Sex expression of an immobile coral-inhabiting snail,<Emphasis Type="Italic"> Quoyula monodonta</Emphasis>

To test the hypotheses explaining the sex expression of the immobile snail Quoyula monodonta, which inhabits the surfaces of the branching coral, Pocillopora eydouxi, the size, sex, gonad development, penis length and the composition of neighboring individuals were investigated between November 1994 and August 1995 in southern Taiwan. Although the snails often aggregated and formed patches, more than 50% were solitary. Females were larger than males both within a snail patch and in the whole population, but the overlap in size range was wide. The males were generally accompanied by females, whereas most females were solitary. Females were rarely (6%) found in the same patch with another female, but 35% of the males had male neighbors. Most juveniles found were also solitary. The composition pattern within a patch cannot be explained by random sampling. Gonad development of an individual was dependent on the presence and the sex of its neighbors within the same patch; the penis length of males also depended on the presence of neighbors. These phenomena suggest that an individual is sensitive to its neighbors. No individuals in the process of sex change were ever found from histological studies of the gonads. Neither the hypothesis that the sex of recruits determines their habitats, nor the hypothesis that there is strict protandric sex-change is supported. The results, however, are all clearly compatible with the hypothesis that the snail has labile sex expression. In the presence of existing females in a patch, recruits are more likely to develop into males, whereas recruits starting a new patch grow to a larger size before developing into females directly. The labile sex expression of Q. monodonta is the only such report in neogastropods.     

Host effect on size structure and timing of sex change in the coral-inhabiting snail<Emphasis Type="Italic"> Coralliophila violacea</Emphasis>

The distribution, size and reproductive characteristics of the snail Coralliophila violacea (Lamarck), which inhabits the surface of both the branching coral Porites nigrescens and the massive corals P. lobata and P. lutea, were surveyed to examine the host effect on: (1) population structure and (2) reproductive characteristics, including the size at sex change of symbionts. On branching hosts, most snails were solitary, whereas on massive hosts, most had formed multiple-snail patches. Significantly smaller snails as well as proportionally more females were found on branching than on massive hosts. Furthermore, the fecundity of the females on the branching hosts was significantly lower than that on the massive hosts. The size at sex change (male to female) of the snails was smaller on the branching hosts than on massive hosts. Patch composition differences can partly explain the smaller size at sex change for snails on branching hosts; however, there was also evidence that host morphology had a significant effect on the timing of sex change.Communicated by T. Ikeda, Hakodate     

Do sex-changing male snails use mate choice to get a jump on their “size advantage”?

Individuals of species that change sex from male to female may gain a “size advantage” from that sex change; that is, as males become larger, they become female, thus increasing their fecundity with their size. However, males could also gain an early and different reproductive size advantage by choosing large females as mates. While male preference for large females has been observed in many dioecious species, we know little about male size preference in sex-changing species. In choice experiments, we examined whether males of two congeneric species of marine sex-changing snails, Crepidula fornicata and C. convexa, chose large females over small ones as partners. We also used choice tests to see whether males of C. fornicata, a species whose members form long-term, multi-animal stacks, would choose two females in a stack over a single female. Surprisingly, males of neither species showed a preference for large females, in spite of the documented fecundity advantage associated with large female-size. Males of C. fornicata chose slightly, but not significantly, more single females than stacks, suggesting that neither number nor size drives mate choice in these animals. Key factors that may influence this lack of size preference include long association time, the likelihood of sperm competition, and the cost of extended mate search; it may also be that sex-change itself, the very factor that creates female-biased sexual size dimorphism in these species, prevents size preference, as males may gain sufficient reproductive advantage from eventually becoming large females themselves to offset any benefit of choosing large females.     

Variation in size at sex-change among natural populations of the protandrous hermaphrodite, <Emphasis Type="Italic">Crepidula fornicata</Emphasis> (Gastropoda,Calyptraeidae)

Protandrous hermaphrodites are predicted to change sex from male to female when relative reproductive fitness of females surpasses that of males. How size at sex transition varies with population, mating group and individual parameters was investigated for five populations of the protandrous hermaphrodite slipper snail, Crepidula fornicata. The populations varied for density, size distribution, average mating group size and sex ratio. Size at sex-change was correlated with the population sex ratio. Comparisons of multiple hypotheses revealed that variables predicting the sex of a snail vary among positions in the mating group. The variables included body size, the relative size of the snail sitting atop the focal snail and population density. Our data support the conclusions that size at sex-change (and by inference, the size at which one sex has relatively greater fitness) is not fixed for these hermaphrodites and that individual size, social conditions and population differences all influence variation in relative fitness.     

