Offspring sex ratio response to host size in the parasitoid wasp Spalangia endius

The host size model, an adaptive model for maternal manipulation of offspring sex ratio, was examined for the parasitoid wasp Spalangia endius. In a Florida strain, as the model predicts, daughters emerged from larger hosts than sons, but only when mothers received both small and large hosts simultaneously. The pattern appeared to result from the mother's ovipositional choice and not from differential mortality of the sexes during development. If sex ratio manipulation is adaptive in the Florida strain, it appears to be through a benefit to daughters of developing on large hosts rather than through a benefit to sons of developing on small hosts. Both female and male parasitoids were larger when they developed on larger hosts. For females, developing on a larger host (1) increased offspring production, except for the largest hosts, (2) increased longevity, (3) lengthened development, and (4) had no effect on wing loading. For males, development on a larger host had no effect on any measure of male fitness – mating success, longevity, development duration, or wing loading. In contrast, a strain from India showed no difference in the size of hosts from which daughters versus sons emerged, although both female and male parasitoids were larger when they developed on larger hosts. These results together with previous studies of Spalangia reveal no consistent connection between host-size-dependent sex ratio and host-size-dependent parasitoid size among strains of S. endius or among species of Spalangia. Received: 28 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

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

Sex ratio and oviposition responses to host age and the fitness consequences to mother and offspring in the parasitoid wasp Spalangia endius

In the parasitoid wasp Spalangia endius more offspring and a greater proportion of daughters were oviposited in, and emerged from 0-day-old versus 3-day-old hosts. Offspring that developed on the younger hosts (1) were larger at adulthood, (2) developed more quickly, (3) had higher survivorship to adulthood, and (4) were more often able to chew their way out of the host. Sons and daughters did not differ in how host age affected their size, development rate, or survivorship. The greater proportion of daughters from the younger hosts may be adaptive, as described by the host quality model (a variant of the Trivers and Willard hypothesis). It is adaptive if greater size or more rapid development has a more positive effect on a daughter’s than a son’s fitness and the positive effect is large enough to compensate for sons being trapped disproportionately to daughters in the older hosts. Despite greater success at drilling the younger hosts, mothers did not try to drill them sooner or more often. Having previously oviposited on the older rather than the younger hosts had no detrimental effect on the mother’s subsequent longevity or offspring production. Received: 8 March 2000 / Revised: 9 June 2000 / Accepted: 24 June 2000     

Sex ratio, sexual dimorphism and mating structure in bethylid wasps

Sexual dimorphism has been linked to parasitoid mating structure by several authors. In turn mating structure has an important influence on predicted sex ratio optima. Here we test the relationship between sexual dimorphism and sex ratio using data from 19 species of bethylid wasps. Using phylogenetically based comparative methods we confirm the findings of a previous cross-species analysis that sex ratio (proportion of males) is strongly and negatively correlated with clutch size. Using cross-species comparisons we show an additional positive correlation of sex ratio and relative male size, as predicted. The relationship however is not significant when using phylogenetically based methods. The cross-species result is largely due to differences between two bethylid sub-families: the Epyrinae have relatively large males and relatively high sex ratios, whereas the Bethylinae have relatively small males and lower sex ratios. Our study illustrates the benefits and drawbacks of using cross-species versus phylogenetically based comparisons. Received: 13 May 1997 / Accepted after revision: 12 January 1998     

Contradictory results in sex ratio studies: populations do not necessarily differ

In avian sex ratio studies, results often differ between species and between populations within species. Some researchers argued that positive results were simply statistical artefacts and that sex ratio adjustment did not exist. However, many of the proposed mechanisms of sex ratio adjustment result in costly laying gaps. In these cases, females laying large clutches may restrict the sex manipulation to the first egg of the clutch. Consequently, detectability of sex ratio adjustment on the level of the clutch can be low. Though obvious, this fact is often neglected in the literature. Using simulations, I show that the proportion of undetected sex manipulation can be surprisingly high when the manipulation is restricted to the first egg. If the sample size is 50 broods, there is 47% and 71% chance in 6- and 12-egg clutches, respectively, that sex manipulation is undetected. Even with large samples (n = 100), the figures are 15% and 46%. These data suggest that nonsignificant results for clutch sex ratios do not necessarily mean that sex is not manipulated in a portion of the brood, e.g. in first-laid eggs. Hence, whenever possible, data on laying order-specific sex manipulation should also be collected. Without such data, contradictory results on brood sex ratios should be interpreted cautiously.     

