Sex ratio varies with egg investment in the red-necked phalarope (Phalaropus lobatus)

Fisher’s sex ratio theory predicts that on average parents should allocate resources equally to the production of males and females. However, when the cost/benefit ratio for producing males versus females differs, the theory predicts that parents may bias production, typically through underproduction of the sex with greater variation in fitness. We tested theoretical predictions in the red-necked phalarope, a polyandrous shorebird with sex-role reversal. Since females are larger and therefore potentially more expensive to produce and may have greater variation in reproductive success, we predicted from Fisher’s hypothesis a male bias in population embryonic sex ratio, and from sex allocation theory, female biases in the clutches of females allocating more resources to reproduction. We measured eggs and chicks and sexed 535 offspring from 163 clutches laid over 6 years at two sites in Alaska. The embryonic sex ratio of 51.1 M:48.9 F did not vary from parity. Clutch sex ratio (% male) was positively correlated with clutch mean egg size, opposite to our prediction. Within clutches, however, egg size did not differ by sex. Male phalarope fitness may be more variable than previously thought, and/or differential investment in eggs may affect the within-sex fitness of males more than females. Eggs producing males were less dense than those producing females, possibly indicating they contained more yolk relative to albumen. Albumen contributes to chick structural size, while yolk supports survivorship after hatch. Sex-specific chick growth strategies may affect egg size and allocation patterns by female phalaropes and other birds.     

Multiple paternity, sperm storage, and reproductive success of female and male painted turtles (Chrysemys picta) in nature

When females receive no direct benefits from multiple matings, concurrent multiple paternity is often explained by indirect genetic benefits to offspring. To examine such possibilities, we analyzed genetic paternity for 1,272 hatchlings, representing 227 clutches, from a nesting population of painted turtles (Chrysemys picta) on the Mississippi River. Goals were to quantify the incidence and distribution of concurrent multiple paternity across clutches, examine temporal patterns of sperm storage by females, and deduce the extent to which indirect benefits result from polyandrous female behaviors. Blood samples from adult males also allowed us to genetically identify the sires of surveyed clutches and to assess phenotypic variation associated with male fitness. From the genetic data, female and male reproductive success were deduced and then interpreted together with field data to evaluate possible effects of female mating behaviors and sire identity on offspring fitness. We document that more than 30% of the clutches were likely fathered by multiple males, and that presence of multiple paternity was positively correlated with clutch size. Furthermore, the data indicate that the second male to mate typically had high paternity precedence over the first.     

Sexual dimorphism in house sparrow eggs

Recent evidence has revealed an apparently high degree of control by female birds over the physiological aspects of their reproduction and offspring sex allocation, consistent with adaptive hypotheses of sex allocation and differential investment in their offspring. In the house sparrow, we investigated possible mechanisms that may be used by females to enhance the fitness returns from a reproductive effort. Using molecular techniques, we demonstrate that house sparrow eggs containing male embryos are significantly larger than those containing female embryos. We also found that male embryos were laid randomly with respect to laying order. We speculate that this sexual dimorphism of eggs is adaptive, because male house sparrows show greater variance in condition-dependent reproductive success than females. More important, the result provides further evidence of the ability of females to detect or control ovulation of either male or female ova and to differentially invest in one sex over the other. Received: 19 January 2000 / Revised: 29 June 2000 / Accepted 20 July 2000     

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.     

Interactive effects of male and female age on extra-pair paternity in a socially monogamous seabird

Females sometimes obtain older sires for their offspring through extra-pair interactions, but how female age influences paternity is largely unexplored and interactive effects across the age span of both sexes have not been analyzed. To test whether female choice of sire age varies with female age in the blue-footed booby (Sula nebouxii), we examined associations between ages of both partners and the probability of extra-pair paternity (EPP) in 350 broods of parents up to 22 years old in a single breeding season. Extra-pair paternity enables a female to select an alternative sire for her offspring and could function to avoid or achieve particular combinations of parental ages. A male age?×?female age interaction revealed that in young females (≤4 years), EPP decreased with increasing age of the social partner, whereas in old females (≥8 years), it increased. Moreover, sires of extra-pair (EP) chicks of young females paired to young males were on average 6.33 years older than the females’ social partners. Since female boobies control copulatory access, this pattern could imply that young females choose old sires for their proven genetic quality and that old females avoid very old males because matings with them may risk infertility or genetic defects in offspring. Taking female age into account and observing across the whole age span may be necessary for understanding female age-based mate choice.     

