Can voluntary nutritional gifts in seminal flow evolve?

In many species that have internal fertilization, seminal flow includes various elements and materials in addition to the fertilizing sperm. The roles of these components are unknown. One hypothesis is that they are nutritional gifts to the female as a paternal investment. We made game theoretical models from the point of view of sperm competition among males and examined this hypothesis. The evolutionary stable strategy (ESS) of the models showed that (1) when multiple mating in the female is relatively low, gifts in seminal flow or other types of gift may evolve, at least, a male does not resist a female digesting his sperm; and (2) the ratio of nutritional gift rapidly decreases to zero with the probability or frequency of multiple mating in the female. Hence, we concluded that nutritional gifts in seminal flow can exist only in species in which sperm competition among males is rare. Furthermore, we concluded that if an element or material eminently decreases when sperm competition becomes intense, it may be a gift as a paternal investment; if not, other hypotheses concerned with more agonistic roles are more feasible. Received: November 6, 2000 / Accepted: January 10, 2001     

Love the one you're with: proximity determines paternity success in the barnacle Tetraclita rubescens

A species' mating system sets limits on the strength of sexual selection. Sexual selection is widespread in dioecious species, but is less well documented in hermaphrodites, and may be less important. We used four highly polymorphic microsatellite markers to assign paternity to broods of the hermaphroditic eastern Pacific volcano barnacle Tetraclita rubescens. These data were used to describe the species' mating system and to examine factors affecting male reproductive success. Tetraclita can sire broods over distances of 11.2 cm, but proximity to the sperm recipient had a highly significant effect on the probability of siring success. There was no effect of body size or the mass of male reproductive tissues on siring success. Broods showed relatively low frequencies of multiple paternity; even at high densities, 75% of broods had only one father. High frequencies of single‐paternity broods imply either that this species does not compete via sperm displacement, or that sperm displacement is extremely effective, potentially explaining the lack of a positive relationship between male investment and paternity. In addition, there was low variance in siring success among individuals, suggesting a lack of strong sexual selection on male traits. Low variance among sires and the strong effect of proximity are probably driven by the unusual biology of a sessile copulating species.     

Is male-infant caretaking related to paternity and/or mating activities in wild Barbary macaques (Macaca sylvanus)?

In species with a promiscuous mating system, the functions of male-infant caretaking remain unclear in the absence of genetic paternity tests. We tested paternal investment and hypotheses concerning reproductive tactics in wild groups of Barbary macaques, including results of genetic paternity tests. Our study revealed that male-infant caretaking was not related to the probability of paternity. In principle, males could use access to females to estimate paternity. However, we found that mating success was not related to paternity, so males could invest in infants that they had not sired, and caretaking of non-offspring was actually observed. Accordingly, males might be 'deceived' with respect to their paternal investment. In that case, one would expect a positive relation between mating success and the subsequent rate of male caretaking of infants. Such a relation is also lacking, leading to comprehensive rejection of the paternal investment hypothesis in Barbary macaques. By contrast, there was evidence that males showing infant care achieved higher mating frequencies than other males with the mothers of the relevant infants. Thus, male Barbary macaques do not show a 'mate-then-care' pattern, but they do exhibit a 'care-then-mate' pattern.     

Copulatory Plug Displacement Evidences Sperm Competition in Lemur catta

Male displacement of copulatory (sperm) plugs from female vaginas provides further evidence for sperm competition in ring-tailed lemurs (Lemur catta), a gregarious prosimian species with a multimale, multifemale mating system. During two mating seasons, I studied two groups of free-ranging ring-tailed lemurs on St. Catherines Island, GA, USA. I observed 22 mating pairs in which males achieved penile intromission. Copulatory plug displacement by males occurred in 9 cases. Plugs were displaced during copulation by male penes upon withdrawl following deep vaginal thrusting. In every case of copulatory plug displacement, the male displacing a plug mated to ejaculation with the estrous female. In a mating system in which females typically mate with more than one male during estrous, often in succession, copulatory plug displacement may function to disrupt or preclude other males' successful insemination of estrous females. The effects of sperm plug displacement on paternity in Lemur catta are unknown, as no study had heretofore documented copulatory plug displacement in this species. The first-male mating advantage suggested for Lemur catta should be re-evaluated where mating order is known, and copulatory plug displacement during mating, or lack thereof, is identified. Because there is a tendency for first-mating males to mate-guard for longer periods of time in Lemur catta, the latency period between the first mate's ejaculation and that of subsequent mates may be an important determinant of male fertilization success.     

