglected. Here, we test whether males influence offspring sex ratios in the parasitoid wasp Nasonia vitripennis. We show that some of the variation in observed sex ratios can be attributed to males when comparing the affect of male strain on sex ratio. We did not find among-male variation in sex ratio with a less powerful experiment using males from only one strain or an effect of male mating environment. Our data suggest that males can influence female sex ratios and contribute to the variation around the sex ratios optimal for females. However, the influence is not large, suggesting that females have more influence on sex allocation than do males. We conclude by considering whether male influences on sex ratio represent differences in male reproductive competence or deliberate attempts by males to increase their fitness by influencing daughter production.     

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.     

Patterns of sperm use in the scorpionfly Panorpa germanica L. (Mecoptera: Panorpidae)

Sperm competition can be a powerful selective force in the evolution of reproductive strategies and mating systems. In studies on sperm competition, patterns of sperm use are typically reported as the mean species value of P ₂, determined as the proportion of offspring sired by the second male to copulate with a doubly mated female. However, the within-species variance in P ₂ has mostly been ignored, although taking this variance into account may be crucial for understanding the underlying mechanisms of sperm competition. Paternity analysis among the offspring of doubly mated females of Panorpa germanica (Mecoptera, Panorpidae) revealed a relationship between relative copulation durations of both males and the proportion of offspring each male will sire. This correlation between proportional copulation durations and paternity suggests mixing of sperm from different males inside the female’s spermatheca. Yet, sperm mixing appears to be incomplete, as paternity was overall slightly shifted towards the second male on average fathering a higher proportion of the offspring than its relative copulation duration would predict in case of complete sperm mixing. For individual males, however, the outcome of sperm competition is rather unpredictable as the intraspecific variance in P ₂ was found to be very high, irrespective of copulation durations. Possible causes of the observed variance in P ₂ and the partial last male sperm precedence are discussed.     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

Male-male interference competition decreases spawning rate in the European bitterling (<Emphasis Type="Italic">Rhodeus sericeus</Emphasis>)

We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.Communicated by J. Krause     

Effect of sperm depletion and starvation on female mating behavior in the water strider,Aquarius remigis

The resolution of intersexual conflict over mating should be dependent on the current state of each individual. In this study, I used a factorial design to examine the influence of two physiological factors, sperm depletion and food deprivation, on resistance to mating by females of the water strider, Aquarius remigis. Females employ several different mate-resisting tactics during an encounter with a male. Five measures of female resistance to mating were identified: jumping, rolling, dunking, time spent dunking, and struggle duration. Jump, roll, and dunk rates were highly correlated with each other and combined into one metric of resistance to mating (PC1) using principal components analysis. Time per dunk (T/D) and struggle duration were also analyzed. Discrete male behaviors during the struggle could not be identified. Two measures of female resistance, PC1 and T/D, were significantly lower in sperm-depleted females than in sperm-replenished females. Struggle duration did not differ between the two treatments. Starvation had no effect on any of the measures of resistance. Sperm depletion significantly enhanced the probability of mating (54% vs. 24% for replenished females), while starvation had no effect on the probability of mating. I pooled all the females and compared females that mated with those that did not mate. Nonmating females resisted significantly more than mating females in all three measures of resistance. Path analysis indicated that PC1 was the only measure of resistance that was significantly negatively related to the probability of mating. Almost half (46%) of sperm-depleted females showed no resistance to males, while only 3% of sperm-replenished females were nonresistant. When nonresisters were removed from the analysis, sperm depletion had no effect on any of the measures of female resistance to mating and no effect on the probability of mating. In A. remigis, female resistance appears to be a yes/no phenomenon with respect to sperm depletion and not affected directly by starvation. Received: 2 September 1994/Accepted after revision: 9 September 1995     

Phenotypic correlates and survival consequences of male mating success in lek-mating greater prairie-chickens (<Emphasis Type="Italic">Tympanuchus cupido</Emphasis>)

Female choice and male–male aggression are two modes of sexual selection that can lead to elaboration of male morphological and behavioral traits. In lek-mating species, male mating success is often strongly skewed, and it is puzzling why variation in male traits is still observed given directional female choice. If male traits correlated with reproductive success are honest signals of male quality, there may be survival costs associated with the expression of those traits. In this study, we examined whether morphological, behavioral, and territorial traits are correlated with male mating success and survival in the lek-mating greater prairie-chicken (Tympanuchus cupido). We introduce a novel application of multinomial discrete choice models for analysis of female mate choice behavior. We found that behavioral and territorial attributes showed 6.5 times more variability among males than morphological traits. Both display and aggressive behaviors were strong predictors of male mating success, suggesting that both female choice and male–male aggression were important in determining mating success among male greater prairie-chickens. Moreover, annual survival of male prairie-chickens was independent of mating success and male traits. Females appear to be choosing males based on behavioral traits where large variation exists between males (coefficient of variation >30%). Behavioral traits were the most important factor in determining mating success of male prairie-chickens, but the mechanism underlying this relationship is unknown. In the future, experimental manipulations of male hormones or parasite loads could bridge the proximate mechanisms and ultimate consequences of factors mediating male mating success in lek-mating grouse.     