Experimental studies on water strider mating dynamics: spatial variation in density and sex ratio

Summary We used field surveys, field experiments and experiments in artificial pools to study the effects of variation in sex ratio and density on mating dynamics of a stream water strider, Aquarius remigis. Our field survey documented the existence of hot spots, sites of higher than average total gerrid density, a male-biased sex ratio, and higher than average female mating activity. Female gerrids frequently changed sites, usually moving upstream, perhaps to spread their eggs among many sites. Male gerrids showed two movement strategies: some males frequently changed sites, while other males were stationary at hot spots. Surprisingly, smaller males tended to be stationary at hot spots. A field manipulation of the availability of refuges for females to avoid harassment by males supported the notion that males prefer hot spots because they are sites where a scarcity of refuge for females makes it relatively easy for males to intercept females. Experiments in plastic pools compared sites with 20 males: 5 females (simulating hot spots) to pools with 5 males: 5 females. The rate of male harassment of females was higher in 20:5 pools as compared to 5:5 pools. In response to increased male harassment, females reduced their activity on the water and increased their time spent out of the water and thus unable to forage. Males showed a large male mating advantage (LMMA) in 5:5 pools, but, surprisingly, not in 20: 5 pools. This pattern can explain the field observation that small males prefer hot spots. A behavioral mechanism that can explain the LMMA is as follows. Mating occurs when males overcome female resistance. Larger males have a mating advantage over smaller males if females resist heavily. Increased harassment (e.g., in 20:5 pools as compared to 5:5 pools) might result in reduced female resistance to males and thus a reduced LMMA. Females also showed some non-random mating by size that might reflect an interplay between male preference for large females and the avoidance of males by large females. Correspondence to: A. Sih     

Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. I: Population structure of snails and corals

Despite their potential importance in structuring reef communities, invertebrate corallivores and their population structures are poorly understood. We found distinct differences in the population structures (length-frequency distribution and sex ratio) of the corallivorous gastropod Coralliophila abbreviata residing on two coral-host taxa, Montastraea spp. and Acropora palmata, in the Florida Keys. In each of two survey years, around 50% of the Montastraea spp. colonies were infested, with a mean snail density of eight snails per infested colony (range 1–45), while around 20% of A. palmata colonies harbored three snails per infested colony (range 1–23). Variation in patterns of snail occurrence was also observed within a host taxon. A. palmata occurred in low- and high-density stands (0.4 and 1 colony m^–2, respectively, at the initial survey) at different sites. Hurricane Georges struck the area in September 1998. When resurveyed in 1999, density of colonies in low-density stands had decreased by 75% to 0.1 colonies m^–2. This decrease was accompanied by a doubling in the proportion of colonies infested with snails (from 19% to 46%) and an increase in snail density per infested colony (from 3.7±3.3 SD to 5.4±4.6 SD) as snails apparently concentrated on surviving A. palmata. In contrast, sites with high density A. palmata stands (thickets) retained colony densities of about ~1 colony m^–2 among years, while snail infestation increased only from 9% to 14% of colonies surveyed and snail density essentially remained unchanged (from 2.7±1.8 to 2.9±1.9 snails per infested colony). Snails collected from Montastraea spp. were shorter than those from A. palmata in low-density stands and were longest on A. palmata in thickets. On both host taxa, female snails were longer than males. The sex ratio of snails on Montastraea spp. hosts was even (1:1), while that of snails on A. palmata was skewed (70% males). Factors that could explain observed differences in size structure and sex ratio between Coralliophila populations on the two coral host taxa include: differential susceptibility to predators, influence of host tissue nutritional quality and/or secondary metabolite content, and genetic differences (cryptic species). The host-specific characteristics of C. abbreviata populations imply that the impact of gastropods on reef communities will vary with the coral species composition.     