Do Male Cook Strait Giant Weta Prudently Allocate Sperm?

Sperm production is costly and so males are expected to prudently allocate sperm to matings in a manner that maximizes their fitness. Sperm competition hypotheses predict that when facing increased sperm competition risk males should increase their investment in ejaculates. In contrast, when facing high future mating opportunities, males are expected to decrease their sperm investment in the current mating. This is because males should keep in reserve an amount of sperm proportional to their expected future mating opportunities. We experimentally tested whether male Cook Strait giant weta (Anostostomatidae: Orthoptera: Deinacrida rugosa) phenotypically adjust their investment in ejaculates in relation to their perceived risk of sperm competition and future mating opportunities. D. rugosa is a large flightless orthopteran insect in which males pass multiple spermatophores to females during a day-long mating bout. Contrary to expectation, we found that low female availability (i.e. increased sperm competition risk) had no effect on male resource allocation to sperm (i.e. number of spermatophores) compared to controls whereas, contrary to expectation, males experiencing high female availability increased their ejaculate investment by transferring significantly more spermatophores to their mates. Our results might be a consequence of males being insensitive to increased presence of rival males, reducing their allocation to sperm under increasingly risky circumstances, or due to females prolonging copulations when their perceived future mating opportunities are low.     

Social cues of sperm competition influence accessory reproductive gland size in a promiscuous mammal

Theory predicts that males should increase overall investment in ejaculate expenditure with increasing levels of sperm competition. Since ejaculate production is costly, we may expect males to tailor their reproductive investment according to anticipated levels of sperm competition. Here, we investigate plasticity in ejaculate investment in response to cues of population average levels of sperm competition in a promiscuous mammal, the bank vole (Myodes glareolus). We manipulated the social experience of experimental subjects during sexual development via differential exposure to the odour of rival males, to simulate conditions associated with relatively high or low average levels of sperm competition. Males exposed to a high level of competition developed larger major accessory reproductive glands (seminal vesicles) than those that experienced a low level of competition, suggesting that an increased investment in the production of copulatory plugs and/or mating rate may be beneficial at relatively high sperm competition levels. However, investment in sperm production, testis size and sperm motility were not altered according to social experience. Our findings emphasize the importance of non-sperm components of the ejaculate in mammalian postcopulatory sexual selection, and add to the growing evidence linking plasticity in reproductive traits to social cues of sperm competition.     

Transition from monogyny to polygyny in Nephila senegalensis (Araneae: Nephilidae) is not accompanied by increased investment in sperm

A common male adaptation to prevent sperm competition is the placement of a mating plug. Such plugs are considered as an extreme investment if they comprise parts of the genital systems and render the male sterile. Genital mutilation occurs in monogynous spiders of several families and may co‐occur with permanent sperm depletion, meaning that sperm production is terminated once males become mature. Within the orb‐web spider genus Nephila, monogynous mating strategies are considered ancestral, although some species have reverted to a polygynous mating strategy. Although genital mutilation does not occur in these species, permanent sperm depletion (PSD) remained. We compared investment in sperm between an effectively plugging (Nephila fenestrata Thorell, 1859) and a closely‐related nonplugging species [Nephila senegalensis (Walckenaer, 1841)]. Sperm investment should be higher in N. senegalensis because males are able to mate with several females, whereas N. fenestrata males can only achieve a maximum of two copulations, generally performed with the same female. The absence of a plugging mechanism in N. senegalensis and the inability to monopolize females by means of mating plugs results in a higher risk of sperm competition. Thus, we predicted higher investment in sperm producing tissue and larger sperm storage organs in males of N. senegalensis compared to N. fenestrata. We examined the testes and deferent ducts of both species for size and cell‐quality differences using light and transmission electron microscopy and analyzed the volume of the sperm reservoir in the male copulatory organ (i.e. spermophor) using X‐ray microcomputed tomography. In contrast to our prediction, the lumen of testes, deferent ducts, and spermophor of N. senegalensis males were significantly smaller than in N. fenestrata.     

