THE EFFECTS OF PREDATION RISK ON MATING SYSTEM EXPRESSION IN A FRESHWATER SNAIL

Environmental effects on mating system expression are central to understanding mating system evolution in nature. Here, I report the results from a quantitative‐genetic experiment aimed at understanding the role of predation risk in the expression and evolution of life‐history and mating‐system traits in a hermaphroditic freshwater snail (Physa acuta). I reared 30 full‐sib families in four environments that factorially contrast predation risk and mate availability and measured age/size at first reproduction, growth rate, a morphological defense, and the early survival of outcrossed/selfed eggs that were laid under predator/no‐predator conditions. I evaluated the genetic basis of trade‐offs among traits and the stability of the G matrix across environments. Mating reduced growth while predation risk increased growth, but the effects of mating were weaker for predator‐induced snails and the effects of predation risk were weaker for snails without mates. Predation risk reduced the amount of time that individuals waited before self‐fertilizing and reduced inbreeding depression in the offspring. There was a positive among‐family relationship between the amount of time that individuals delayed selfing under predation risk and the magnitude of inbreeding depression. These results highlight several potential roles of enemies in mating‐system expression and evolution.     

Life-history plasticity and inbreeding depression under mate limitation and predation risk: cumulative lifetime fitness dissected with a life table response experiment

Environmental effects on the evolution of mating systems are increasingly discussed, but we lack many examples of how environmental conditions affect the expression and consequences of alternative mating systems. Variation in mate availability sets up a trade-off between reproductive assurance and inbreeding depression, but the consequences of both mate limitation and inbreeding may depend on other environmental conditions. Predation risk is common under natural conditions, and known to affect allocation to reproduction, but we know little about the effects of isolation and inbreeding under predation risk. We reared selfed and outcrossed hermaphroditic freshwater snails (Physa acuta) in four environments (predator cues present or absent crossed with mating partners available or not) and quantified life-history traits and cumulative lifetime fitness. Our results confirm that isolation from mates can increase longevity and growth, resulting in higher lifetime fecundity. Thus, we observed no evidence for mate limitation of reproduction. However, reproduction under isolation (i.e., selfing) resulted in inbreeding depression, which should counteract the benefits of selfing. Inbreeding depression in fitness occurred in both predator and no-predator environments, but there was no overall change in inbreeding depression with predator cues. This represents, to our knowledge, the first empirical estimate of the effect of predation risk on inbreeding depression in an animal. Cumulative fitness was most influenced by early survival and especially early fecundity. As predation risk and inbreeding (both ancestral and due to a lack of mates) reduced early fecundity, these effect are predicted to have important contributions to population growth under natural conditions. Therefore life-history plasticity (e.g., delayed reproduction) is likely to be very important to overall fitness.     

Mate Familiarity Affects Pairing Behaviour in a Long‐Term Monogamous Lizard: Evidence from Detailed Bio‐Logging and a 31‐Year Field Study

Long‐term monogamy is most prevalent in birds but is also found in lizards. We combined a 31‐year field study of the long‐lived, monogamous Australian sleepy lizard, Tiliqua rugosa, with continuous behavioural observations through GPS data logging, in 1 yr, to investigate the duration of pair bonds, rates of partner change and whether either the reproductive performance hypothesis or the mate familiarity hypothesis could explain this remarkable long‐term monogamy. The reproductive performance hypothesis predicts higher reproductive success in more experienced parents, whereas the mate familiarity hypothesis suggests that effects of partner familiarity select for partner retention and long‐term monogamy. Rates of partner change were below 34% over a 5‐yr period and most sleepy lizards formed long‐term pair bonds: 31 partnerships lasted for more than 15 yr, 110 for more than 10 yr, and the recorded maximum was 27 yr (ongoing). In the year when we conducted detailed observations, familiar pairs mated significantly earlier than unfamiliar pairs. Previous pairing experience (total number of years paired with previous partners) had no significant effect. Early mating often equates to higher reproductive success, and we infer that is the case in sleepy lizards. Early mating of familiar pairs was not due to better body condition. We propose two suggestions about the proximate mechanisms that may allow familiar pair partners to mate earlier than unfamiliar partners. First, they may have improved coordination of their reproductive sexual cycles to reach receptivity earlier and thereby maximise fertilisation success. Second, they may forage more efficiently, benefiting from effective information transfer and/or cooperative predator detection. Those ideas need empirical testing in the future. Regardless of the mechanism, our observations of sleepy lizard pairing behaviour support the mate familiarity hypothesis, but not the reproductive performance hypothesis, as an explanation for its long‐term monogamous mating system.     