Colony level sex allocation in a polygynous and polydomous ant

The colony-level sex allocation pattern of eusocial Hymenoptera has attracted much attention in recent studies of evolutionary biology. We conducted a theoretical and empirical study on this subject using the dolichoderine ant Technomyrmex albipes. This ant is unusual in having a dispersal polymorphism in both males and females. New colonies are founded by an alate female after mating with one or more alate males in the nuptial flight. In mature colonies, the reproductive role of the foundress queen is taken over by wingless offspring (supplementary reproductives). Mature colonies are extremely polygynous, with many wingless queens reproducing through intea-colonial mating with wingless males (inbreeding), and producing both alate and wingless sexuals. The population sex ratio of wingless sexuals was found to be extremely female-biased, while the population allocation ratio of alates was almost 1:1. This result suggests that there is local mate competition among wingless sexuals. A specific model for this extraordinary life cycle predicted that the asymmetry of regression relatedness (b _f/b _m) will disappear during the first few generations of wingless reproductives after the foundress dies. If colonies begin to produce alates after several wingless generations, this undermines the hypotheses for intercolonial sex ratio variation based on the relatedness asymmetry. We compared the magnitude of variation in sex ratios and other characteristics between two levels (within-colony-inter-nest and between-colony). Although there was considerable within-colony variation in all the examined characteristics, between-colony variances were always larger. This means that allocation is important at the whole-colony level, not that of the nest. There was no apparent correlation between the sex ratio of alates and colony size. Furthermore, partial correlation analysis indicated that neither the number of workers nor investment in alates explained the variation in the sex ratio of alates. The only factor which was significantly correlated with the sex ratio of alates was the sex ratio of wingless sexuals (a positive correlation). We conclude that both the alate and wingless sex ratios may be influenced by a common primary sex ratio at the egg stage, the variance of which may have genetic components. In the wingless sexuals, partial correlation analysis indicated that colony size and the number of workers explained the sex allocation ratio. The number of wingless females was strongly (positively) correlated with the total investment in wingless sexuals, while the number of males showed no such correlation. There is, however, no convincing explanation for the variation in sex allocation ratio of wingless sexuals, because the estimates of investment in wingless males may have a large sampling error. Correspondence to: K. Tsuji     

Selective males and ardent females in pipefishes

Summary In the pipefishes Syngnathus typhle and Nerophis ophidion, males have been shown to limit female reproductive rate, and females to compete for access to males. Hence, these species fit the criteria for sex-role reversal. Males brood the eggs and provide the offspring with nutrients, oxygen and an osmoregulated environment. Moreover, in S. typhle both sexes prefer a larger mate when given a choice. Sexual selection theory predicts that males should be more choosy than females, and that was experimentally demonstrated in this study. We predicted that S. typhle males should be less eager to copulate than S. typhle females with an unattractive (i.e. small) mate. We measured eagerness as the time from the start of the experiment until copulation occurred. Males with unattractive partners took significantly longer to copulate than females with unattractive partners. Moreover, females invariably initiated the courtship dance, and resumed it quicker after copulation than did the males, again suggesting reproductive hesitation in males. Neither male nor female size per se was correlated with time until copulation. In N. ophidion, where we have previously shown that males prefer larger to smaller females, we found that females did not select males with regard to size. Our results are consistent both with earlier findings (males limit female reproduction and females compete for males) and with operational sex ratios in nature: in seven annual field samples in June, the numbers of S. typhle females with ripe eggs always significantly exceeded numbers of receptive males. Hence, the potential cost of being choosy in terms of lost matings is much higher in females than in males. In conclusion, S. typhle females were somewhat choosy, but less so than males, whereas N. ophidion females were not choosy at all. Correspondence to: A. Berglund     

Male-male competition selects for delayed sex change in the protandrous shrimp<Emphasis Type="Italic"> Pandalus latirostris</Emphasis>