The function of nuptial feeding in insects: a review of empirical studies

Nuptial feeding encompasses any form of nutrient transfer from the male to the female during or directly after courtship and/or copulation. In insects, nuptial gifts may take the form of food captured or collected by the male, parts, or even the whole of the male's body, or glandular products of the male such as salivary secretions, external glandular secretions, the spermatophore and substances in the ejaculate. Over the past decade, there has been considerable debate over the current function of nuptial feeding in insects. This debate has centred on the issue of whether nuptial gifts function as paternal investment (i.e. function to increase the fitness and/or number of the gift-giving male's own offspring) or as mating effort (i.e. function to attract females, facilitate coupling, and/or to maximize ejaculate transfer), although the two hypotheses are not mutually exclusive. In the present article, evidence for the potential of nuptial gifts to function as either paternal investment, mating effort, or both is reviewed for each form of nuptial feeding in each insect taxon for which sufficient data are available. Empirical evidence suggests that many diverse forms of nuptial feeding in different insect taxa function, at least in part, as mating effort. For example, nuptial prey and salivary masses in the Mecoptera, regurgitated food in Drosophila (Diptera), hind-wing feeding in Cyphoderris (Orthoptera) and the secretion of the male's cephalic gland in Neopyrochroa (Coleoptera) and Zorotypus (Zoraptera) appear to function to entice females to copulate and/or to facilitate coupling. Nuptial prey and salivary masses in the Mecoptera also appear to function to maximize ejaculate transfer (which is also a form of mating effort), as do nuptial prey in Empis (Diptera), external glandular secretions in Oecanthus and Allonemobius (Orthoptera) and the spermatophylax in gryllids and tettigoniids (Orthoptera). Large spermatophores in, for example, the Lepidoptera and Coleoptera, also appear to be maintained by selection on the male to maximize ejaculate transfer and thereby counter the effects of sperm competition. In contrast to the large amount of evidence in support of the mating effort hypothesis, there is a relative lack of good evidence to support the paternal investment hypothesis. Certain studies have demonstrated an increase in the weight and/or number of eggs laid as a result of the receipt of larger gifts, or a greater number of gifts, in tettigoniids, gryllids, acridids, mantids, bruchid beetles, drosophilids and lepidopterans. However, virtually all of these studies (with the possible exception of studies of the spermatophylax in tettigoniids) have failed to control adequately for hormonal substances in the ejaculate that are known to affect female reproductive output. Furthermore, in at least four tettigoniids (but not in the case of two species), three lepidopterans, a drosophilid and probably also bruchid beetles and bittacids, evidence suggests that the male has a low probability of fertilising the eggs that stand to benefit from his nuptial gift nutrients. Therefore, the hypothesis that paternal investment might account for the function of nuptial gifts in general is not supported.     

The unexpected but understandable dynamics of mating,paternity and paternal care in the ocellated wrasse

Although theory generally predicts that males should reduce paternal care in response to cues that predict increased sperm competition and decreased paternity, empirical patterns are equivocal. Some studies have found the predicted decrease in male care with increased sperm competition, while even more studies report no effect of paternity or sperm competition on male care. Here, we report the first example, to our knowledge, of paternal care increasing with the risk and intensity of sperm competition, in the ocellated wrasse (Symphodus ocellatus). Theory also predicts that if paternal care varies and is important to female fitness, female choice among males and male indicators traits of expected paternal care should evolve. Despite a non-random distribution of mating success among nests, we found no evidence for female choice among parental males. Finally, we document the highest published levels of extra-pair paternity for a species with exclusive and obligate male care: genetic paternity analyses revealed cuckoldry at 100 per cent of nests and 28 per cent of all offspring were not sired by the male caring for them. While not predicted by any existing theory, these unexpected reproductive patterns become understandable if we consider how male and female mating and parental care interact simultaneously in this and probably many other species.     