An adaptive annual rhythm in the sex of first pigeon eggs

When the reproductive value of male and female offspring varies differentially, parents are predicted to adjust the sex ratio of their offspring to maximize their fitness (Trivers and Willard, Science 179:90–92, 1973). Two factors have been repeatedly linked to skews in avian offspring sex ratio. First, laying date can affect offspring sex ratio when the sexes differ in age of first reproduction, such that the more slowly maturing sex is overproduced early in the season. Second, position of the egg in the laying sequence of a clutch may affect sex ratio bias since manipulating the sex of the first eggs may be least costly to the mother. We studied both factors in two non-domesticated pigeon species. Both the Wood pigeon (Columba palumbus) and the Rock pigeon (Columba livia) have long breeding seasons and lay two-egg clutches. In the field, we determined the sex of Wood pigeon nestlings. In Rock pigeons, housed in captivity outdoors, we determined embryo sex after 3 days of incubation. On the basis of their sex-specific age of first reproduction, we predicted that males, maturing at older age than females, should be produced in majority early and females later in the year. This was confirmed for both species. The bias was restricted to first eggs. Rock pigeons produced clutches throughout the year and show that the sex of the first egg followed an annual cycle. To our knowledge, this study presents the first evidence of a full annual rhythm in adaptive sex allocation in birds. We suggest that this reflects an endogenous seasonal program in primary sex ratio controlled by a preovulatory mechanism.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Experimental assessment of ecological and phenotypic factors affecting male mating success and polyandry in northern watersnakes, Nerodia sipedon

To resolve conflicting field observations regarding the action of sexual selection, we used breeding experiments and paternity analysis of the 927 resulting offspring to assess how male size, condition, tail length, genetic similarity to the female, and variation in operational sex ratio (OSR) affected male reproductive success and the incidence of polyandry in northern watersnakes (Nerodia sipedon). Only size affected male mating success. Large males were more successful, but only when male size varied substantially and competition among males was intense (i.e., male-biased OSR). The conditional nature of the size advantage may explain why studies of free-living watersnakes have produced inconsistent results regarding the relationship between male size and mating success. Size differences between males did not affect the proportion of offspring each male sired within multiply sired litters. We found positive size-assortative mating, but only when the OSR was female biased, suggesting that smaller males had improved access to females when competition among males was reduced, but that competition with larger males still restricted mating opportunities of small males to less preferred, smaller females. Most litters (58%) were multiply sired and larger females were more likely to produce multiply sired litters, similar to free-living watersnakes. There was no association between the incidence of multiple paternity and OSR, however, suggesting that polyandry is not simply a function of opportunity, with females passively waiting for males to court them.     

Sex allocation and paternity in a cooperatively breeding passerine: evidence for the male attractiveness hypothesis?

Sex allocation theory predicts that female birds with high-quality mates will benefit from producing more sons, since sons will inherit their father’s superior traits and enjoy a great reproductive success, whereas females with low-quality mates will benefit from producing more daughters, since the variance in reproductive success among daughters is typically lower. The male attractiveness hypothesis may apply to extra-pair paternity (EPP) because socially monogamous females routinely mate with higher quality males outside the pair bond. We test these predictions using the Tibetan ground tit (Pseudopodoces humilis), a sexually monomorphic, socially monogamous, facultatively cooperative breeder. There was greater variation in actual reproductive success among males than females due to EPP. An excess of sons was detected for bi-parental (i.e., non-cooperative) broods wherein EPP was mainly sired by bi-parental males. The pattern was attributed to male-biased sex ratios produced for both EPP and within-pair offspring within the same broods. The reason for the latter case might be a random allocation of more offspring to sons by the potentially EPP-exposed females that have an inability to control fertilization by specific males. In cooperative broods where EPP mostly resulted from within-group helpers of presumed low-quality, as indicated by their failure in acquiring a social mate, there was a non-significant tendency for EPP offspring to be daughters and for within-pair offspring in the same broods to be unbiased. These results support the EPP-related male attractiveness hypothesis especially in terms of the overproduction of sons. Offspring produced through quasi-parasitism was unbiased towards either sex, suggesting a weak female choiceness with respect to the quality of host males.     