Female mediation of sperm competition in the millipede Alloporus uncinatus (Diplopoda: Spirostreptidae)

The tropical spirostreptid millipede Alloporus uncinatus has a polygynandrous mating pattern that is apparently shaped by sperm competition (Telford and Dangerfield 1990, 1993a). In the present study radioisotopic labelling of ejaculates was used to quantify the temporal effects of double mating sequences on sperm precedence patterns. Ejaculates of successive males mix completely within the sperm storage organs and are concentrated close to the site of fertilisation, the spermathecal-oviduct junction. When matings follow each other immediately (experiment 1), equal volumes of the ejaculates of each male are stored within the spermathecae, and both males have equal probability of paternity. Where matings are separated by a 24-h delay (experiment 2), the volumetric contribution of the first male is reduced by 54.8 % and last male precedence operates. Although the distal ends of the gonopods play a primary role in the redistribution of ejaculates (Barnett and Telford 1994), by using a single mating sequence we show that they are not responsible for the reduction in ejaculate volume. The most likely explanation is absorption and/or ejection of the first male's ejaculate by the female. Genital functional morphology is used to reconstruct the mechanism of ejaculate distribution.     

The cost of sexual coercion and heterospecific sexual harassment on the fecundity of a host-specific, seed-eating insect (Neacoryphus bicrucis)

Two species of seed-eating true bugs, Neacoryphus bicrucis (Heteroptera: Lygaeidae) and Margus obscurator (Heteroptera: Coreidae) co-occur on ragwort, Senecio tomentosus, in southern Georgia, USA. Males of both species sometimes engage in chases and protracted grappling with females that flee initial mountings. Sometimes genital coupling occurs while the wriggling female is restrained in the male's grasp. Chases, grappling, and mounting attempts are misdirected toward heterospecific females, heterospecific males, or conspecific males. In a laboratory study, confinement of mated N. bicrucis females with either conspecific or heterospecific males reduced fecundity by approximately one-half relative to mated females confined only with other females. Perhaps as a consequence of this, N. bicrucis females frequently leave areas of high host plant density, where they prefer to oviposit, when males are abundant. The abundance of each species is positively correlated with host plant density but the two species rarely occur together on the same plants. This may be an effect of heterospecific courtship which induces the flight of N. bicrucis more than the flight of M. obscurator. The laboratory results suggest that copulations following chases and grappling represent sexual harassment, not a mechanism of active female choice for a vigorous mate. As sexual harassment imposes high fitness costs that favor abandonment of host plants, it may, when misdirected, incidentally limit habitat use by ecologically similar species. Received: 3 October 1998 / Received in revised form: 21 January 1999 / Accepted: 14 February 1999     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

Differential use of stored sperm during oviposition in the damselfly Calopteryx splendens xanthostoma (Charpentier)

Female Calopteryx splendens xanthostoma (Charpentier) (Odonata), have two discrete sperm storage organs, the bursa copulatrix and the paired spermathecae. We used the Random Amplified Polymorphic DNA(RAPD) profiling technique to determine from which sperm storage organ spermatozoa were used to fertilise eggs during behaviourally distinct oviposition bouts. During oviposition bouts following remating sperm from the bursa copulatrix are largely used to fertilise eggs, but when females avoid remating before an oviposition bout the sperm used to fertilise eggs are derived mainly from the paired spermathecae. Our observations suggest that female C. s. xanthostoma can avoid male paternity assurance mechanisms by a combination of behavioural and anatomical adaptation: the reason(s) they do so is unclear. Received: 28 November 1995/Accepted after revision: 5 August 1996     

Patterns of sexual cohabitation and female ejaculate storage in the American lobster (<Emphasis Type="Italic">Homarus americanus</Emphasis>)

Little is known of the time and ejaculate allocation strategies during mating of American lobster, Homarus americanus. This study investigated sexual cohabitation and female ejaculate accumulation patterns in a laboratory mating experiment, as well as female seminal receptacle load in exploited populations in the waters of the Magdalen and Anticosti Islands, in eastern Canada. In the laboratory experiment, the length of sexual cohabitation was proportionate to female size for large but not for small males. Also, large males cohabited with pre- and postmolt females longer than small males. These different time investment strategies can be explained by different mutual benefits. In the field and laboratory, larger females accumulated more ejaculate than smaller ones. This suggests that male lobsters tailor ejaculate to female size, a reliable index of her reproductive potential. Moreover, similarly-sized females accumulated more ejaculate when mated with large compared to small males. Comparison of receptacle loads between wild-mated and laboratory-mated females suggests that the former were mating mainly with smaller males, although some evidence of positive size-assortative mating existed, especially at the less exploited Anticosti site. The results are discussed in the context of evolutionary theory and of proposed management measures to increase egg production in exploited populations.Communicated by T. Czeschlik     

Mate preferences by male guppies (Poecilia reticulata) in relation to the risk of sperm competition

We investigated male mate preferences in relation to the perceived risk of sperm competition in the guppy (Poecilia reticulata), a freshwater fish with a promiscuous mating system. Our laboratory experiments revealed that male mate choice behaviour is not influenced by the presence of rival males that are merely in close proximity to a potential mate, as there was no significant difference in the amount of time that males spent with females that were recently either alone or in close proximity to four rival males. Male mate choice behaviour was, however, strongly influenced by the presence of rival males in a second experiment, where those rivals were permitted to copulate with one of the females. In that situation, males spent significantly more time with, and directed significantly more sigmoid courtship displays toward, females that they had recently seen alone compared to females they had seen receiving forced copulations from up to four rival males. Our results therefore indicate that male guppies are sensitive to the risk of sperm competition and alter their mate choice behaviour in an adaptive fashion.Communicated by K. Lindström