Mate choice in a hermaphrodite: you won't score with a spermatophore

Sexual selection has played a major role in shaping the wide variety of mating patterns found in species with separate sexes, but little is known about its effects on simultaneous hermaphrodites. However, many hermaphrodites possess complex reproductive systems and mating behaviour is often elaborate, suggesting that some form of mate assessment takes place. We found that the marine slug Aeolidiella glauca, a simultaneous hermaphrodite with reciprocal external sperm transfer via spermatophores, shows a unique mate choice behaviour by avoiding mating with conspecifics already carrying a spermatophore received during the previous mating. Current mating status did not seem to affect this behaviour, because both slugs that had mated 2-3 days before our mate choice trials and slugs that had been isolated for 4-6 weeks avoided spermatophore-carrying partners. There are two obvious reasons why slugs should avoid recently mated partners. First, they may reduce the risk of getting a partner depleted in self-sperm. Second, the risk of sperm competition may be decreased. Histological investigations of sperm reserves suggest that sperm depletion did not influence our mate choice experiments. Most slugs had sufficient sperm stored for spermatophore production. Therefore, the most likely explanation for A. glauca's peculiar mate choice is that, by avoiding a recently mated partner, a sperm donor may reduce its risk of being subjected to sperm competition.     

Females of the Cellar Spider Discriminate Against Previous Mates

Mate choice for novel partners should evolve when remating with males of varying genetic quality provides females with fitness‐enhancing benefits. We investigated sequential mate choice for same or novel mating partners in females of the cellar spider Pholcus phalangioides (Pholcidae) to understand what drives female remating in this system. Pholcus phalangioides females are moderately polyandrous and show reluctance to remating, but double‐mated females benefit from a higher oviposition probability compared to single‐mated females. We exposed mated females to either their former (same male) or a novel mating partner and assessed mating success together with courtship and copulatory behaviours in both sexes. We found clear evidence for mate discrimination: females experienced three‐fold higher remating probabilities with novel males, being more often aggressive towards former males and accepting novel males faster in the second than in the first mating trial. The preference for novel males suggests that remating is driven by benefits derived from multiple partners. The low remating rates and the strong last male sperm precedence in this system suggest that mating with novel partners that represent alternative genotypes may be a means for selecting against a former mate of lower quality.     

Age‐dependent effects of predation risk on reproductive success in a freshwater snail

Reproductive performance is often age‐dependent, showing patterns of improvement and/or senescence as well as trade‐offs with other traits throughout the lifespan. High levels of extrinsic mortality (e.g., from predators) have been shown to sometimes, but not always, select for accelerated actuarial senescence in nature and in the lab. Here, we explore the inductive (i.e., plastic) effects of predation risk (i.e., nonlethal exposure to chemical cues from predators) on the reproductive success of freshwater snails (Physa acuta). Snails were reared either in the presence or absence of chemical cues from predatory crayfish and mated early in life or late in life (a 2 × 2 factorial design); we measured egg hatching and early post‐hatching survival of their offspring. Both age and predation risk reduced reproductive success, illustrating that predation risk can have a cross‐generational effect on the early survival of juveniles. Further, the decline in reproductive success was over three times faster under predation risk compared to the no‐predator treatment, an effect that stemmed from a disproportionate, negative effect of predation risk on the post‐hatching survival instead of hatching rate. We discuss our results in terms of a hypothesized consequence of elevated stress hormone levels.     