The male and female perspective in the link between male infant care and mating behaviour in Barbary macaques

Infant care from adult males is unexpected in species with high paternity uncertainty. Still, males of several polygynandrous primates engage in frequent affiliative interactions with infants. Two non‐exclusive hypotheses link male infant care to male mating strategies. The paternal investment hypothesis views infant care as a male strategy to maximize the survival of sired offspring, while the mating effort hypothesis predicts that females reward males who cared for their infant by preferably mating with them. Both hypotheses predict a positive relationship between infant care and matings with a particular female. However, the paternal investment hypothesis predicts that increased matings come before infant care whereas the mating effort hypothesis predicts that infant care precedes an increase in matings. Both hypotheses are usually tested from the perspective of the proportion of matings and care that individual females engage in and receive, rather than from the perspective of the care and mating behaviour of individual males. We tested the relationships between care and mating from both female and male perspectives in Barbary macaques. Mating predicted subsequent care and care predicted subsequent mating when viewed from the male but not the female perspective. Males mainly cared for infants of their main mating partners, but infants were not mainly cared for by their likely father. Males mated more with the mothers of their favourite infants, but females did not mate more with the main caretakers of their infants. We suggest that females do not choose their mating partners based on previous infant care, increasing paternity confusion. Males might try to increase paternal investment by distributing the care according to their own instead of female mating history. Further, males pursue females for mating opportunities based on previous care.     

Securing paternity in spiders? A review on occurrence and effects of mating plugs and male genital mutilation

Low female mating frequencies often appear to be cases of direct male induction that can oppose female interests. Mating plugs are most obvious means leading to low degrees of multiple mating in females. In spiders, mating plugs are formed by a variety of amorphous materials, by the breakage of the male sperm transferring organ, or by the whole male that functions as a mating barrier. Our compilation of the available information on the presence of the various types of mating plugs suggests that plugs predominantly occur in entelegyne spiders. In this group, plugs do not interfere with oviposition since separate openings for insemination and oviposition are present. In contrast, mating plugs seem to be rare in haplogyne spiders that do not possess separate openings. The available experimental studies on the function of the different types of plugs suggest that plugs can be considered as male adaptations to avoid sperm competition. However, females in some cases were shown to have evolved means to prevent or control male manipulation or may selectively favour plug production in specific males, an aspect which has largely been neglected. In order to understand plug evolution and function we need to explore the morphological, behavioural and biochemical aspects involved and extend our approach to interactions between the sexes.     

Mating Plugs in Polyandrous Giants: Which Sex Produces Them,When, How and Why?

Background

Males usually produce mating plugs to reduce sperm competition. However, females can conceivably also produce mating plugs in order to prevent unwanted, superfluous and energetically costly matings. In spiders–appropriate models for testing plugging biology hypotheses–mating plugs may consist of male genital parts and/or of amorphous covers consisting of glandular or sperm secretions. In the giant wood spider Nephila pilipes, a highly sexually dimorphic and polygamous species, males are known to produce ineffective embolic plugs through genital damage, but nothing is known about the origin and function of additional conspicuous amorphous plugs (AP) covering female genitals.

Methodology

We tested alternative hypotheses of the nature and function of AP in N. pilipes by staging mating trials with varying degrees of polyandry. No APs were ever formed during mating trials, which rules out the possibility of male AP formation. Instead, those females that oviposited produced the AP from a liquid secreted during egg sac formation. Polyandrous females were more likely to lay eggs and to produce the AP, as were those that mated longer and with more total insertions. Our further tests revealed that, in spite of being a side product of egg sac production, AP, when hardened, prevented any subsequent copulation.

Conclusions

We conclude that in the giant wood spider (Nephila pilipes), the amorphous mating plugs are not produced by the males, that repeated copulations (most likely polyandrous) are necessary for egg fertilization and AP formation, and that the AP represents a female adaptation to sexual conflict through prevention of unwanted, excessive copulations. Considering the largely unknown origin of amorphous plugs in spiders, we predict that a similar pattern might be detected in other clades, which would help elucidate the evolutionary interplay of various selection pressures responsible for the origin and maintenance of mating plugs.     

