Dual function of allopreening in the cooperatively breeding green woodhoopoe, Phoeniculus purpureus

Allopreening is a widespread but little-studied phenomenon in birds and is particularly prevalent in species where individuals are forced into close proximity. Such a situation facilitates the transfer of ectoparasites between individuals and allopreening has therefore been proposed to serve a hygienic function. In addition, allopreening might theoretically play a role in social communication. Green woodhoopoes (Phoeniculus purpureus) are cooperatively breeding birds that roost communally every night in a tree cavity and are thus susceptible to high ectoparasite loads. Our results suggest that allopreening of the head and neck (“head”), which cannot be efficiently self-preened, serves a primarily hygienic function: all individuals, irrespective of sex, dominance status, and group size, donated and received similar rates of head allopreening and terminated a similar proportion of bouts in which they were involved. Furthermore, there was a high occurrence of reciprocation and head allopreening occurred at a constant rate throughout the year. In contrast, allopreening of the rest of the body, which is accessible to the recipient itself, is likely to serve a primarily social function: body allopreening rates were higher in larger groups; dominant individuals received more body allopreening and terminated a significantly higher proportion of bouts than subordinates; and subordinates donated body allopreening at a higher rate than dominants. Moreover, bouts initiated by dominants were more likely to be reciprocated than those initiated by subordinates and body allopreening rates varied seasonally. Allopreening in the green woodhoopoe is therefore likely to serve a dual function, depending on the part of the body involved.     

Long-term mate and territory fidelity in neotropical buff-breasted wrens (Thryothorus leucotis)

Among sedentary species, mate and site fidelity may be imposed by year-round territoriality and high annual survivorship, both of which theoretically lead to few opportunities to move between mates and breeding territories. We examined mate and territory fidelity in sedentary neotropical buff-breasted wrens (Thryothorus leucotis) in relation to the opportunities to divorce (vacancies on neighbouring territories), breeding experience and site of origin (within vs outside local breeding population). Overall, buff-breasted wrens showed high rates of mate and territory fidelity between years, with individuals significantly more faithful to their territories than to their partners. Mate and territory fidelity was not forced upon buff-breasted wrens, as most individuals had opportunities to switch mates and territories. However, this was not uniform among all individuals. Individuals that had bred together displayed 100% mate and territory fidelity despite the fact that males and females in almost all experienced pairs had opportunities to switch mates and territories. Local recruits (philopatric offspring that acquired territories in the local breeding population) also displayed mate and territory fidelity, but fidelity occurred because recruits rarely had the opportunity to divorce prior to initiating their first breeding attempt. By contrast, almost half of all territorial individuals that originated outside the local breeding population (immigrants) had opportunity to divorce, with approximately two thirds of them switching mates and territories. Divorces occurred within the first 5 months of pairing, suggesting that immigrants may have sampled mates and territories before forming permanent partnerships.     

No evidence for acoustic mate-guarding in duetting buff-breasted wrens (Thryothorus leucotis)

There are few empirical tests of the acoustic mate-guarding hypothesis for the function of duetting in birds. This hypothesis states that when females are fertile, males initiate many songs or answer most of their mates solo songs to form duets and repel rival males seeking extra-pair copulations. We tested the hypothesis by comparing song initiation and answer rates of males and females in socially monogamous buff-breasted wrens (Throthorus leucotis) during pre-fertile and fertile periods. During pre-fertile periods, males often sang for short periods before being answered by their mates, yet first duets were formed earlier relative to dawn and more duets were given during the dawn chorus on pre-fertile than fertile mornings. Males initiated more songs during pre-fertile than fertile periods, whereas there was no difference between stages in female song initiation rates. The proportion of songs answered by individuals of both sexes did not differ between breeding stages. Other mate-guarding behaviours, such as frequent copulation and maintaining close proximity to mates when fertile, did not appear to be important in this species, as no copulations were observed and there was no difference in the time pairs spent in close proximity when females were fertile or not. Parentage analysis revealed that only 3% of 31 broods had young that were likely the result of extra-pair paternity. These findings do not support the acoustic mate-guarding hypothesis, and suggest that the low rate of extra-pair paternity in buff-breasted wrens was maintained without the use of acoustic or traditional paternity guards.Communicated by R. Gibson     