Influence of rearing and experimental temperatures on predator avoidance behaviour in a freshwater pulmonate snail

1. Predicted increases in the temperature of freshwaters is likely to affect how prey species respond to predators. We investigated how the predator avoidance behaviour of the freshwater pulmonate snail Lymnaea stagnalis is influenced by the temperature at which it was reared and that at which behavioural trials were carried out. 2. Crawl‐out behaviour of juvenile snails from two populations (high predation risk versus low predation risk) reared at either 15 or 20 °C was assessed in response to predation cues (predatory fish kairomones and conspecific alarm cues) in behavioural trials at both 15 and 20 °C. 3. Trial temperature had a significant effect on the time that snails spent in avoidance, regardless of their population of origin. Crawl‐out behaviour was greater during behavioural trials at 15 °C, but there was no effect of trial temperature on the speed with which animals showed avoidance behaviour. 4. There was no interactive effect of rearing temperature (RT) and trial temperature, but the effect of RT on avoidance behaviour did differ between populations. For an RT of 15 °C, snails from the South Drain (high risk) population showed a more rapid and longer avoidance response than those from the Chilton Moor (low risk) population. In contrast, for snails reared at 20 °C, there was no difference between populations for the duration of the avoidance response and snails from Chilton Moor crawled out faster than those from South Drain. 5. Hence, whilst (predictable) differences relative to natural predation threat in crawl‐out behaviour were apparent at 15 °C, raising the developmental temperature to 20 °C eliminated or, in the case of latency, reversed these differences. This suggests that L. stagnalis populations that cohabit with predatory fish and experience high developmental temperatures may have a reduced ability to respond to fish predation risk.     

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine,the tree swallow Thachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallow Tachycineta bicolor using 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.     

Trade-off between mating and predation risk in the marine snail, Littorina plena

Abstract. We investigated the hypothesis that predation risk affects mating decisions in the intertidal snail Littorina plena in Bamfield Inlet, Northeast Pacific. First, we conducted a field tethering experiment to test the assumption that mating pairs of snails are more susceptible to predation than solitary individuals, and then performed a laboratory experiment to quantify the effect of predation threat on the propensity of snails to form mating pairs. Our results support the hypothesis, in that "mating pairs" were more frequently killed than single snails in the field, and snails were less likely to form mating pairs in the laboratory when simulated predation risk was high (chemical cues from crushed conspecifics were added to the water) than when it was low (no risk cues were added to the water). In contrast to several earlier studies, we found no effect of individual size on snail susceptibility to predation, perhaps because our two size classes were contiguous and snails within them were not dissimilar enough. The results of the behavioral experiment were consistent with this lack of individual size effect on snail vulnerability; both size classes of snails showed a significant and similar tendency to decrease mating when predation risk was high. Taken together, the results of this and recent studies indicate that predators can considerably affect the behavior of littorinid snails, including their movement patterns, feeding, and reproduction. We argue that greater consideration should be given to how marine invertebrates trade off predation risk and activities related to reproduction.     

Mate preference for novel partners in the cricket Gryllus bimaculatus

Abstract .1. To benefit from the putative genetic advantages of multiple mating with multiple partners, female insects would be expected to select against mating with the same male twice when another potential partner is present.
2. This paper examined whether female Gryllus bimaculatus (Gryllidae) preferred to mate with a novel partner over a partner with which they had mated previously.
3. Females presented with a choice preferred significantly to mate with novel males over previously mated males, and preferred to do so even when the potentially confounding influence of male–male competition was controlled for.
4. The potential advantages of such a mate choice pattern and possible ways in which the choice is mediated are discussed.     