Variation in offspring sex ratio among individual Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>) females of different quality

The Trivers–Willard model predicts that in polygynous species, superior-quality females will maximize their fitness by producing male offspring. Using a sample of 1,780 Weddell seal (Leptonychotes weddellii) pups recorded over 31 years, we investigated relationships between offspring sex ratio and maternal age, reproductive experience, an index of maternal lifetime reproductive output, and annual environmental variations. We found evidence that females with higher index of lifetime reproductive output were more likely to produce male than female offspring but found only weak evidence that large-scale environmental variations influenced sex ratios. Our results suggest that mothers manipulate offspring sex to maximize their own fitness, and inherent maternal quality may influence offspring sex. These findings support the Trivers–Willard sex-allocation model. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Body size and sexual selection in the koala

Sexual selection is often characterized by polygynous breeding systems, size dimorphism, and skewed operational sex ratios. Koalas are sexually dimorphic in multiple domains, yet are absent from the literature on sexual selection and the structure of their mating system is unclear. We provide the first documentation of the strength of sexual selection in koalas by using microsatellite markers to identify sires. We combine the genetic data with morphological data in order to assess the role of body size in regulating reproductive output. During our 4-year study, 37% of males were identified as possible sires. Males were significantly larger than females, with sires heavier than non-sires. Male body mass correlated with annual reproductive output, with Crow’s Index of Opportunity for Selection revealing that variation in male reproductive success was threefold higher than that of females. Since it appears that male koalas rarely engage in physical confrontations over access to females, size dimorphism could be based upon non-agonistic competition and/or female mate choice. We propose that size dimorphism in koalas evolved as a consequence of endurance rivalry promoting vocal sexual advertisements that attract females. We suggest that female choice is a key mediator of male reproductive output.     

Sex allocation in Savi’s warblers <Emphasis Type="Italic">Locustella luscinioides</Emphasis>: multiple factors affect seasonal trends in brood sex ratios

Sex allocation theory predicts that whenever the relative fitness of sons and daughters differ, females should invest more in the sex with the greatest fitness return. In this study, we evaluated the influence of various ecological factors on the brood sex ratio (BSR) of Savi’s warblers (Locustella luscinioides) across several breeding seasons. There was a slight but significant female production bias at the population level, which is consistent with the ‘local resource competition’ hypothesis, as the breeding density is very high and females are more prone to disperse. We found that there was a significant decline in BSR during the breeding season, but no influence of male size, female size, social status nor extra-pair paternity were detected. The seasonal decline in BSR was further evaluated by assessing the within- and between-female effects, which indicated that multiple factors were operating simultaneously in our study population. First, there was a significant within-female decline in BSR, which was consistent with the decline in female condition due to the reproductive effort associated with multiple brooding (supporting the Trivers and Willard hypothesis). Second, a significant decline in BSR with the laying date of first clutches of different pairs indicated that male and/or female qualities are also associated with the seasonal variation in BSR. Finally, a comparison between the sex of the youngest nestling with the remaining ones did not suggest any bias, indicating that females do not compensate for the increased mortality of the last nestling (caused by asynchronous hatching) by producing a male from the last laid egg.     