Divorce and extrapair mating in female black-capped chickadees (Parus atricapillus): separate strategies with a common target

Patterns of divorce and extrapair mating can provide insights into the targets of female choice in free-living birds. In resident, site-faithful species with continuous partnerships, the better options and the incompatibility hypotheses provide the most likely explanations for divorce. Extrapair mating can be explained by a number of hypotheses often making similar predictions. For example, the good genes and future partnerships hypo- theses predict similar patterns if males with good genes also make the best future partners. By considering both divorce and extrapair mating, it may be possible to distinguish between these comparable hypotheses. We examined natural patterns of divorce and extrapair mating in a long-term study of black-capped chickadees (Parus atricapillus). Out of 144 partnerships over 8 years, we observed 11 divorces and 38 faithful pairs between seasons. Females usually divorced between their first and second breeding seasons for males of higher social rank than their previous partners, had similar reproductive success prior to divorce as females who retained their previous partners, and did not divorce on the basis of previous reproductive success. These results confirm earlier experimental evidence that females divorce for better options. Females who divorced were significantly more likely to have had mixed-paternity broods prior to divorce than females who stayed with their previous partners. There was no evidence that females divorced in favour of previous extrapair partners. These results support the good genes hypothesis for extrapair mating, suggesting that female chickadees use divorce and multiple mating as separate strategies sharing a common target. Received: 4 February 2000 / Revised: 20 July 2000 / Accepted: 4 September 2000     

Seasonal variation in the duetting behaviour of rufous-and-white wrens (<Emphasis Type="Italic">Thryothorus rufalbus</Emphasis>)

Seasonal variation in animal signalling behaviour has been well documented and has contributed much to our understanding of male signals. In contrast, we know little about seasonal variation in female signals or signals produced jointly by males and females, such as the vocal duets of birds. Here, we examine how singing behaviour changes in relation to time of year and breeding stage in rufous-and-white wrens (Thryothorus rufalbus), neotropical songbirds where both males and females sing and where breeding partners coordinate songs to produce vocal duets. We recorded a colour-marked population of birds over an extended time period encompassing multiple breeding stages. Across all time frames and breeding stages, males sang at higher rates than females and male solos were more common than duets or female solos. Males and females showed divergent seasonal patterns of singing. Females sang more often early in the year, during the pre-breeding season, and female song tapered off as the breeding season progressed. Duetting followed a parallel pattern, which resulted from females showing less duet responsiveness to their partner’s songs later in the year. Male independent song rate peaked at the onset of the rainy season – a time when females become fertile – and males showed the highest level of duet responsiveness during this period. Our results suggest that early in the year, duets appear to be cooperative displays, functioning in joint territory defence and/or the coordination of breeding activities. When females are fertile, however, increased duet responsiveness by males is consistent with mate or paternity guarding.     

Influence of kinship on helping behavior in Galápagos mockingbirds

Summary The social organization of the Galápagos mockingbird (Nesomimus parvulus) in unusual in that groups frequently include more than one breeding pair (plural breeding), and helping behavior is flexible: some birds neither breed nor help, while others do both. To investigate the influence of kinship on helping behavior, I categorized each bird as a helper or non-helper with respect to each nest within its group where it had an opportunity to help. The incidence of helping varied with relatedness: more birds helped when nestlings available to be fed were close relatives than when not. This result was independent of a higher incidence of helping among males than among females and of variation with age among males. Proportionally more nonbreeding than breeding males helped, but breeding and nonbreeding females helped equally infrequently; breeders helped most often after their own nests failed. The incidence of helping was highest among birds with opportunities to feed offspring of breeders that had fed the potential helper as a nestling, suggesting a mechanism for kin discrimination based on associative learning. Juveniles with opportunities to choose among alternative recipients preferentially fed closely related nestlings, but insufficient information was available to determine if adults also did so. Kinship did not influence the rate at which nestlings were fed by helpers. Juveniles fed nestlings at lower rates than did adult helpers, but helping effort was otherwise unaffected by age, sex, or relatedness. Limitation of help to former feeders functions as a mechanism for directing aid to relatives in a plural breeding system where degrees of kinship vary among potential recipients within the same group.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