In most protandrous species, male size advantage is generally regarded as unimportant in determining the timing of protandrous sex change. In pandalid shrimp, the size/age of male sex change often fluctuates among years and populations, but the adaptive significance of late reversing males (LRMs) is not well understood. This study experimentally examined the adaptive significance of LRMs in the protandrous pandalid shrimp Pandalus latirostris Rathbun. Field and laboratory studies were carried out in August–September of 1998–2002 on P. latirostris in Notoro Lagoon, Japan (44°03′N; 144°10′E). Mature females that had molted (i.e. mate receptive) were tethered in the field and their mating behavior with wild males was observed. Copulations occurred with a single male at a time, although other males could sequentially mate with a tethered female. Because tethered females rejected male approaches, males had difficulty transferring their spermatophores. In the laboratory, males copulated with non-tethered, recently molted females for only 15 min after molting. Recently molted females are wary of potential predators, since their soft exoskeleton makes them particularly vulnerable. Fast access by males enhanced fertilization success in this shrimp. The effect of male size on mating success in the laboratory was examined. Both small and large males successfully inseminated females in the absence of competitors. In experiments where large, medium, and small males competed for a female, however, larger males guarded females longer than smaller males, until the female molted and became receptive. Moreover, large males were more successful at copulating once molting occurred. These results imply that male-male competition drives delayed sex change in some situations.     

Population characteristics of the deep-sea lobsters Polycheles typhlops and Stereomastis sculpta (Decapoda: Polychelidae) in a bathyal mud community of the Mediterranean Sea

Distribution and abundance as a function of depth, and population parameters such as sex and size structure of the population and aspects of reproductive biology have been studied in the deep-sea polychelid lobsters Polycheles typhlops and Stereomastis sculpta. Samples were taken by otter trawl in the Balearic Sea, a deepsea basin in the northwestern Mediterranean Sea from 1985 to 1989. In both species the largest size classes were dominated by females. In S. sculpta, male and female sizes were very similar. In P. typhlops, ovigerous females and those with external spermatophores were 23 and 25 mm in carapace length (CL), respectively; males with external spermatophores were 17 mm in CL. In S. sculpta, ovigerous females and those with external spermatophores were 24 and 19 mm CL, respectively, and males with external spermatophores 19 mm CL. Highest densities of P. typhlops occurred along the middle slope at depths between 500 and 1000 m. Only small-sized individuals were found at the deepest depths sampled; some recruitment must therefore occur in waters much deeper than those usually inhabited by the adult population. The sex-ratio was 1:1 in most samples, but in some of the shallowest samples females predominated. The depth distribution range of S. sculpta was 981 to 2253 m: densities clearly increased with increasing depth. There were no apparent variations in size distribution as a function of depth. Since very few adult males and females were captured, the population in the survey area would seem to be comprised mainly of juveniles. As a function of depth, females were numerically dominant only in some of the shallowest samples taken in the distribution range of this species. There is strict habitat partitioning between the two species. In both species, the variation in the sex-ratio as a function of depth suggests differential migration between the sexes, probably related to egg incubation and hatching.     

Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae)

Summary Colonies of Anelosimus eximius in Panama had an average sex ratio of 0.15±sd 0.09, i.e. about five females for each male. The sex ratio in egg sacs reared was even lower (0.08±0.01), as was that of immatures in newly founded colonies (0.12±0.05). The possible mechanisms responsible are discussed. Mature colonies had an average ratio of 17 females and 2 males for each egg sac present (range: 2–91 females, 0.2–8.2 males) and contained a large proportion of females which were not inseminated but which presumably help. Since both sexes are diploid, arrhenotoky can be ruled out and it is assumed that some females do not come to reproduction, the proportion depending on the availability of resources. This mechanism may enable entire colonies to survive lean times.     

Dispersal: an alternative mating tactic conditional on sex ratio and body size

Summary Small male milkweed beetles are less successful at obtaining mates than are larger males. Larger males usually win fights and prevent smaller males from obtaining mates and from choosing larger more fecund females as mates. When sex ratios are male-biased, smaller males are particularly likely to experience these mating disadvantages. It follows that smaller males should be especially responsive to their local competitive environment and behave so as to minimize the mating disadvantages of their smaller size. This paper tests the hypothesis that smaller males disperse from host plant patches with male-biased sex ratios and remain in patches with female-biased sex ratios more readily than larger males.Results show both larger and smaller males disperse from patches with male-biased sex ratios more frequently than from patches with femalebiased sex ratios. As predicted, however, small males are more likely to disperse from patches with male-biased sex ratios and remain in patches with female-biased sex ratios than are larger males.The data also show that smaller males dispersing from patches with male-biased sex ratios obtain more matings than non-dispersing males.For milkweed beetles, moving between patches can be viewed as an alternative mating tactic conditional on male body size and local sex ratio.     