Relatedness and sex ratio in a primitively eusocial halictine bee

Lasioglossum laevissimum was studied in Calgary, Alberta, where it is eusocial with one worker brood. Estimates of relatedness were obtained among various categories of nestmate based upon four polymorphic enzyme loci, two of which exhibited significant levels of linkage disequilibrium. Relatedness estimates among workers and among reproductive brood females were very close to the expected 0.75 value that obtains when nests are headed by one, singly mated queen. However, relatedness between workers and the reproductive brood females they reared was significantly lower than 0.75. A low frequency of orphaning with subsequent monopolisation of oviposition by one worker brood female in orphaned nests may explain these results. Workers were significantly more and queens significantly less closely related to male reproductives than expected if all males were to have resulted from queen-laid eggs. Orphaning and worker-produced males contribute to this result. The sex investment ratio was 1:2.2 in favour of females, in excellent agreement with the predictions based upon relative relatednesses between workers and reproductive brood males and females. Adaptive intercolony variation in investment ratios was detected: the sex ratio was more heavily female-biased in nests in which the relative relatedness asymmetry between workers and reproductive brood was more female-biased. The study species is the most weakly eusocial hymenopteran for which relatedness estimates and sex ratio data are available. With high relatedness among nestmates and a strongly female-biased sex ratio, this study suggests the importance of indirect fitness contributions in the early stages of social evolution. Correspondence to: L. Packer     

Precise sex allocation, local mate competition, and sex ratio shifts in the parasitoid wasp Trichogramma pretiosum

We determined the sex, order, and clutch size of eggs laid by the parasitoid wasp, Trichogramma pretiosum Riley, in the eggs of one of its natural hosts, Trichoplusia ni (Hübner). The parasitoid allocated sex non-randomly to hosts in the laboratory with a variance significantly less than that of a binomial (random) distribution, our null model. More clutches of two or more eggs contained a single male egg as the second or third egg laid than would be expected by chance and none contained two or more male eggs. T. pretiosum also increased the sex ratio (% male) of its offspring with increasing foundress numbers by increasing the frequency of male offspring as the second egg in a two-egg clutch allocated to unparasitized hosts and as the single egg allocated to previously parasitized hosts. These results indicate that T. pretiosum allocates the sex of its offspring precisely. Precise sex allocation is favored under local mate competition because it reduces variation in the number of sons per patch thus maximizing the number of inseminated daughters emigrating from the patch. Similar combinations of female and male offspring emerged from T. ni eggs parasitized by T. pretiosum in the field, again with a sex ratio variance less than that expected for a binomial distribution. These results strongly suggest that this parasitoid species manifests local mate competition.     

Biased primary sex ratio in the bushcricket Poecilimon veluchianus,an insect with sex chromosomes

Offspring sex ratio at hatching was examined in the bushcricket Poecilimon veluchianus. Offspring sex ratios varied significantly between females (Fig. 1). Low mortality prior to sex determination established that this heterogeneity was already present in the primary offspring sex ratio. Sperm age and female age had no influence on offspring sex ratio (Fig. 2). Male age at copulation, however, correlated significantly with offspring sex ratio (Fig. 3). There were two types of males: one type produced predominantly daughters when young and an increasing proportion of sons with age. The other type produced, independent of age, 1:1 offspring sex ratios (Fig. 4). The two types of males seem to occur in approximately equal numbers. Sex ratio variation (1) may adaptively compensate for local sex ratio biases caused by sex-specific motility, or (2) it may be adaptive if there is a sex-differential effect of laying date on offspring fitness. Received: 14 March 1996/Accepted after revision: 24 June 1996     

Unattractiveness of mated females to males in the parasitoid wasp Spalangia endius