Relatedness, polyandry and extra-group paternity in the cooperatively-breeding white-browed scrubwren (Sericornis frontalis )

We used DNA fingerprinting to examine the genetic parentage and mating system of the cooperatively breeding white-browed scrubwren, Sericornis frontalis, in Canberra, Australia. Our analyses revealed a remarkable variety of mating tactics and social organization. Scrubwrens bred in pairs or multi-male groups that consisted of a female and two or more males. Females were always unrelated to the pair male or alpha (dominant) male. Among multi-male groups we found three different mating tactics. Firstly, when alpha and beta (subordinate) males were unrelated, they usually shared paternity in the brood. This resulted in both males gaining reproductive benefits directly. Secondly, when beta males were not related to the female but were related to the alpha males, beta males sired offspring in some broods. In this situation, beta males gained reproductive benefits both directly and potentially indirectly (through the related alpha male). Thirdly, when beta males were related to the female or both the female and alpha male, they remained on their natal territory and did not sire any offspring. Thus beta males gained only indirect reproductive benefits. Overall, when group members were related closely, the dominant male monopolized reproductive success, whereas when the members were not related closely the two males shared paternity equally. This positive association between monopolization of reproduction and relatedness is predicted by models of reproductive skew, but has not been reported previously within a single population of birds. Other cooperatively breeding birds with both closely related and unrelated helpers may show a similar variety of mating tactics. Finally, we found that extra-group paternity was more common in pairs (24% of young) than in multi-male groups (6%), and we discuss three possible reasons for this difference. Received: 21 May 1996 / Accepted after revision: 14 December 1996     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

Evolution of extreme-mating behaviour: patterns of extrapair paternity in a species with forced extrapair copulation

Sexual conflict can result in the evolution of extreme mating strategies, including forced copulation. Forced extrapair copulation (FEPC) is generally rare among birds, but is common in re-introduced populations of the hihi (Notiomystis cincta), a socially monogamous, New Zealand endemic, endangered passerine. The aim of this study was to understand the patterns of extrapair paternity in a population where the majority of EPC is forced and to examine the factors, in particular female-specific, influencing the proportion of offspring fathered by extrapair males (EPP—extrapair paternity) and the number of males siring extrapair offspring within a brood (EPM) in this species. Using 8 years of comprehensive paternity, life-history and demographic information for 485 breeding attempts, we show that the frequency of EPP is dependent on (1) social male and female age, (2) the month the female fledged, (3) breeding density and (4) whether it was their first or second reproductive event of the season. In addition, we show that both EPP and EPM are negatively associated with breeding synchrony and clutch size is the most important predictor of EPM. Understanding the drivers of EPP and EPM in species with FECP is important because these are strong determinants of variance in reproductive success and the maintenance of extreme mating behaviour.     

No evidence that sperm morphology predicts paternity success in wild house wrens

Postcopulatory sexual selection (PCSS) in internally fertilizing vertebrates is a topic of great interest, yet relatively little is known about the characteristics of sperm and ejaculates that confer an advantage in PCSS. In this study, we investigated several measures of sperm morphology that potentially contribute to fertilization success under PCSS. We tested whether sperm morphology related to success in PCSS (via extra-pair paternity) in house wrens (Troglodytes aedon). We found no evidence that sperm morphology differed between extra-pair sires and the within-pair males they cuckolded, nor that sperm morphology correlated with the proportion of within-pair offspring sired, the number of extra-pair offspring sired, or the total annual reproductive success. Male behavioral strategies may affect the probability that their sperm compete with other males’ sperm and that their sperm succeed under competition. Effects of these behavioral strategies, as well as differences between males in sperm number, could mask the effects of sperm morphology on the outcome of PCSS. Despite moderate levels of extra-pair paternity, selection on sperm may be relatively weak in house wrens. Further work is needed to understand general patterns in how sperm morphology relates to fertilization success within species.     

