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Dispersal influences evolution, demography, and social characteristics but is generally difficult to study. Here we combine long-term demographic data from an intensively studied population of superb fairy-wrens (Malurus cyaneus) and multivariate spatial autocorrelation analyses of microsatellite genotypes to describe dispersal behavior in this species. The demographic data revealed: (1) sex-biased dispersal: almost all individuals that dispersed into the study area over an eight-year period were female (93%; n = 153); (2) high rates of extragroup infidelity (66% of offspring), which also facilitated local gene dispersal; and (3) skewed lifetime reproductive success in both males and females. These data led to three expectations concerning the patterns of fine-scale genetic structure: (1) little or no spatial genetic autocorrelation among females, (2) positive spatial genetic autocorrelation among males, and (3) a heterogeneous genetic landscape. Global autocorrelation analysis of the genotypes present in the study population confirmed the first two expectations. A novel two-dimensional local autocorrelation analysis confirmed the third and provided new insight into the patterns of genetic structure across the two-dimensional landscape. We highlight the potential of autocorrelation analysis to infer evolutionary processes but also emphasize that genetic patterns in space cannot be fully understood without an appropriate and intensive sampling regime and detailed knowledge of the individuals genotyped.

Despite great interest in the use of extra-pair mating as a tool for examining female choice and intersexual selection, the underlying assumption of female control has proved di¤cult to verify empirically. We combined microsatellite genotyping and radiotelemetry of fertile females in order to investigate mate choice in superb fairy-wrens Malurus cyaneus, the bird with the highest known rate of extra-pair fertilization. All ¢ve females radio tracked during the peak of fertility, two to four days before the ¢rst egg is laid, undertook pre-dawn forays. All extra-pair young produced by the female were sired by a male visited during their forays, indicating that females control extra-pair fertilizations. In a larger sample of paternity data, some broods were sired by two extra-group males. In virtually all the cases the territory of the two sires were on an identical linear trajectory from the female's territory. This again suggests that extra-group paternity in superb fairy-wrens is directly linked to female extra-territorial forays. In other species mixed paternity has been taken to indicate that females attempt to insure against infertile pairings or try to maximize the genetic diversity of their brood. However, in fairy-wrens the likelihood of multiple extra-group paternity increased greatly as females traversed more territories in order to mate, perhaps suggesting that females which foray further are more likely to have di¤culties locating the preferred male.

Most birds breed in pairs but at least 3% of passerine species are cooperative breeders, whereby more than two adults help to raise the young. The general rarity of cooperative breeding has led to the assumption that cooperative behaviour has evolved from the ancestral trait of pair breeding. However, it has been suggested that pair breeding may be the derived state in some taxa. The primary aim of this research was to test this suggestion using the genus Acanthiza, which contains examples of both cooperatively and pair breeding species. Mitochondrial DNA sequences were used to construct a phylogenetic hypothesis for the tribe containing Acanthiza, the Acanthizini. The breeding behaviour of the species sequenced was determined from records in the literature; where there were no such data the frequency of another social behaviour, flocking, was used as an indicator of breeding behaviour. The mapping of breeding systems onto the phylogeny led to the conclusion that cooperative breeding is the ancestral state in the Acanthizini, with pair breeding evolving twice in the genus Acanthiza. Models explaining the occurrence of cooperative breeding in terms of broad environmental factors or life history do not appear to be applicable to the genus Acanthiza. The pair breeding Acanthiza species cluster into two clades, suggesting some influence of phylogenetic history on the occurrence of the different breeding systems. Combining the results of this study with other data suggests the tendency to breed cooperatively could be ancestral in the superfamily Meliphagoidea.