Despite common stereotypes, males are not always indiscriminate and eager when it comes to mating. In the parasitoid wasp Spalangia endius, the initial response of males to females was almost always one of apparent excitement; however, this was followed by a clear preference for virgin females over mated females in both no-choice and choice situations. The no-choice data were collected from videotapes of male-female pairs of all possible combinations of mated and virgin individuals. Neither female nor male mating status had a significant effect on likelihood of, or time until, contact or male courtship fanning. However, a males first retreat was sooner when the female was mated than when she was virgin; mated males exhibited their first retreat sooner than did virgin males; and mated females were less likely to be mounted than were virgin females. In addition to the videotapes, male-choice experiments were performed. When given a choice of a virgin and a mated female, both virgin and mated males were more likely to mount and copulate with the virgin. The difference in response to virgin versus mated females seemed to be less in virgin males than in mated males, perhaps due to virgin males greater eagerness to mate: when a virgin male and a mated male were presented with a dead virgin female, the virgin male was usually the first to respond to the female. That males preferentially retreated from and avoided mounting mated females appears to be adaptive given that mated females rarely copulated.Communicated by N. Wedell     

Maternal investment, sex-differential prospects, and the sex ratio in wild house mice

In a population of first-generation offspring from wild-caught house mice (Mus musculus domesticus), previous evidence suggested that male fitness is more strongly affected by an increase in body weight than female fitness. This paper shows that in these mice the young are weaned at heavier weights the smaller the litter and the better the maternal body condition. These effects persisted into adulthood and were less pronounced in female young. However, contrary to expectation from conventional sex ratio theory, maternal condition and litter size had no detectable effect on sex ratios. Also, litter size did not affect sex ratios in two populations of laboratory-kept, wild-caught western (M. m. domesticus) and eastern house mice (M. m. musculus). Wild house mice, therefore, appear not to adaptively manipulate the sex ratio of offspring. It is argued that this absence of sex ratio trends might not be maladaptive, but rather that models currently used to predict sex ratio trends in rodents may not be valid. Received: 13 March 1997 / Accepted after revision: 9 August 1997     

Asymmetric larval competition in the parasitoid wasp <Emphasis Type="Italic">Nasonia vitripennis</Emphasis>: a role in sex allocation?

Sex allocation theory offers excellent opportunities for testing how animals adjust their behaviour in response to environmental conditions. A major focus has been on instances of local mate competition (LMC), where female-biased broods are produced to maximise mating opportunities for sons. However, the predictions of LMC theory can be altered if there is both local competition for resources during development and an asymmetry between the competitive abilities of the sexes, as has been seen in animals ranging from wasps to birds. In this paper, we test the extent to which asymmetric larval competition alters the predictions of LMC theory in the parasitoid wasp Nasonia vitripennis. We found that the body size of both sexes was negatively correlated with the number of offspring developing within the host. Further, we found that when faced with high levels of competition, the body size of females, but not males, was influenced by the sex ratio of the competing offspring; females were smaller when a higher proportion of the brood was female. This asymmetric competition should favour less biased sex ratios than are predicted by standard LMC theory. We then develop a theoretical model that can be parameterised with our data, allowing us to determine the quantitative consequences of the observed level of asymmetric larval competition for sex allocation. We found that although asymmetric competition selects for less biased sex ratios, this effect is negligible compared to LMC. Furthermore, a similar conclusion is reached when we re-analyse existing data from another parasitoid species where asymmetric larval competition has been observed; Bracon hebetor. Consequently, we suspect that asymmetric larval competition will have its greatest influence on sex ratio evolution in species that have smaller clutches and where local mate competition is not an issue, such as birds and mammals.     

Differential predator escape performance contributes to a latitudinal sex ratio cline in a migratory shorebird

Sexual segregation outside the mating season is common in vertebrates, and has been attributed to sexual differences in predator escape performance in ungulates and fish, but not in birds. Here, we tested the hypothesis that sex- and latitude-specific predator escape performance underlies the differential nonbreeding distribution of western sandpipers (Calidris mauri), a migratory shorebird. Females overwinter further south along the American Pacific coast, creating a latitudinal cline in sex ratio. Escape performance is reduced with increasing body mass, and birds generally carry less fat further south. Western sandpipers with poor escape performance were therefore predicted to prefer southern sites to reduce the risk of mortality posed by predators. Data from four nonbreeding latitudes showed that wing loading, used as an index of escape performance, was overall higher for females, and that it decreased with latitude in both sexes. Within latitudes, wing loading was lower at smaller, and presumably more dangerous, sites. Flight response to a predatory attack was longer in the south. Mortality risk offers a novel and candidate explanation for differential distribution patterns in western sandpipers and possibly other avian migrants.     

The response of an insect parasitoid,Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation

Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.     

