Differential Mating Success Research Articles

Evolution of sociability by artificial selection.

There has been extensive research on the ecology and evolution of social life in animals that live in groups. Less attention, however, has been devoted to apparently solitary species, even though recent research indicates that they also possess complex social behaviors. To address this knowledge gap, we artificially selected on sociability, defined as the tendency to engage in nonaggressive activities with others, in fruit flies. Our goal was to quantify the factors that determine the level of sociability and the traits correlated with this feature. After 25 generations of selection, the high-sociability lineages showed sociability scores about 50% higher than did the low-sociability lineages. Experiments using the evolved lineages indicated that there were no differences in mating success between flies from the low and high lineages. Both males and females from the low lineages, however, were more aggressive than males and females from the high lineages. Finally, the evolved lineages maintained their sociability scores after 10 generations of relaxed selection, suggesting no costs to maintaining low and high sociability, at least under our settings. Sociability is a complex trait, which we currently assess through genomic work on the evolved lineages.

Pleistocene divergence in the absence of gene flow among populations of a viviparous reptile with intraspecific variation in sex determination.

Polymorphisms can lead to genetic isolation if there is differential mating success among conspecifics divergent for a trait. Polymorphism for sex‐determining system may fall into this category, given strong selection for the production of viable males and females and the low success of heterogametic hybrids when sex chromosomes differ (Haldane's rule). Here we investigated whether populations exhibiting polymorphism for sex determination are genetically isolated, using the viviparous snow skink Carinascincus ocellatus. While a comparatively high elevation population has genotypic sex determination, in a lower elevation population there is an additional temperature component to sex determination. Based on 11,107 SNP markers, these populations appear genetically isolated. “Isolation with Migration” analysis also suggests these populations diverged in the absence of gene flow, across a period encompassing multiple Pleistocene glaciations and likely greater geographic proximity of populations. However, further experiments are required to establish whether genetic isolation may be a cause or consequence of differences in sex determination. Given the influence of temperature on sex in one lineage, we also discuss the implications for the persistence of this polymorphism under climate change.

Towards a predictive conservation biology: the devil is in the behaviour.

One of the most important challenges in conservation biology is to predict the viability of populations of vulnerable and threatened species. This requires that the demographic stochasticity strongly affecting the ecological and evolutionary dynamics of especially small populations is correctly estimated and modelled. Here, we summarize theoretical evidence showing that the demographic variance in population dynamics is a key parameter determining the probability of extinction and also is directly linked to the magnitude of the genetic drift in the population. The demographic variance is dependent on the mating system, being larger in a polygynous than in monogamous populations. Understanding factors affecting intersexual differences in mating success is therefore essential in explaining variation in the demographic variance. We hypothesize that the strength of sexual selection, for example, quantified by the Bateman gradient, may be a useful predictor of the magnitude of the demographic stochasticity and hence the genetic drift in the population. We provide results from a field study of moose that support this claim. Thus, including central principles from behavioural ecology may increase the reliability of population viability analyses through an improvement of our understanding of factors affecting stochastic influences on population dynamics and evolutionary processes. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.

Sex differences in local adaptation: what can we learn from reciprocal transplant experiments?

Local adaptation is of fundamental interest to evolutionary biologists. Traditionally, local adaptation has been studied using reciprocal transplant experiments to quantify fitness differences between residents and immigrants in pairwise transplants between study populations. Previous studies have detected local adaptation in some cases, but others have shown lack of adaptation or even maladaptation. Recently, the importance of different fitness components, such as survival and fecundity, to local adaptation have been emphasized. Here, we address another neglected aspect in studies of local adaptation: sex differences. Given the ubiquity of sexual dimorphism in life histories and phenotypic traits, this neglect is surprising, but may be partly explained by differences in research traditions and terminology in the fields of local adaptation and sexual selection. Studies that investigate differences in mating success between resident and immigrants across populations tend to be framed in terms of reproductive and behavioural isolation, rather than local adaptation. We briefly review the published literature that bridges these areas and suggest that reciprocal transplant experiments could benefit from quantifying both male and female fitness components. Such a more integrative research approach could clarify the role of sex differences in the evolution of local adaptations.This article is part of the theme issue 'Linking local adaptation with the evolution of sex differences'.