Summary Between 1988 and 2001, we studied social relationships in the superb fairy‐wren Malurus cyaneus (Latham), a cooperative breeder with male helpers in which extra‐group fertilizations are more common than within‐pair fertilizations. Unlike other fairy‐wren species, females never bred on their natal territory. First‐year females dispersed either directly from their natal territory to a breeding vacancy or to a foreign ‘staging‐post’ territory where they spent their first winter as a subordinate. Females dispersing to a foreign territory settled in larger groups. Females on foreign territories inherited the territory if the dominant female died, and were sometimes able to split the territory into two by pairing with a helper male. However, most dispersed again to obtain a vacancy. Females dispersing from a staging post usually gained a neighbouring vacancy, but females gaining a vacancy directly from their natal territory travelled further, perhaps to avoid pairing or mating with related males. Females frequently divorced their partner, although the majority of relationships were terminated by the death of one of the pair. If death did not intervene, one‐third of pairings were terminated by female‐initiated divorce within 1000 days. Three divorce syndromes were recognized. First, females that failed to obtain a preferred territory moved to territories with more helpers. Secondly, females that became paired to their sons when their partner died usually divorced away from them. Thirdly, females that have been in a long relationship divorce once a son has gained the senior helper position. Dispersal to avoid pairing with sons is consistent with incest avoidance. However, there may be two additional benefits. Mothers do not mate with their sons, so dispersal by the mother liberates her sons to compete for within‐group matings. Further, divorcing once their son has become a breeder or a senior helper allows the female to start sons in a queue for dominance on another territory. Females that do not take this option face constraints on their ability to recruit more sons into the local neighbourhood.

Summary 1.Correlational studies of reproductive success are plagued by difficulty over the direction of causation. For example, improved reproductive success with age can result from increased experience or reproductive effort, or selection against low-quality phenotypes that survive poorly. An association between supernumeraries and reproductive success in cooperative breeders can arise either because supernumeraries boost productivity, or productive territories accumulate supernumeraries. 2. Paired comparisons of parents sampled with and without supernumeraries have recently been widely applied to quantify help. However, Dickinson & Hatchwell (2004) have argued that this approach is flawed. They conjectured that those groups that gain supernumeraries are a biased superior sample of those that initially lack supernumeraries, while groups that lose supernumeraries will be a sample of inferior cooperative groups. They predict that these biased comparisons will underestimate the effect of help. 3. This conjecture has neither been explored theoretically, nor empirically tested. We use data from a 19-year study of the superb fairy-wren Malurus cyaneus to examine the conjecture and derive predictors of annual reproductive success in this species. 4. We introduce statistical models of reproductive success based on a zero-inflated Poisson link function to identify three strong correlates of reproductive success: high spring rainfall, progress from the first to later years of life, and acquisition of supernumeraries. 5. First year females that died after breeding and those that survived to breed again had similar productivity. As female productivity improves with age, increased reproductive skill or effort is implicated rather than selection against inferior phenotypes. 6. We argue that the Dickinson-Hatchwell conjecture does not constrain paired comparisons in M. cyaneus . The dominant male and breeding female gain no immediate fecundity advantage from supernumeraries. 7. Effects on the future survival of dominants are even more difficult, as while helpers could enhance survival of dominants, a territory that facilitates survival should also accumulate philopatric supernumeraries. Males, the philopatric sex, did not survive better on territories with supernumeraries. However, females, the dispersive sex, had higher survival as the number of supernumeraries increased, because helpers allowed them to reduce the costs of reproduction. These data exacerbate the paradox posed by previously reported costs that supernumeraries impose on dominant males.

Females often mate with several males before producing offspring. Field studies of vertebrates suggest, and laboratory experiments on invertebrates confirm, that even when males provide no material benefits, polyandry can enhance offspring survival. This enhancement is widely attributed to genetic benefits that arise whenever paternity is biased towards males that sire more viable offspring. Field studies suggest that post-mating sexual selection biases fertilization towards genetically more compatible males and one controlled experiment has shown that, when females mate with close kin, polyandry reduces the relative number of inbred offspring. Another potential genetic benefit of polyandry is that it increases offspring survival because males with more competitive ejaculates sire more viable offspring. Surprisingly, however, there is no unequivocal evidence for this process. Here, by experimentally assigning mates to females, we show that polyandry greatly increases offspring survival in the Australian marsupial Antechinus stuartii. DNA profiling shows that males that gain high paternity under sperm competition sire offspring that are more viable. This beneficial effect occurs in both the laboratory and the wild. Crucially, there are no confounding non-genetic maternal effects that could arise if polyandry increases female investment in a particular reproductive event because A. stuartii is effectively semelparous. Our results therefore show that polyandry improves female lifetime fitness in nature. The threefold increase in offspring survival is not negated by a decline in maternal lifespan and is too large to be offset by an equivalent decline in the reproductive performance of surviving offspring.

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