Trail following,speed and fractal dimension of movement in a marine prosobranch,Littorina littorea,during a mating and a non-mating season

We quantitatively studied movement behaviour in the gastropod Littorina littorea in laboratory experiments during periods of their non-mating season (November 1992) and their mating season (April–May 1993). Snails were collected in 1992 and 1993 from one boulder shore on the north west coast of Sweden. In a comparison between the two seasons (one non-mating and one mating) we measured trail complexity of unsexed snails using fractal dimesion, the degree of mucus trail following (coincidence index of marker and tracker trails) and average movement speed of marker and tracker snails. We found no differences in fractal dimension and coincidence index of trails between the two seasons. Tracker snails moved, however, significantly faster than marker snails during both seasons. This could not be explained by trackers, when locomoting, using the mucus trail deposited by the marker to increase their speed, since there was no correlation between coincidence index and tracker speed. During the mating season we also conducted trail complexity, trail following and speed experiments comparing the behaviour of males and females. There was no difference between males and females in the fractal dimension of movement, nor was there any difference between the mean speed of male and female snails, although male marker snails tended to move faster than female marker snails. Males tracking other males, females tracking other females and females tracking males followed trails about equally long distances (i.e., coincidence indices did not differ). In contrast, males following female mucus trails showed a significantly higher degree of trail following than the other sex combinations. This new finding may suggest that females of L. littorea release pheromones in their mucus trails and that males are able to identify them.     

Estimation of age in the sex-changing, coral-inhabiting snail Coralliophila violacea from the growth striae on opercula and a mark–recapture experiment

Change of sex in the coral-inhabiting snail Coralliophila violacea (Lamarck) may occur in a wide range of sizes in the field. One proposed explanation for this is that the snails change their sex at a certain age and that individuals have different growth rates caused by microhabitat differences. In this study, we attempt to establish a method to determine the age of this snail and age at sex change. The growth striae on the operculum were studied and compared to the age estimated by the Gompertz growth function based on growth data obtained from mark-recapture experiments in southern Taiwan. There is a significant correlation between the number of striae on the operculum and the age estimated from the Gompertz growth function, and the relationship is 1:1. These results suggest that the number of striae on the operculum can be used as an age index, with each stria representing 1 year of age. The age of sex change of this snail, according to our estimates by both stria number and aperture-length inferences, occurs between 4 and 6 years old. Growth rates of the snails are negatively correlated to size. Furthermore, individuals undergoing sex-change grow faster than males and females.     

Correlates of male mating success in two species of Anthidium bees (Hymenoptera : Megachilidae)

Summary Field observations were made on the mating behavior of two congeneric species of solitary bees, Anthidium porterae in an arid grassland and A. palliventre in a coastal sand dune habitat. Males of both species exhibited resource defence polygyny and defended hostplants to gain access to females foraging for nectar and pollen. The mating frequencies of marked and measured resident (territorial) males were monitored during periods of continuous observation, following which measurements of territory size and floral resources were obtained. Mating success of A. palliventre males was strongly influenced by territory characteristics: Males that defended small areas with a few rich hostplant patches mated more often than males that held larger territories containing many hostplant patches of low floral density. Large males generally held high-quality sites and thus had a mating advantage over smaller individuals. In A. porterae, on the other hand, male mating success was unrelated to any measure of territory quality. Copulation frequency and male size were positively correlated, however, apparently due to the increased ability of large males to seize and hold females for mating. The two species also differed in the incidence of non-territorial, sneaky males. While absent in A. palliventre, sneaky males accounted for 12% of all mating observed in A. porterae. Males of A. porterae that displayed sneaky tactics mated, on average, as often as resident males. Offprint requests to: E.M. Villalobos     

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

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

Sex ratios in bumble bee colonies: Complications due to orphaning?