Courtship activity, copulation & insemination success in a mosquito vector fed a herbal aphrodisiac: Implications for sterile insect technology.

Background & objectives:In sterile insect technology (SIT), mating competitiveness is a pre-condition for the reduction of target pest populations and a crucial parameter for judging efficacy. Still, current SIT trials are being hindered by decreased effectiveness due to reduced sexual performance of released males. Here, we explored the possible role of a herbal aphrodisiac in boosting the mating activity of Aedes aegypti.Methods:Males were fed one of two diets in this study: experimental extract of Eurycoma longifolia (MSAs) and sugar only (MSOs). Differences in life span, courtship latency, copulation activity and mating success were examined between the two groups.Results:No deaths occurred among MSA and MSO males. Life span of MSOs was similar to that of MSAs. The courtship latency of MSAs was shorter than that of MSOs (P<0.01). MSAs had greater copulation success than MSOs (P<0.001). In all female treatments, MSAs mated more than MSOs, but the differences in rate were significant only in the highest female density (P<0.05). In MSAs, mating success varied significantly with female density (P<0.01), with the 20-female group (P<0.01) having the lowest rate. Single MSA had better mating success at the two lowest female densities. In MSOs, there were no significant differences in mating success rate between the different female densities.Interpretation & conclusions:Our results suggested that the herbal aphrodisiac, E. longifolia, stimulated the sexual activity of Ae. aegypti and may be useful for improving the mating competitiveness of sterile males, thus improving SIT programmes.

Do males with higher mating success invest more in armaments? An across‐populations study in damselflies

1. Males with higher mating success would be expected to invest more in traits that facilitate mating, leading to steeper allometry of those traits with respect to body size. Across‐population studies following latitudinal variation in male mating success are an excellent study system to address this question.2. Males of the damselfly Lestes sponsa were used to investigate whether the allometric patterns of the length and width of the anal appendages, used for grasping the female prior to mating, corresponded to male mating success. Across a large latitudinal gradient, it was hypothesised that there is a larger investment in the grasping apparatus, i.e. a steeper allometric slope, following higher mating success.3. Behavioural observations in field enclosures showed the highest mating success at high latitude, while there were no significant differences in mating success between the central and low latitudes. Positive allometry was found for the length of the anal appendages in high‐latitude males, while central‐ and low‐latitude males showed no significant regressions of the traits on body size.4. These results partially support the hypothesis, as high‐latitude, more successful males invested more in the length (but not the width) of the grasping apparatus than did central‐ and low‐latitude males. Therefore, higher mating success might be facilitated by larger investment in armaments. Intraspecific studies on allometric patterns of traits that participate in mating success might offer new insights into the role of those traits in the reproductive behaviour of a species.

Ecological and Social Factors Constrain Spatial and Temporal Opportunities for Mating in a Migratory Songbird.

Many studies of sexual selection assume that individuals have equal mating opportunities and that differences in mating success result from variation in sexual traits. However, the inability of sexual traits to explain variation in male mating success suggests that other factors moderate the strength of sexual selection. Extrapair paternity is common in vertebrates and can contribute to variation in mating success and thus serves as a model for understanding the operation of sexual selection. We developed a spatially explicit, multifactor model of all possible female-male pairings to test the hypothesis that ecological (food availability) and social (breeding density, breeding distance, and the social mate's nest stage) factors influence an individual's opportunity for extrapair paternity in a socially monogamous bird, the black-throated blue warbler, Setophaga caerulescens. A male's probability of siring extrapair young decreased with increasing distance to females, breeding density, and food availability. Males on food-poor territories were more likely to sire extrapair young, and these offspring were produced farther from the male's territory relative to males on food-abundant territories. Moreover, males sired extrapair young mostly during their social mates' incubation stage, especially males on food-abundant territories. This study demonstrates how ecological and social conditions constrain the spatial and temporal opportunities for extrapair paternity that affect variation in mating success and the strength of sexual selection in socially monogamous species.