Rank-related partner choice in the fission–fusion society of the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>)

When social partners vary in their relative value, individuals should theoretically initiate partnerships with conspecifics of the highest value. Here, we tested this prediction in a wild population of spotted hyenas (Crocuta crocuta). Crocuta live in complex, fission–fusion societies structured by dominance hierarchies in which individuals vary greatly in their value as social companions. Because patterns of association among Crocuta reflect social preferences, we calculated association indices (AIs) to examine how social rank influences intrasexual partner choice among unrelated adults of both sexes. The highest-ranking individuals were generally most gregarious in both sexes. Females associated most often with dominant and adjacent-ranking females. Females joined subgroups based on the presence of particular conspecifics such that subordinates joined focal females at higher rates than did dominants. Dominants benefit from associations with subordinates by enjoying priority of access to resources obtained and defended by multiple group members, but the benefits of these associations to subordinates are unknown. To investigate this, we tested three hypotheses suggesting how subordinates might benefit from rank-related partner choice among unrelated females. We found that subordinates who initiated group formation benefited by gaining social and feeding tolerance from dominants. However, rates at which dominants provided coalitionary support to subordinates did not vary with AIs. Overall, our data resemble those documenting patterns of association among cercopithecine primates. We consider our results in light of optimal reproductive skew theory, Seyfarth’s rank attractiveness model, and biological market theory. Our data are more consistent with the predictions of Seyfarth’s model and of biological market theory than with those of skew theory.     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Unusually high levels of extrapair paternity in a duetting songbird with long-term pair bonds

Previous studies suggest that extrapair young are very rare or absent in socially monogamous avian species that produce vocal duets. These results are generally consistent with functional hypotheses suggesting that duets may signal commitment between partners, or aid males as a paternity guard to ensure genetic as well as social monogamy. Additionally, species that exhibit social monogamy with the same partner across multiple breeding seasons tend to exhibit low levels of extrapair paternity, so duetting species that mate for life may be particularly likely to exhibit genetic monogamy. This study examined the social and genetic mating systems of California towhees (Pipilo crissalis), a duetting species thought to have life-long pair bonds. Observation of a color-banded population confirmed that California towhees exhibit long-term social monogamy. Known social families were genotyped at four microsatellite loci with high allelic diversity. Unexpectedly, paternity exclusion indicated that at least 13 of 31 (42%) nests contained extrapair young. All chicks exhibited maternal alleles, but 21 of 81 (26%) young were not the offspring of social fathers. Thus, in contrast to previous work, this study documents high frequencies of extrapair young among socially monogamous duetting birds with long-term pair bonds.     

Parental facilitation: parent-offspring relations in communally breeding birds

Summary A view of parent-offspring relations is developed that stresses an increased role for mutualism and tolerance rather than conflict in evolutionary trends toward sociality. For example, in Aphelocoma jays, parental tolerance of offspring increases from asocial to social species, contrary to what parental manipulation or parent-offspring conflict models have suggested. A simple model is presented in which parents can leave more grand-offspring while having fewer offspring through a mutualistic relationship between parent and offspring. Consistent with the model are the following observations: (1) At the extreme of sociality, in Mexican jays, A. ultramarina, both sexes commonly inherit their natal territory, thus retaining productive territories largely under control of genotypes that tolerate persistence of young in the territory. Sons and daughters may breed on their natal territory in the presence of their mothers and presumed fathers. (2) Jays banded as nestlings in the study area achieved higher reproductive success when breeding on their natal territory than those that moved to other territories. It is hypothesized that plural breeding can evolve as an extension of the trend leading to singular breeding via a non-altruistic reduction in the number of offspring per parent. The concept of parental facilitation may prove to be important for many highly social types of animals, including insects, birds and mammals.     