Effect of mating on survival at low temperature in females of the Japanese common grass yellow,Eurema mandarina

1. In some insects that overwinter as adults, mating occurs both before and after overwintering. Two hypotheses have been proposed to explain the adaptive significance of pre‐overwintering copulation of females. One is the bet‐hedging hypothesis, which explains pre‐overwintering copulation as a preparation for less chance of mating in the following spring. The other is the nuptial gift hypothesis, which states that secretions derived from males increase overwintering success of females. 2. In Eurema mandarina, both diapause autumn‐ and non‐diapause summer‐form male adults emerge with autumn‐form female adults in the last generation in a year. Most autumn‐form females mate with summer‐form males before winter, and re‐mate with autumn‐form males in the following spring. Because autumn‐form females have sufficient chances for mating after overwintering, the nuptial gift hypothesis has been regarded as the more probable hypothesis. 3. To test the nuptial gift hypothesis, the survival period was compared under short‐day conditions at 10 °C between mated and unmated females that had been reared on sucrose solution at 25 °C for 15–21 days. The mated females had significantly greater longevity than the unmated females, supporting the nuptial gift hypothesis. Body size also affected the survival period. 4. The results suggest that the nuptial gift is an important factor for the evolution of pre‐overwintering copulation in species in which females mate both before and after overwintering.     

Repeated mating with the same male increases female longevity and fecundity in a polyandrous leaf beetle Galerucella birmanica (Coleoptera: Chrysomelidae)

Female multiple mating (polyandry) is widespread across Insecta, even if mating can be costly to females. To explain the evolution and maintenance of polyandry, several hypotheses, mainly focusing on the material (direct) and/or the genetic (indirect) benefits, have been proposed and empirically tested in many species. Considering only the direct benefits, repeatedly‐mated females are expected to exhibit the same fitness as multiply‐mated females under the same mating frequency. In the present study, we compare the fitness of females received monandrous repeated mating (MM) and polyandrous multiple mating (PM) in a polyandrous leaf beetle Galerucella birmanica and assess female mate preference with regard to polyandry or monandry. Our data indicate that the longevity and the egg‐laying duration of MM females are significantly longer than that of PM females. MM females produce significantly more hatched eggs than PM females over their lifetime under the same mating frequency, which results from the high hatching rate of eggs produced by MM females. PM females mated with novel virgin males in the second mating suffer decreased longevity and lifetime fecundity compared with PM females mated with novel mated males in the second mating. Once‐mated females are more likely to re‐mate with familiar males than novel males. By contrast to expectations, the results of the present study suggest that repeated mating provides females with more direct benefits than multiple mating in G. birmanica, and females prefer to re‐mate with familiar males. The possible causes of this finding are discussed.     

The evolutionary dynamics of adaptive virginity,sex‐allocation,and altruistic helping in haplodiploid animals

In haplodiploids, females can produce sons from unfertilized eggs without mating. However, virgin reproduction is usually considered to be a result of a failure to mate, rather than an adaptation. Here, we build an analytical model for evolution of virgin reproduction, sex‐allocation, and altruistic female helping in haplodiploid taxa. We show that when mating is costly (e.g., when mating increases predation risk), virginity can evolve as an adaptive female reproductive strategy. Furthermore, adaptive virginity results in strongly divergent sex‐ratios in mated and virgin queen nests (“split sex ratios”), which promotes the evolution of altruistic helping by daughters in mated queen nests. However, when helpers evolve to be efficient and increase nest production significantly, virgin reproduction is selected against. Our results suggest that adaptive virginity could have been an important stepping stone on the pathway to eusociality in haplodiploids. We further show that virginity can be an adaptive reproductive strategy also in primitively social haplodiploids if workers bias the sex ratio toward females. By remaining virgin, queens are free to produce sons, the more valuable sex in a female‐biased population. Our work brings a new dimension to the studies linking reproductive strategies with social evolution.     