Correlational selection on personality and social plasticity: morphology and social context determine behavioural effects on mating success.

Despite a central line of research aimed at quantifying relationships between mating success and sexually dimorphic traits (e.g., ornaments), individual variation in sexually selected traits often explains only a modest portion of the variation in mating success. Another line of research suggests that a significant portion of the variation in mating success observed in animal populations could be explained by correlational selection, where the fitness advantage of a given trait depends on other components of an individual's phenotype and/or its environment. We tested the hypothesis that interactions between multiple traits within an individual (phenotype dependence) or between an individual's phenotype and its social environment (context dependence) can select for individual differences in behaviour (i.e., personality) and social plasticity. To quantify the importance of phenotype- and context-dependent selection on mating success, we repeatedly measured the behaviour, social environment and mating success of about 300 male stream water striders, Aquarius remigis. Rather than explaining individual differences in long-term mating success, we instead quantified how the combination of a male's phenotype interacted with the immediate social context to explain variation in hour-by-hour mating decisions. We suggest that this analysis captures more of the mechanisms leading to differences in mating success. Males differed consistently in activity, aggressiveness and social plasticity. The mating advantage of these behavioural traits depended on male morphology and varied with the number of rival males in the pool, suggesting mechanisms selecting for consistent differences in behaviour and social plasticity. Accounting for phenotype and context dependence improved the amount of variation in male mating success we explained statistically by 30-274%. Our analysis of the determinants of male mating success provides important insights into the evolutionary forces that shape phenotypic variation. In particular, our results suggest that sexual selection is likely to favour individual differences in behaviour, social plasticity (i.e., individuals adjusting their behaviour), niche preference (i.e., individuals dispersing to particular social conditions) or social niche construction (i.e., individuals modifying the social environment). The true effect of sexual traits can only be understood in interaction with the individual's phenotype and environment.

Courtship behavior differs between monogamous and polygamous plovers

Courting, accessing, and/or competing for mates are involved in sexual selection by generating differences in mating success. Although courtship behavior should reflect intensity of mating competition and sexual selection, studies that compare courtship behavior across populations/species with different mating systems subject to differing degrees of mating competition are scanty. Here, we compare courtship behavior between two closely related plover species (Charadrius spp.): a polygamous population of snowy plovers and a socially monogamous population of Kentish plovers. Consistently with expectations, both males and females spent more time courting in the polygamous plover than in the monogamous one. In addition, courtship behavior of males relative to females increased over the breeding season in the polygamous plover, whereas it did not change in the monogamous one. Our results therefore suggest that courtship behavior is a fine-tuned and informative indicator of sexual selection in nature.

Colour polymorphism torn apart by opposing positive frequency-dependent selection, yet maintained in space.

Polymorphic warning signals in aposematic species are enigmatic because predator learning and discrimination should select for the most common coloration, resulting in positive frequency-dependent survival selection. Here, we investigated whether differential mating success could create sufficiently strong negative frequency-dependent selection for rare morphs to explain polymorphic (white and yellow) warning coloration in male wood tiger moths (Parasemia plantaginis). We conducted an experiment in semi-natural conditions where we estimated mating success for both white and yellow male moths under three different morph frequencies. Contrary to expectations, mating success was positively frequency-dependent: white morph males had high relative fitness when common, likewise yellow morph males had high relative fitness when instead they were common. We hence built a model parameterized with our data to examine whether polymorphism can be maintained despite two sources of positive frequency dependence. The model includes known spatial variation in the survival advantage enjoyed by the yellow morph and assumes that relative mating success follows our experimentally derived values. It predicts that polymorphism is possible under migration for up to approximately 20% exchange of individuals between subpopulations in each generation. Our results suggest that differential mating success combined with spatial variation in predator communities may operate as a selection mosaic that prevents complete fixation of either morph.