Adult Responses to Larval Population Size in the Almond Moth, Cadra cautella

Juvenile population size may affect the potential for future mating opportunities and therefore potentially sperm competition; this may favour ontogenetic adjustments in sperm production. Theory predicts that males should optimize their ejaculatory investment in accordance with the risk of sperm competition. Evidence for these theories is typically revealed in males of highly polyandrous species. Whether such responses to environmental cues exist for females, or are maintained in mildly polyandrous species in which most females do not re-mate, is unknown. Male lepidopterans produce normal, fertilizing sperm (eupyrene) and non-fertilizing (apyrene) sperm. Apyrene sperm are associated with reduced female receptivity, suggesting a role in sperm competition. We tested the effect of juvenile population size on life-history parameters and reproductive investment in the mildly polyandrous almond moth, Cadra cautella , a species in which current male ejaculate traits suggest previous selection for paternity protection consistent with a sperm-competitive environment. Larvae were reared at high (H) or low population sizes (L). We recorded larval development time, adult longevity and male gametic investment. Our results show a response by adults to signals in the juvenile environment. H males transferred more apyrene, but not eupyrene sperm. We also examined potential trade-offs between somatic characters and reproductive behaviours. Larval duration was longer for H individuals, females and heavier individuals. Further, H females and L males lived longer than L females. Our data are consistent with the theory that males should adjust their reproductive investment in accordance with sperm competition risk.     

Value of male remating and functional sterility in redback spiders

In the Australian redback spider, Latrodectus hasselti, males typically use their paired copulatory organs (palps) to copulate twice with a single female then sacrifice themselves to their cannibalistic mates in a strategy that increases their paternity in that one mating, but leads to death. This type of terminal investment in one mating is predicted only if the expected value of future matings is low for males relative to the value of repeated mating with the same female. In this laboratory study, we quantified the reproductive value of mating more than once with the same female (repeated mating) and mating with more than one female (multiple mating) for male redback spiders. We tested two natural selection hypotheses for repeated mating, sperm limitation and reproductive insurance, but found no support for either hypothesis. We show that, in the absence of sperm competition or cannibalism, male lifetime reproductive output is the same whether a male copulates once, twice, or several times with a given female. Repeated mating does not increase the probability of successful fertilization, nor does it increase the number of offspring produced in successful matings. Although male repeated mating is not favoured because of increased fertility of mates, other studies suggest it may be important in sperm competition. Here we show that the relative reproductive value of the first two copulations is very high for redback males because they are functionally sterile after each palp has been used once; nonvirgin males are unable to father offspring. Functional sterility and repeated mating by male redbacks may be favoured by the same factors that lead to male sacrifice behaviour: ecological constraints on multiple mating combined with competitive benefits of maximal investment in the first mating. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Effects of mating status on copulation investment by male bushcricket Gampsocleis gratiosa (Tettigoniidae, Orthoptera)

Male’s copulation investment, including spermatophore and sperm investment were very high in the Chinese bushcricket Gampsocleis gratiosa. The effects of mating status of both males and females on male’s copulation investment were examined in this study. The fresh weight of spermatophylax increased positively with the weight of males’ body. This indicated that the nutritional investment during copulation depended on male’s quality. Spermatophore investment showed insignificant differences in every copulation protocols. This finding supported the paternal investment hypothesis, that is, males contributed to their offspring with little attention to their partners. Sperm releasing per ejaculation varied significantly among the trials. Males decreased 54.19% sperm in second mating than in its first mating, demonsrated that males regarded the first mating highly, and were more prudent in subsequent mating. These males’ strategies may contribute to the viability of the offspring.     

Effects of mating status on copulation investment by male bushcricket Gampsocleis gratiosa (Tettigoniidae, Orthoptera)

Male’s copulation investment, including spermatophore and sperm investment were very high in the Chinese bushcricket Gampsocleis gratiosa. The effects of mating status of both males and females on male’s copulation investment were examined in this study. The fresh weight of sper-matophylax increased positively with the weight of males’ body. This indicated that the nutritional in-vestment during copulation depended on male’s quality. Spermatophore investment showed insig-nificant differences in every copula...~     

A mating plug protein reduces early female remating in Drosophila melanogaster

Mating plugs are formed within the female reproductive tract during mating from male ejaculate constituents or even from male genitalia themselves. Across species, mating plugs have roles in sperm storage and the prevention of female remating. In the fruitfly Drosophila melanogaster, accessory gland proteins such as the sex peptide are known to reduce female remating, however this effect can take some time to establish, hence other ejaculate components must also be involved. We hypothesised a role for the PEBII mating plug protein in the prevention of early female remating. Using RNA interference we produced PEBII knockdown males. We found that these males were significantly less able to prevent female remating in the 4 h following mating. The mating plugs produced by PEBII knockdown males also showed lower levels of autofluorescence in the first 10 min after the start of mating, suggesting they differed in composition to those of control males. Reduced levels of PEBII had no effect, however, on fecundity, progeny production or egg-adult viability in the first 24 after mating, suggesting there were no short-term effects of PEB II on sperm transfer, storage or use. Our results show that PEBII has a subtle but significant role in the prevention of early female remating.     