Size‐dependent mating pattern in a nuptial gift‐giving insect

The reproductive interests of females and males often diverge in terms of the number of mating partners, an individual’s phenotype, origin, genes, and parental investment. This conflict may lead to a variety of sex‐specific adaptations and also affect mate choice in both sexes. We conducted an experiment with the bush‐cricket Pholidoptera griseoaptera (Orthoptera, Tettigoniidae), a species in which females receive direct nutritional benefits during mating. Mated individuals could be assigned due to the genotype of male spermatodoses, which are stored in the female’s spermatheca. After 3 weeks of possible copulations in established mating groups which were random replications with four females and males we did not find consistent assortative mating preference regarding to body size of mates. However, our results showed that the frequency of within‐pair copulations (192 analyzed mating events in 128 possible pairwise combinations) was positively associated with the body size of both mated individuals with significant interaction between sexes (having one mate very large, association between body size and the number of copulations has weaken). Larger individuals also showed a higher degree of polygamy. This suggests that body size of this nuptial gift‐giving insect species is an important sexual trait according to which both sexes choose their optimal mating partner.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

The structure of the bursa copulatrix in virgin and mated snails, Helisoma duryi (Mollusca): role of acid phosphatase in reproduction

Abstract. The fine structure of the bursa copulatrix of the virgin snails has been compared with that of mated snails. One of the noticeable changes after mating is an increase in the number of the Golgi and the secretory vesicles. Since some of the vesicles react positively for acid phosphatase it is suggested that this enzyme activity increases following mating. The bursa lumen of the virgin snail contains gel-like materials devoid of spermatozoa, however, following mating, the lumen is full of semen containing live spermatozoa and bacteria. The source of bacteria in the lumen is not known. Acid phosphatase activity is significantly higher in the luminal content of mated snails than in the virgin snails. The activity is higher in the lumen than in the epithelial cells, suggesting that the enzyme is secreted into the lumen where it is utilized for extracellular degradation of spermatozoa. Following mating, the spermatozoa are motile in the lumen of the bursa for ∼3–7 d, but become immobile and finally undergo extracellular digestion so that intact spermatozoa are not recognizable by day 10. The use of castrated snails in mating experiments suggest that individuals of Helisoma duryi reproduce by cross fertilization and that the bursa may act as the holding organ from where the spermatozoa are periodically transported to the carrefour over ∼7 d. At day 10 following mating, however, autosperms appear in the hermaphroditic duct awaiting the next mating.     

Female Gryllus vocalis Field Crickets Gain Diminishing Returns from Increasing Numbers of Matings

Although female insects generally gain reproductive benefits from mating frequently, females do not mate unlimited numbers of times. This study asks whether the limit on female mating rate is imposed by trade‐offs between reproduction and survival. Female Gryllus vocalis were given the opportunity to mate 5, 10, or 15 times with novel males, and the effects on daily fecundity (egg production), fertility (proportion of eggs that were fertilized), and female post‐experimental longevity were measured. Females that mated 10 times laid more eggs and had a higher proportion of fertile eggs than females that mated 5 times. However, females that mated 15 times did not lay significantly more eggs or have a higher proportion of fertile eggs than females that mated 10 times. Although number of matings did not affect the date that females laid their last egg, mating more times was associated with a prolonged period of laying fertile eggs. Number of matings did not affect female post‐experimental longevity. Thus, there was no trade‐off between female reproductive effort and survival, even when females mated very large numbers of times. When females were allowed to mate ad libitum, the average number of times that females mated was greater than the number of times that confers maximal fitness. The lack of cost to mating explains why females might be willing to mate beyond the point of diminishing reproductive returns.     