Sexual selection according to Darwin: A response to Padian and Horner's interpretation

The interpretation of exaggerated structures in the vertebrate fossil record has been hampered by disagreement over the definition of sexual selection and how it relates to sexual dimorphism. Previous assertions that Darwin placed a requirement of sexual dimorphism on sexual selection are mistaken. Instead, Darwin describes variation within one sex and the exertion of a struggle (expressed as intrasexual competition, intersexual mate choice, or both) as the necessary components of sexual selection. The use of structures by one sex to attract mates or repel rivals for mates occurs independently of any existing sexual differences. Differential mating success is also a requirement of Darwinian sexual selection. Mutual sexual selection is a legitimate concept that was described by Darwin. Sexual selection remains a viable explanatory hypothesis for the presence of exaggerated structures of extinct organisms and need not be dismissed summarily, but should not be employed without support as a default hypothesis.

Sexual selection hinders adaptation in experimental populations of yeast

Sexual selection, the suite of processes that lead to differential mating success among individuals, probably influences the evolutionary trajectory of populations. Because sexual selection often shifts traits away from their survival optima, strong sexual selection pressures are thought to increase potential for population extinction, especially during environmental change. Sexual selection pressures may also increase the opportunity for speciation by accelerating the generation of pre-zygotic isolation among populations. These relationships remain largely untested experimentally. Here, we allow populations of baker's yeast, Saccharomyces cerevisiae, to evolve for approximately 250 generations with altered sex ratios in order to test the effect of the strength of sexual selection on the fate of populations. We find that populations experiencing stronger sexual selection are less able to adapt to a novel environment compared with populations experiencing weaker sexual selection or no sex, and that strong sexual selection erases the benefits of sexual reproduction. This pattern persists when fitness is assayed in a closely related environment. We also identify a trend that may suggest the beginning of pre-zygotic isolation between populations experiencing stronger sexual selection, though this is not statistically significant. These results highlight the importance of sexual selection in shaping macroevolutionary patterns and biodiversity.

Mating success and oviposition of a butterfly are not affected by non-lethal tissue sampling

Non-lethal methods of tissue sampling are increasingly used for genetic studies of insect species and the effects of this approach have long been assumed to be minimal. Tissue removal has the potential to influence insect reproductive behaviours such as mate recognition, courtship or oviposition but the effects of non-lethal sampling on reproductive success have not been widely and adequately tested. Here, we test potential effects of both wing-clipping and leg removal on reproductive behaviours of the cabbage white butterfly (Pieris rapae). We conducted a total of 93 male and 59 female mating trials, and found no significant differences in mating success between treated (i.e., tissue removed) and control individuals in either sex. We also monitored the number and location of eggs laid by 58 females. We found no significant differences in egg-laying behaviour among leg removed and control individuals. Power analysis indicated that we had sufficient statistical power to detect moderate effects of treatment on both mating and oviposition. Our study provides the most comprehensive examination to date of the effects of non-lethal sampling on reproductive behaviours in a butterfly/insect species, and supports the contention that tissue sampling is non-detrimental. To fully comprehend the general impacts of tissue sampling on butterfly reproductive behaviour however, additional similar studies need to be conducted on a variety of species with differing mating behaviours. Only through meta-analysis, may it be possible to detect more subtle effects of tissue removal which cannot be revealed within a single study due to sample size limitations.