Precise sex ratio in a parasitic wasp: the result of counting eggs

Summary The parasitic wasp Nesolynx albiclavus maintains a very skewed offspring sex ratio (ca. 4% males). We studied proximate, causal factors of the son-daughter choice mechanism to obtain insight into the precision of the procedures and the underlying rules. We analyzed the number of males in clutches produced during sessions of 12 or 24 h. The results show that the timing of the next male egg depends on the time elapsed and the number of eggs laid since the previous one. The influence of the passage of time per se during non-laying periods was reduced in an experiment with 4-h periods. The data show that the first male egg is produced by a fairly rigid process: about the sixth egg in a run will be left unfertilized. No further male egg will appear in the next 10–20 eggs if they are laid in rapid succession.     

Local mate competition and precise sex ratios in Telenomus fariai (Hymenoptera: Scelionidae), a parasitoid of triatomine eggs

Telenomus fariai is a gregarious endoparasitoid of the eggs of several species of Triatominae (Hemiptera) with a high degree of sibmating: males fertilize their sisters inside the host egg before emergence or emerge first and copulate with their sisters as these emerge. Our results show that, when laying alone, T. fariai behaves adaptively, minimizing offspring mortality and conforming to the prediction of local mate competition (LMC) theory by laying a single male, which is sufficient to fertilize all the sisters. When more than one wasp was placed with one host, sex ratios still conformed to LMC predictions but, despite the decreasing number of eggs laid per wasp, clutch size could not be completely adjusted to avoid mortality. This is not surprising, as superparasitism is rare in the field. Offspring production was independent of the contacts between conspecifics but was affected by the number of mothers laying on a single host egg. The sex of the progeny was precisely determined: a female produced one male per clutch when laying on both unparasitized or previously parasitized hosts. On the other hand, a mother produced less daughters when superparasitizing. Under crowded conditions, the number of eggs laid per female wasp and per host decreased as the number of mothers increased. Developmental mortality also increased with the number of T. fariai eggs per host, determining a maximum of approximately 14 emerged adults. Host resources per individual affected male and female adult size with similar intensity, and male adult mortality was slightly higher than that for females. These results, and previous findings, suggest that T. fariai attains Hamiltonian sex ratios by laying one male and a variable number of females, and that the detection of chemical marks left by conspecifics provides information on the number of foundresses sharing a patch. Received: 4 February 2000 / Received in revised form: 19 April 2000 / Accepted: 20 May 2000     

Fitness consequences of body-size-dependent host species selection in a generalist ectoparasitoid

In insect parasitoids, offspring fitness is strongly influenced by the adult females choice of host, particularly in ectoparasitoids that attack non-growing host stages. We quantified the fitness consequences of size-dependent host species selection in Dirhinus giffardii, a solitary ectoparasitoid of tephritid fruit fly pupae. We first showed a positive correlation between the size of emerged D. giffardii wasps and the size of their host fruit fly species (in order of decreasing size): Bactrocera latifrons, B. cucurbitae, B. dorsalis or Ceratitis capitata. We then manipulated individual wasps to show that the parasitoid preferred to attack the largest (B. latifrons) to the smallest (C. capitata) host species when provided with a choice, and laid a greater proportion of female eggs in B. latifrons than in C. capitata. There were no differences in developmental time or offspring survival between individuals reared from these two host species. Finally, we compared the foraging efficiency of large versus small wasps (reared from B. latifrons vs C. capitata) under two different laboratory conditions: high versus low host habitat quality, given that realized fecundity in parasitoids may be influenced by either egg-limited or time-limited factors. Under both conditions, large wasps parasitized more hosts than did small ones as a consequence of high searching efficiency in the host-poor habitat, and high capacity for adjusting egg maturation in response to host availability in the host-rich habitat. Considering the flexibility of body growth, the apparent lack of cost of achieving large body size in either development or survival, and the strong dependence of realized reproductive success on a females size, we argue that body size may be a key to understanding evolution of host species selection in ectoparasitoids. We also discuss constraints upon the evolution of size-dependent host species selection in parasitoids.Communicated by D. Gwynne     

Contradictory findings in studies of sex ratio variation in roe deer (Capreolus capreolus)