The evolution of sperm length in moths

Sperm form and function remain poorly understood despite being of fundamental biological importance. An instructive approach has been to examine evolutionary associations across comparable taxa between sperm characters and other, potentially selective reproductive traits. We adopt this approach here in a comparative study examining how sperm lengths are associated with male and female reproductive characters across moths. Primary data have revealed Lepidoptera to be an ideal order for examination: there is profound variation in the dimensions (but not organization) of the reproductive traits between closely related species which all share a monophyletic ancestry, for example, eupyrene sperm length varies from 110 to 12,675 microm. Eupyrene (normal fertilizing) and apyrene (anucleate and non-fertile) sperm lengths are positively correlated across taxa and both sperm types show positive associations with mating pattern (as measured by the residual testis size). At fertilization, eupyrene sperm must migrate down the often elongated female spermathecal duct from storage to unite with the ovum. Across taxa, the elongation of this duct is associated with increased eupyrene sperm length, suggesting a positive female influence on sperm size since longer, more powerful sperm may be selected to migrate and/or compete successfully down greater ductal lengths. Apyrene sperm length is not associated with female reproductive tract dimensions. However, we found a positive relationship between the residual testis volume and spermathecal volume, suggesting coevolution between male investment in spermatogenesis and the extent of the female sperm storage capacity. Within males, there is a positive association between the two organs which form the ejaculate-containing spermatophore: the testes and the accessory gland. The 'trade-up' in investment to these components is discussed in relation to paternal investment and mating patterns.     

The timing and interval of mate encounter affects investment during mating

The benefits obtained from mating are usually condition‐dependent, favouring the evolution of flexible investment during copulation; for example, in terms of invested time, energy or sperm. Flexible investment strategies are predicted to depend on the likelihood of acquiring alternative mates and therefore they should depend on the timing of mate encounter. However, scarce experimental evidence for this hypothesis exists. In the present study, we manipulated the time delay until first mating and the interval between first and second mating in the polygynandrous common lizard Zootoca vivipara. We determined treatment effects on fertilization success and copulation duration, with the latter being a proxy for investment in mating and for the quantity of transferred sperm. The duration of the second copulation decreased with increasing inter‐mating interval and depended on the fertilization success of first mates. The former provides evidence for time‐dependent investment strategies, most likely resulting from the progression of the female's reproductive cycle. The fertilization success of first mates increased with increasing inter‐mating interval and was higher when females were closer to ovulation, showing that flexible investment strategies significantly affected male reproductive success. This indicates fertilization assurance, which may mitigate the negative effects of low population density on reproductive success (e.g. Allee effects).     

Ejaculate depletion patterns evolve in response to experimental manipulation of sex ratio in Drosophila melanogaster

We assessed the extent to which traits related to ejaculate investment have evolved in lines of Drosophila melanogaster that had an evolutionary history of maintenance at biased sex ratios. Measures of ejaculate investment were made in males that had been maintained at male-biased (MB) and female-biased (FB) adult sex ratios, in which levels of sperm competition were high and low, respectively. Theory predicts that when the risk of sperm competition is high and mating opportunities are rare (as they are for males in the MB populations), males should increase investment in their few matings. We therefore predicted that males from the MB lines would (1) exhibit increased investment in their first mating opportunities and (2) deplete their ejaculates at a faster rate when mating multiply, in comparison to FB males. To investigate these predictions we measured the single mating productivity of males from three replicates each of MB and FB lines mated to five wild-type virgin females in succession. In contrast to the first prediction, there was no evidence for differences in productivity between MB and FB line males in their first matings. The second prediction was upheld: mates of MB and FB males suffered increasingly reduced productivity with successive matings, but the decline was significantly more pronounced for MB than for FB males. There was a significant reduction in the size of the accessory glands and testes of males from the MB and FB regimes after five successive matings. However, the accessory glands, but not testes, of MB males became depleted at a significantly faster rate than those of FB males. The results show that male reproductive traits evolved in response to the level of sperm competition and suggest that the ability to maintain fertility over successive matings is associated with the rate of ejaculate, and particularly accessory gland, depletion.