The relationship between sexual selection and speciation

The intriguing relationship between sexual selection and speciation has sparked an increasing number of papers in recent years that have attempted to draw connections between the two processes. These include a myriad of theoretical and empirical studies, as well as several influential reviews (e.g. Price, 1998; Panhuis et al., 2001, Kirkpatrick and Ravigne, 2002; Ritchie, 2007; Maan and Seehausen, 2011). In particular, many researchers have speculated on whether sexual selection can lead to speciation, either alone, or in conjunction with ecological processes. Such speculation is tempting in part because of the obvious differences in secondary sexual characters that often exist between closely related species. Indeed one of the earliest and most influential of all sexual selection models, Lande’s (1981) paper “Models of speciation by sexual selection on a polygenic trait” centered its title on the role that sexual selection may play in speciation. What, however, is the evidence of a causal relationship between sexual selection and speciation? It turns out to be surprisingly ambiguous. Many have attempted to address a relationship by comparative studies, a process that can be problematic because of both methodological concerns and the fact that the presence of sexual selection is inferred by proxy (Kraaijeveld et al., 2011). Kraaijeveld et al. (2011) point out that these studies only sometimes support a correlation between sexual selection and speciation, and using a meta–analysis find only weak support for a relationship across multiple datasets. A large number of individual experimental studies have also examined various aspects of the intersection of sexual selection and speciation, but it is often unclear what would constitute a true test of the role of sexual selection in speciation, and unambiguous tests of this role are essentially absent (large scale literature analysis by Safran, Mendelson, Scordato and Symes, unpublished data). In particular, the relative roles of ecological factors and sexual selection often appear very intertwined and difficult to tease apart (case studies reviewed in Ritchie, 2007; Maan and Seehausen, 2011). Some studies have attempted to assess whether there is a difference between mate choice and species recognition (e.g. Kozak et al., 2009), but these have also found mixed results. Theoretical studies, which focus on the case of speciation with gene flow in particular, have also demonstrated a very mixed relationship between sexual selection and speciation. Many studies examining the evolution of premating isolation couch the language in terms of sexual selection, simply assuming that species recognition and sexual selection are one and the same in this regard. While this is strictly accurate by the definition of sexual selection as differential mating success, it does not address the spirit of the question of whether sexual selection processes within a species are ultimately responsible for speciation. Several theoretical studies have focused on whether sexual selection alone could account for sympatric speciation, but the general conclusion is that this is unlikely (e.g., van Doorn et al., 2004). Intriguingly, the theoretical literature has also in recent years developed an idea largely absent from the empirical literature, namely that the role of sexual selection in speciation with gene flow may often be inhibitory rather than driving (see discussion in Servedio and Kopp, 2012, this issue). Specifically, sexual selection may often eliminate variation necessary for speciation (e.g., Kirkpatrick and Nuismer, 2004), or lead to stabilizing selection that can counter divergent ecological selection to prevent speciation with gene flow (e.g., Matessi et al., 2001; Otto et al., 2008; Pennings et al., 2008).

By any name, female-female competition yields differential mating success.

By any name, female-female competition yields differential mating success.

Influence of behavior and mating success on brood-specific contribution to fish recruitment in ponds

One source of uncertainty in predicting the response of populations to exploitation is individual differences within a population in both vulnerability to capture and contribution to population renewal. For species with parental care, individuals engaged in nesting behavior are often targeted for exploitation, but predicting outcomes of this nonrandom vulnerability will depend in part on an understanding of how parental traits are related to potential for brood contribution to the population. Variation in brood-specific contribution to recruitment of largemouth bass (Micropterus salmoides), a fish species with extended parental care, was quantified to determine if differences in mating success, parental care behaviors, and timing of reproduction influenced offspring recruitment. Dependence of these relationships on brood predation was tested in communities that differed in the presence of bluegill, Lepomis macrochirus, an important nest predator. Daily snorkel surveys were conducted in experimental ponds during spring to monitor male spawning and parental care behaviors in populations of largemouth bass. Tissue samples collected from larvae in nests were used to develop brood-specific DNA fingerprints for determining nest origins of fall recruits. Largemouth bass spawning period in bluegill ponds was longer and more variable in duration, with lower, more variable mating success, than in ponds without bluegill. In all populations, only one or two broods provided the majority of recruits, and these were broods produced during the earliest days of spawning by the oldest, largest males. In bluegill ponds, brood contribution from earliest nests also increased with brood size. Earliest nesters were the oldest males, and recruits from these nests were often above average in body size. Offspring needed to be guarded to at least swim-up larval stage to contribute any recruits. Termination of parental protection before offspring were free swimming mainly occurred with broods guarded by smaller males in ponds with brood predators. These age- and size-specific differences in timing of spawning and duration of parental care are consistent with influences of residual reproductive value and energetic constraints on reproductive behavior. Furthermore, these patterns of individual contribution to recruitment imply that fisheries that selectively target either nesting individuals or larger, older males could potentially decrease recruitment at the population scale.