Patterns of sex ratio variation and maternal investment reported in the literature are often inconsistent. This could be due to intra- and inter-specific variation in social systems, but may also be a result of the a posteriori nature of much of this type of analysis or the testing of models which are inappropriate. Two recent papers reported directly opposed results concerning variation in offspring sex ratios in relation to maternal condition in roe deer, interpreting the results as support for the Trivers and Willard model and for the local resource competition hypothesis, respectively. In this paper, we present data on offspring sex ratios and early juvenile body weight from two long-term studies of this species to test predictions arising from these two models concerning sex biases in litter composition and maternal care. First, we observed no consistent pattern of sex differences in an index of weaning weight or body weight at 1 month old in either population, indicating a lack of sex bias in maternal care. However, in one population, higher maternal body weight was associated with higher juvenile body weight of daughters, but not of sons. Secondly, we found a negative, but not statistically significant, relationship between maternal body weight and litter sex ratio such that heavier females tended to produce more daughters and lighter females to produce more sons. These results indicate that roe females which have additional investment potential available do not invest it in sons, as predicted by the Trivers and Willard model. Our results may provide some support that roe deer are subject to local resource competition acting at the level of the individual mother; however, the fact that particular trends in sex ratio data can be explained in functional terms provides no indication that they are actually adaptive. Received: 9 December 1997 / Accepted after revision: 11 November 1998     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication     

Sex allocation and local mate competition in Old World non-pollinating fig wasps

The populations of many species are structured such that mating is not random and occurs between members of local patches. When patches are founded by a single female and all matings occur between siblings, brothers may compete with each other for matings with their sisters. This local mate competition (LMC) selects for a female-biased sex ratio, especially in species where females have control over offspring sex, as in the parasitic Hymenoptera. Two factors are predicted to decrease the degree of female bias: (1) an increase in the number of foundress females in the patch and (2) an increase in the fraction of individuals mating after dispersal from the natal patch. Pollinating fig wasps are well known as classic examples of species where all matings occur in the local patch. We studied non-pollinating fig wasps, which are more diverse than the pollinating fig wasps and also provide natural experimental groups of species with different male morphologies that are linked to different mating structures. In this group of wasps, species with wingless males mate in the local patch (i.e. the fig fruit) while winged male species mate after dispersal. Species with both kinds of male have a mixture of local and non-local mating. Data from 44 species show that sex ratios (defined as the proportion of males) are in accordance with theoretical predictions: wingless male species<wing-dimorphic male species<winged male species. These results are also supported by a formal comparative analysis that controls for phylogeny. The foundress number is difficult to estimate directly for non-pollinating fig wasps but a robust indirect method leads to the prediction that foundress number, and hence sex ratio, should increase with the proportion of patches occupied in a crop. This result is supported strongly across 19 species with wingless males, but not across 8 species with winged males. The mean sex ratios for species with winged males are not significantly different from 0.5, and the absence of the correlation observed across species with wingless males may reflect weak selection to adjust the sex ratio in species whose population mating structure tends not to be subdivided. The same relationship is also predicted to occur within species if individual females adjust their sex ratios facultatively. This final prediction was not supported by data from a wingless male species, a male wing-dimorphic species or a winged male species. Received: 27 July 1998 / Received in revised form: 11 January 1999 / Accepted: 16 January 1999     

Operational sex ratio versus gender density as determinants of copulation duration in the walnut fly,Rhagoletis juglandis (Diptera: Tephritidae)

In laboratory and field studies of the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), we assessed the effect of operational sex ratio on copulation duration and partitioned the sex ratio effect into component effects due to male density and female density. In our first laboratory experiment, results were clearly consistent with theoretical expectation: increases in male density were associated with significant increases in copulation duration while increases in female density were associated with significant decreases in copulation duration. These component effects yielded a striking composite effect of operational sex ratio (OSR) on copulation duration in which male-biased ratios were associated with low frequencies of short copulations and female-biased ratios were associated with high frequencies of short copulations. Consistent with a priori expectations concerning costs of territorial behavior, the effect of male density on copulation duration was stronger than that of female density. There was no significant interaction between the effects of gender density on copulation duration: each gender density contributed additively to the composite OSR effect on copulation duration. In contrast to the effect of OSR, overall density had little effect. Field data corroborated these findings fully and showed additionally that OSR in the vicinity of fruit tended in nature to be male-biased. In a second laboratory experiment, we measured copulation duration for individuals exposed alternately to male-biased and female-biased ratios. Individual flies consistently copulated for longer in male-biased environments than in female-biased ones. We propose that this plasticity permits individuals to track changes in local sex ratio over space and time and respond appropriately. Received: 15 November 1995/Accepted after revision: 27 April 1996