Summary Highly male-biased investment ratios (1:2.86, F:M, biomass wet weight) were found among reproductives reared by 32 Bombus terricola colonies set out in the field and 35 B. melanopygus colonies held in laboratory confinement. The numerical sex ratios were almost identical for the two species (1 queen to 6.1 males). Most of the males were produced early in the season by small weak colonies. It is argued that in a population where some colonies are orphaned and rear only males, female-biased sex ratios are expected among the surviving queen-producing colonies.     

Bi-directional sex change in a coral-dwelling goby

Bi-directional sex change has recently been reported among obligate coral-dwelling gobies of the genus Gobiodon. However, neither the functional role of this pattern of sex change nor the frequency of sex change in either direction in natural populations is known. We investigated the social structure and pattern of sex change of Gobiodon histrio at Lizard Island on the Great Barrier Reef. The social structure of G. histrio within coral colonies usually consisted of a single juvenile or a heterosexual adult pair. The size of adult social groups was not constrained by coral colony size. In contrast to expectations for pair-forming species, G.␣histrio was primarily a protogynous hermaphrodite. All immature G. histrio were females and sex change from female to male occurred readily when two mature females were placed in a coral colony. In addition, male G. histrio were able to change back to females when two mature males were placed in a coral. Sex change from female to male, however, occurred with over twice the frequency of sex change from male to female. Where two males were placed in a coral colony, heterosexual pairs were most frequently re-established by immigration of females from outside the treatment population. This pattern might be predicted if sex change from male to female is more expensive than sex change from female to male for G. histrio. Where sex change is expensive, movement may be favoured over sex change, particularly where coral densities are high and movement among corals incurs little mortality risk. Received: 10 November 1997 / Accepted after revision: 16 May 1998     

Male mating history and female fecundity in the Lepidoptera: do male virgins make better partners?

In insects, large ejaculate and associated materials, including spermatophores, appear to have evolved via sexual selection acting on males to either delay female remating or to increase the rate of egg-laying. It is also possible, however, that females use nutrients transferred during mating to increase their lifetime fecundity. If so, male ejaculate size may also have evolved under natural selection as a form of paternal investment. In Lepidoptera, males with a greater number of prior matings tend to produce smaller spermatophores. However, the reported effects of male mating history on female fecundity vary widely among species. We therefore performed a meta-analysis using data from 29 studies of 25 species. Overall, the reproductive output of females mated to virgin males was significantly higher than that of females mated to sexually experienced males (Hedges d=0.33, P<0.01). A sample size of around 145 females per male mating type is required to detect an effect of this size with 80% statistical power at =0.05 (two-tailed). There was no difference in mean effect size between butterflies/skippers and moths. After controlling for any effect of taxonomic group, however, the mean effect size for polyandrous species was significantly greater than that for monandrous species (Hedges d=0.45 vs 0.25, P=0.01). We then discuss possible reasons why male mating history, presumably acting through its effect on spermatophore size, might have a stronger effect in polyandrous than monandrous species.Communicated by A. Cockburn     

Male copulatory guarding enhances female foraging in a water strider

Summary In the laboratory, females of Gerris remigis foraging singly after being separated from a copulating male averaged 16 times longer to repel male copulatory attempts compared to repelling time when copulating (60 trials). In<10 min after separation, all females either became immobile at the edge (in 34% of all trials), or recopulated and either continued to forage (61%) or swam to the edge and became immobile (5%). Copulating females captured 85% of imitation prey, but single females, harassed by males, captured only 32% of prey.During a census of 2 small streams, in the pools with swimming single males, >99% of all swimming females were copulating, <1% being single. Ninety seven percent of all single females were immobile at the edge, whereas only 11% of all pairs were at the edge. In pools without swimming single males, 32% of all single females were swimming, compared to <1% in pools with swimming males. Introductions of a male into 4 pools with a female swimming singly in each resulted in the females either becoming immobile at the edge or copulating. The 2 streams had a : sex ratio of 1.6:1, and 79% of all females were copulating during the census. Foraging, copulation and copulatory attempts continued at a reduced level during the night.Thus in streams where male G. remigis are attempting to copulate, females can forage effectively only by carrying a copulating male who apparently repels copulatory attempts by other males.