Female-biased size dimorphism in a diapausing caddisfly, Mesophylax aspersus: effect of fecundity and natural and sexual selection

1. The effect of mating success, female fecundity and survival probability associated with intra-sex variation in body size was studied in Mesophylax aspersus, a caddisfly species with female-biased sexual size dimorphism, which inhabits temporary streams and aestivates in caves. Adults of this species do not feed and females have to mature eggs during aestivation. 2. Thus, females of larger size should have a fitness advantage because they can harbour more energy reserves that could influence fecundity and probability of survival until reproduction. In contrast, males of smaller size might have competitive advantages over others in mating success. 3. These hypotheses were tested by comparing the sex ratio and body size of individuals captured before and after the aestivation period. The associations between body size and female fecundity, and between mating success and body size of males, were explored under laboratory conditions. 4. During the aestivation period, the sex ratio changed from 1 : 1 to male biased (4 : 1), and a directional selection on body size was detected for females but not for males. Moreover, larger clutches were laid by females of larger size. Finally, differences in mating success between small and large males were not detected. These results suggest that natural selection (i.e. the differential mortality of females associated with body size) together with possible fecundity advantages, are important factors responsible of the sexual size dimorphism of M. aspersus. 5. These results highlight the importance of taking into account mechanisms other than those traditionally used to explain sexual dimorphism. Natural selection acting on sources of variation, such as survival, may be as important as fecundity and sexual selection in driving the evolution of sexual size dimorphism.

Female competition and its evolutionary consequences in mammals

Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter-group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating-system evolution.

Ginger root oil increases mating competitiveness of sterile Mediterranean fruit fly (Diptera:Tephritidae) in Western Australia

AbstractThe exposure of sterile male Mediterranean fruit fly, Ceratitis capitata (Wiedemann) to the aroma of essential oil derived from ginger root Zingiber officinale Roscoe (GRO) has been shown in field cage trials to increase their mating success. This field cage study compared the mating performance of mass reared sterile Mediterranean fruit fly males prepared for ground release programmes under four different post‐irradiation systems, two of which involved exposure of the flies to GRO aroma. In the first system, irradiated pupae were placed into non‐vented 5‐L paper tubs, individually aromatized by 20 μl of GRO (4 ml/m3) for 96–120 h until the adults were 2–3 days old. In the second system, irradiated pupae were placed in vented 5‐L paper tubs, exposed for 24 h in a GRO aromatized room (0.5 ml/m3) when the adults were 3 days old and contained until required for mating tests at 5 days old. The third and fourth systems were the same as the first two systems respectively, except that the flies were not exposed to GRO aroma. Significant differences in mating success were found among treatments. The exposure of 2 to 3‐day old flies to GRO aroma improved mating performance slightly compared with equivalent non‐exposed flies, but it was still below the minimum value accepted under international standards, while 27% of unexposed 5‐day‐old sterile males mated and had competitiveness (Fried value 0.38) slightly above this minimum level. GRO exposed 5‐day‐old flies had the highest mating percentage (67%) and a level of competitiveness (Fried value 1.81) based on induced sterility above that of wild flies (Fried value 1). The results indicate that the effectiveness of sterile male ground release programmes can be increased significantly by releasing 5‐day‐old sterile males that have been exposed to GRO in an entirely aromatized adult fly holding room.

The influence of persistent individual differences and age at maturity on effective population size.

Ratios of effective populations size, N(e), to census population size, N, are used as a measure of genetic drift in populations. Several life-history parameters have been shown to affect these ratios, including mating system and age at sexual maturation. Using a stochastic matrix model, we examine how different levels of persistent individual differences in mating success among males may affect N(e)/N, and how this relates to generation time. Individual differences of this type are shown to cause a lower N(e)/N ratio than would be expected when mating is independent among seasons. Examining the way in which age at maturity affects N(e)/N, we find that both the direction and magnitude of the effect depends on the survival rate of juveniles in the population. In particular, when maturation is delayed, lowered juvenile survival causes higher levels of genetic drift. In addition, predicted shifts in N(e)/N with changing age at maturity are shown to be dependent on which of the commonly used definitions of census population size, N, is employed. Our results demonstrate that patterns of mating success, as well as juvenile survival probabilities, have substantial effects on rates of genetic drift.