Intra-specific interactions influence egg composition in the lesser black-backed gull (Larus fuscus)

Egg composition, which is under maternal control, can have a profound effect on offspring fitness. The presence of maternal testosterone and carotenoids in avian egg yolk, for example, is thought to enhance the development and competitive ability of the offspring and protect the hatching and growing chick against oxidative stress. Egg quality often differs between females and such variation can be due to differences in maternal social environment, e.g. breeding density. However, this is confounded by the possibility that the quality of individuals breeding in high- or low-density areas may vary. We tested if maternal social environment influences egg composition in a colonial seabird, the lesser black-backed gull (Larus fuscus). To control for confounding effects of female quality, we experimentally manipulated maternal social environment during egg formation. We increased the frequency of intra-specific interactions (i.e. aggressive encounters with conspecifics other than nest mates) in which the females were involved, by placing an elevated platform in their territory. Females that took part in more intra-specific interactions produced a heavier last egg, but the yolk testosterone concentration in eggs laid by control and experimental females did not differ. Differences in yolk testosterone concentration in relation to embryo sex were found neither in the control nor in the experimental group. In contrast, within the control group, eggs with a male embryo contained more carotenoids than eggs with a female embryo. Moreover, experimental females that had been involved in more intra-specific interactions produced female eggs with higher carotenoid levels compared to female eggs of control birds. An experimental increase in carotenoid levels was not observed in eggs containing a male embryo. Our results suggest that intra-specific interactions experienced by female birds during egg formation can influence conditions for embryonic development.Communicated by J. Graves     

Indirect genetic benefits of polyandry in a spider with direct costs of mating

The search for the evolutionary explanation of polyandry is increasingly focused on direct and indirect selection on female resistance. In a polyandrous spider Stegodyphus lineatus, males do not provide material benefits and females are resistant to remating. Nevertheless, polyandrous females may obtain indirect genetic benefits that offset the costs associated with multiple mating. We manipulated the opportunity for females to select between different partners and examined the effect of female mating history (mated once, mated twice, or rejected the second male) on offspring body mass, size, condition, and survival under high- and low-food rearing regimens. We found that multiple mating, not female choice, results in increased female offspring body mass and condition. However, these effects were present only in low-food regimen. We did not find any effects of female mating history on male offspring variables. Thus, the benefits of polyandry depend not only on sex, but also on offspring environment. Furthermore, the observed patterns suggest that indirect genetic benefits cannot explain the evolution of female resistance in this system.     

Paternal age and offspring growth: separating the intrinsic quality of young from rearing effects

Younger individuals are often less successful in reproduction than older ones. This might be because of improving breeding skills with age or because the genetic quality of young or early maternal effects on them vary with parental age. However, no attempt has been made to experimentally separate these processes in vertebrates. We conducted a cross-fostering experiment in collared flycatchers (Ficedula albicollis) in three breeding seasons to disentangle origin- and rearing-related effects of paternal age on chick growth, while controlling for date-specific environmental conditions as well as differences in clutch and brood size. The age of the male at the nest of origin, but not that of the rearing male, had a year-dependent effect on nestling body mass and tarsus length. In two seasons, young of subadult males grew slower in the early linear phase of growth than young of adult males. There was no compensatory growth in the final asymptotic phase, so both body mass and tarsus length before fledging reflected the differential early development. In the remaining year, the age of the male at the nest of origin had no significant effect on chick growth. The environment-dependent origin effect we detected was unexplained by incubation times, hatching asynchrony, chick masses at swapping or previously described age-dependent egg quality patterns. Our results therefore suggest a genotype × environment interaction on the relative development of offspring sired by subadult and adult males. Our results also raise the possibility that female birds may gain genetic benefits by mating with older males. Further studies should identify general patterns of male age-dependent female mate choice and offspring quality in different environmental conditions.     

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.