Old‐male paternity advantage is a function of accumulating sperm and last‐male precedence in a butterfly

Old‐male mating advantage has been convincingly demonstrated in Bicyclus anynana butterflies. This intriguing pattern may be explained by two alternative hypotheses: (i) an increased aggressiveness and persistence of older males during courtship, being caused by the older males' low residual reproductive value; and (ii) an active preference of females towards older males what reflects a good genes hypothesis. Against this background, we here investigate postcopulatory sexual selection by double‐mating Bicyclus anynana females to older and younger males, thus allowing for sperm competition and cryptic mate choice, and by genotyping the resulting offspring. Virgin females were mated with a younger virgin (2–3 days old) and afterwards an older virgin male (12–13 days old) or vice versa. Older males had a higher paternity success than younger ones, but only when being the second (=last) mating partner, while paternity success was equal among older and younger males when older males were the first mating partner. Older males produced larger spermatophores with much higher numbers of fertile sperm than younger males. Thus, we found no evidence for cryptic female mate choice. Rather, the findings reported here seem to result from a combination of last‐male precedence and the number of sperm transferred upon mating, both increasing paternity success.     

Ecology of female mating failure/lifelong virginity: a review of causal mechanisms in insects and arachnids

Sexual reproduction implies binary outcomes of competitive interactions for access to male gametes: lifelong virgin females with null fitness vs. mated females with variable (generally nonzero) fitness. Female mating failure has long remained a dormant concept in sexual selection theory in part because it is acutely maladaptive (lifelong virgins that do not reproduce are strongly selected against) and also due to widespread acceptance of the Bateman–Trivers paradigm (anisogamy and correlated sex roles). Based on recent scientific output on lifelong virginity across multiple taxonomic groups in insects (Coleoptera, Diptera, Hemiptera, Lepidoptera, Odonata, Orthoptera, Strepsiptera), female mating failure has become a mainstay of sexual selection over the last decade. Lifelong virginity and senescence (death) are intertwined processes; old virgin females compensate for increased risk of lifelong virginity by becoming less choosy and increasing investment in mating‐related activities. Low rates of female lifelong virginity (<5%) in most natural populations of insects indicate that sex generally ‘works’ due to selective pressures acting on both males and females to enhance lifetime fitness. Mating failures are most common in insects with female flightlessness; these pressures may lead in evolutionary time to transitionary pathways from sexual reproduction to parthenogenesis. Female mating probability is affected by nonlinear density‐dependent processes dependent upon the scale of observation (mate‐encounter Allee effect at large spatial scales, mating interferences between females at small scales). Mate choice and sex role reversal (females being the active sexual partner) are ubiquitous in insects and arachnids with significant paternal investment, but consequences in terms of female lifelong virginity remain unknown. Logistically, conceptual development of female mating failure in insects is most limited by the lack of broadly applicable methods to assess rates of lifetime virginity among flighted females.     

Polyandry produces sexy sons at the cost of daughters in red flour beetles

Female mating with multiple males within a single fertile period is a common phenomenon in the animal kingdom. Female insects are particularly promiscuous. It is not clear why females mate with multiple partners despite several potential costs, such as expenditure of time and energy, reduced lifespan, risk of predation and contracting sexually transmitted diseases. Female red flour beetles (Tribolium castaneum) obtain sufficient sperm from a single insemination to retain fertility for several months. Nonetheless they copulate repeatedly within minutes with different males despite no direct fitness benefits from this behaviour. One hypothesis is that females mate with multiple partners to provide indirect benefits via enhanced offspring fitness. To test this hypothesis, we compared the relative fitness of F(1) offspring from females mated with single males and multiple males (2, 4, 8, or 16 partners), under the condition of relatively high intraspecific competition. We found that a female mating with 16 males enhanced the relative fitness of F(1) males (in two out of three trials) but reduced F(1) females' fitness (in two independent trials) in comparison with singly mated females. We also determined whether several important fitness correlates were affected by polyandry. We found that F(1) males from mothers with 16 partners inseminated more females than F(1) males from mothers with a single partner. The viability of the eggs sired or produced by F(1) males and females from highly polyandrous mothers was also increased under conditions of low intra-specific competition. Thus, the effects of polyandry on F(1) offspring fitness depend on environmental conditions. Our results demonstrated a fitness trade-off between male and female offspring from polyandrous mothers in a competitive environment. The mechanisms and biological significance of this unique phenomenon are discussed.