Polygyny and male parental care in Savannah sparrows: effects on female fitness

Summary To determine the effects of male mating status on female fitness, we compared the reproductive success, survival, and future fecundity of female Savannah sparrows (Passerculus sandwichensis) mated to monogamous vs. polygynous males in a 5-year study on Kent Island, New Brunswick, Canada. The proportion of males with more than one mate varied from 15 to 43% between years and sites. Polygynous and monogamous males fledged young of equal size in every year of the study. Females who shared paternal care with other females laid as many eggs per clutch and clutches per season as monogamously mated females. In most years polygynously mated females showed no delay in laying a second clutch, and they suffered no reduction in fecundity the following year. Recruitment of a female's offspring into the breeding population was generally independent of her mating status. Fitness costs of being mated to a polygynous male were only apparent in one year of the study, during which females mated to polygynous males had higher over-winter mortality than those mated to monogamous males. That same year, young raised by polygynous males were only one-third as likely to survive to reproductive maturity (as inferred by returns) as those raised by monogamous males. A male's mating status had no effect on his own survivorship. A male's mating status did not necessarily reflect his contributions to raising nestlings, which may partially explain why monogamously and polygynously mated females had equal fitness. At 35 nests the proportion of food deliveries brought by individual males varied from 0 to 75%; on average, males brought fewer than 30% of all food deliveries. Yet parental care by polygynous males was no less than that of monogamous males, at least at the nests of their primary females. Secondary females tended to receive less male assistance during the nestling stage, but their reproductive success was indistinguishable from that of primary females. Females feeding young without male assistance made as many food deliveries/h as did pairs in which males brought at least 30% of all food deliveries. Unassisted females did not suffer diminished fledging success or produce smaller fledglings. The benefits of polygyny for male Savannah sparrows are clear: polygynous males recruit more surviving offspring into the breeding population than monogamous males. The fitness of females, on the other hand, appears to be unaffected by whether their mate was monogamous or polygynous except in occasional years. Polygyny may be maintained in this population by the constraints of a female-biased sex ratio, the inability of females to predict a male's paternal care based on his morphology or behavior, the poor correlation between a male's mating status and his assistance at the nest, and inconsistent natural selection against mating with a polygynous male. Correspondence to: N.T. Wheelwright     

Female-coerced monogamy in burying beetles

The reproductive interests of the sexes often do not coincide, and this fundamental conflict is believed to underlie a variety of sex-specific behavioral adaptations. Sexual conflict in burying beetles arises when a male and female secure a carcass that can support more offspring than a single female can produce. In such a situation, any male attracting a second female sires more surviving offspring than he would by remaining monogamous, whereas the female's reproductive success decreases if a rival female is attracted to the carcass. Monogamously paired males on large carcasses do in fact attempt to attract additional females by means of pheromone emission, whereas males on small carcasses do not. Females physically interfere with male polygynous signaling using various behavioral tactics. We demonstrate that such interference leads to a significant decrease in the amount of time that males spend signaling, according females a means by which to impose monogamy on their mates.     

Population and individual consequences of breeding resource availability in the European bitterling (Rhodeus amarus)

Resource availability may affect both individual fitness and population demography and the effects can interact. We used two experiments to test how breeding resource abundance and its spatial distribution, combined with female abundance, affected male reproductive behavior, population spawning rate, and embryo development and recruitment in the European bitterling (Rhodeus amarus), a small cyprinid fish that lays its eggs in living unionid mussels. In the first experiment, we found that at the population level the abundance of breeding resources (freshwater mussels) was more important for bitterling recruitment than resource spatial distribution (clumped or regular). In contrast at the individual level, (variability in reproductive success) the spatial distribution of resources was more important, but only when resource abundance was not limiting. Territorial males obtained almost exclusive access to fertilizations when resources were abundant and distributed regularly, but were unable to defend large clusters of resources (when rival abundance was always high) and abandoned territoriality. Surprisingly, territorial males remained aggressive and successfully defended their territories when resources were grouped into a single cluster, but at a low abundance. In the second experiment, more rapid embryo development and larger juvenile body size at the end of the growing season was detected at high resource abundance and low female abundance, indicating that early hatched juveniles survived better and hence investment in offspring production early in the season yields a higher fitness pay-offs. The abundance of females in spawning condition was the best overall predictor of the intensity of male reproductive behavior in both experiments.     

Paternity analysis shows experience,not age,enhances mating success in an aquatically mating pinniped,the Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>)

For polygynous mammals with no paternal care, the number of offspring sired is often the sole measure of male reproductive success. The potential for polygyny is highest when resources or other environmental factors such as restricted breeding sites force females to aggregate. In these circumstances, males compete intensely for females and mating success may vary greatly among males, further intensifying selection for those traits that confer an advantage in reproduction. Hence, determinants of male success in competition for females are likely to be under strong sexual selection. Paternity analysis was used in conjunction with measures of age, site fidelity, and behavior during the breeding season to assess variance in male breeding success in Weddell seals (Leptonychotes weddellii) breeding at Turtle Rock, McMurdo Sound (77.727S, 166.85E) between 1997 and 2000. Paternity could be assigned to 177 pups at relaxed or 80% confidence level or 111 pups at strict or 95% confidence levels. Weddell seals at Turtle Rock show a modest degree of polygyny with the greatest number of pups sired by any individual male in a single season equalling 5 or ∼10% of the pups born. Over four consecutive years, most (89.2%) males sired at least one pup. In a generalized linear model (GLM), age and the age first seen at the study site as an adult were unrelated to mating success, but adult experience, either site-specific or elsewhere in McMurdo Sound, over the reproductive life span of males explained nearly 40% of variance in total mating success with 80% confidence and 24% of variance at 95% confidence. While learning where females are likely to be may enhance male reproductive success, aquatic mating reduces the ability of males to monopolize females, and thereby increases equity in mating success.     

Mixed reproductive strategy and mate guarding in a semi-colonial passerine,the swallow Hirundo rustica

Summary Both male and female swallows Hirundo rustica have a mixed reproductive strategy (parental care for offspring and extra-pair couplations). Mate guarding protects females from male harassment and male swallows from being cuckolded. Females hide their fertile period by copulating successfully with their mates for an extended period during first clutches. Males guard in the pre-fertile period, when many unpaired males are present. Early breeding swallows guard more than late breeders since more sexual chases of females by non-mate males take place in the early period. Solitarily breeding females experience few chases by strange males; copulation frequency, length of copulation period and mate guarding is adjusted to a lower level than among colonial birds. Male guarding activity is more intense in the fertile than in the pre-fertile period. Guarding reduces success of extra-pair copulation attempts.Three female swallows each paired and copulated with a single male and later changed to a new male before starting to breed. Extra-pair copulations most often take place between neighbouring swallows in the fertile period of the female. Many old, early breeding males and many young, late breeding females participate in extra-pair couplations. Successful extra-pair copulations peak in the fertile period contrary to success of pair copulations which does not change during the copulation period.     

Seasonal decline in reproductive performance varies with colony size in the fairy martin, <Emphasis Type="Italic">Petrochelidon ariel</Emphasis>

Reproductive success within populations often varies with the timing of breeding, typically declining over the season. This variation is usually attributed to seasonal changes in resource availability and/or differences in the quality or experience of breeders. In colonial species, the timing of breeding may be of particular importance because the costs and benefits of colonial breeding are likely to vary over the season and also with colony size. In this study, we examine the relationship between timing of breeding and reproductive performance (clutch size and nest success) both within and between variable sized colonies (n = 18) of fairy martins, Petrochelidon ariel. In four of these colonies, we also experimentally delayed laying in selected nests to disentangle the effects of laying date and individual quality/experience on reproductive success. Within colonies, later laying birds produced smaller clutches, but only in larger colonies. The general seasonal decline in nest success was also more pronounced in larger colonies. Late laying birds were generally smaller than earlier laying birds, but morphological differences were also related to colony size, suggesting optimal colony size also varies with phenotype. Experimentally delayed clutches were larger than concurrently produced non-delayed clutches, but only in larger colonies. Similarly, delayed clutches were more likely to produce fledglings, particularly later in the season and in larger colonies. We suggest that the reduced performance of late breeding pairs in larger colonies resulted primarily from inexperienced/low quality birds preferring to settle in larger colonies, possibly exacerbated by an increase in the costs of coloniality (e.g., resource depletion and ectoparasite infestations) with date and colony size. These findings highlight the importance of phenotype-related differences in settlement decisions and reproductive performance to an improved understanding of colonial breeding and variation in colony size.     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich