The Australian magpie (Gymnorhina tibicen) is a large, sedentary, omnivorous passerine. In some populations, individuals live in groups, and some of these groups breed cooperatively. The white-backed magpie (G. t. tyrannica) from the south-eastern corner of the continent, has had relatively little study, and few details are known of its mating system, social structure, and method of parental care. I conducted an observational study on a population of white-backed magpies, recording details of their demography, dispersal, breeding system, and parental care. In conjunction, I conducted a genetic analysis of the population, to determine if the genetic mating system matched the observed social system, to detect instances of extra-group mating, and to sex juvenile birds. Extra-pair paternity (EPP) is a common feature of the mating systems of many birds. The rate of EPP may vary between species, races and populations. I made a comparison of extra-group paternity (EGP) rates between two races of the Australian magpie, to determine if similar mating systems were being employed. The two populations had similar social structure, but differed in group size and dispersal. I predicted that dispersal differences would have a profound effect on the rate of EGP between the populations, as the population with the lower rate of dispersal and higher chance of breeding with a close relative would engage in EGPs more frequently. Eight microsatellite loci were used to determine parentage in the white-backed Australian magpie. The rate of EGP was found to be 44%. Dispersal rates were estimated from observational data. Over half of the juvenile magpie cohort from the previous breeding season leave the territorial group. These results contrast sharply with the results found by other researchers in a population of western Australian magpies (G t. dorsalis). In this population, 82% EGP is recorded and dispersal of juveniles is close to nil. The results indicate that dispersal rate is a potentially important predictor of rates of extra-group fertilisations between populations of this species, and suggest that females maximise their reproductive output by avoiding breeding with close kin. The reproductive success of a male bird is often correlated with measurable traits that predict his intrinsic quality. Females are thought to select mates based on their quality to gain their 'good genes'. Male Australian magpies of the white-backed race were trapped in two breeding seasons. Measurements were taken of morphometric and other characteristics in order to discover whether particular traits of males were associated with: a) number of fledglings produced in the territory per season; b) percentage of offspring sired in the territory; and c) whether females select males for their 'good genes'. The only variable that was correlated with number of territorial offspring was feather lice load. Males with high numbers of lice were less likely to produce territorial fledglings in one season and across both seasons. Males of inferior quality may be subject to increased conspecific territorial intrusions, leading to more time spent on defence, more failed breeding attempts, less time allocated to grooming and thus high parasite loads. Males that produced many territorial fledglings were more likely to gain genetic paternity of at least some of them, although again this was significant for only one season. Also, across both seasons, a high number of females in the group was correlated with increased paternity within the group. The general lack of correlation between the variables and level of genetic paternity may be due to females engaging in extra-group mating primarily to avoid breeding with a close relative rather than to choose a quality male. In this scenario, males would not have to be 'high quality', but merely genetically different to the female's social mate. Extra-group paternity (EGP) can affect paternal effort. It may also influence the helping effort of auxiliary birds in cooperatively breeding species. If helping is driven by kin selection, helpers should decline to provision unrelated young. Relatedness becomes difficult to assess however, when females mate outside the group. Alternative rewards may then become important in helper decisions. In my study population of Australian magpies, 38% of fledglings were sired by males outside the territorial group. In a second population (G. t. dorsalis), 82% of fledglings were sired by extra-group males. I observed within-group male and helper feeding effort over three breeding seasons in the first population and obtained data recorded over a single season in the second population. In both populations, males provisioned young regardless of relatedness, as did helpers. Males provisioned less than the nesting female on average. Paternal effort did not reduce with an increase in the rate of EGP between populations. In the population with intermediate levels of EGP, the white-backed magpies, I observed helpers in about half of the sampled territories that produced fledglings. Helpers did not increase the production of young. In the population with high levels of EGP, western magpies, I detected helping behaviour in proportionally more territories. It appears that Australian magpie helpers provide help in order to pay 'rent' and remain on the natal territory. I discuss these results in light of the differences between the two races of magpie and the major theories regarding male parenting decisions and helper activity. Finally, I examine the relatively high rates of EGP's in the Australian magpie from a phylogenetic perspective. Although inbreeding avoidance is strongly supported by this study as the major reason EGP is so common in magpie populations, there may be an element of phylogenetic inertia that maintains the frequency of this behavioural trait. I comment upon the use of single-population estimates of species EGP rates in comparative analyses, given the intraspecific variation discovered between Australian magpie populations. Future directions for the study of mate choice in the Australian magpie are outlined with a proposal to study variation at the major histocompatibility complex between mated pairs.

The evolution of extra-group paternity (EGP) is a contentious issue in evolutionary biology. This thesis examines the factors and adaptive benefits driving EGP in a high-density, group-living population of European badgers (Meles meles). To improve power to assign parentage, I isolated and characterised 21 new polymorphic microsatellite markers. I genotyped 83% of 1410 badger trapped 1987‒2010 using 35 autosomal microsatellite markers. Maternity and paternity were assigned at 80% confidence ca. 82% of individuals. 48% of paternities were extra-group, where 85% were attributable to neighbouring-group males and EGP was detected in 47% of litters; thus badger social group do not correspond with a breeding unit. I tested whether indirect genetic benefits explain these high EGP rates. (1) ‘Good-gene-as-heterozygosity Hypothesis’: Paternal heterozygosity, but not maternal or an individual’s own heterozygosity, associated positively with first-year survival probability. Under benign environmental conditions, cubs fathered by more heterozygous males had a higher first year survival probability. Despite this correlation, the EGP rate per litter correlated with neither average nor maximum within-group heterozygosity of candidate fathers. (2) Fitness benefit Hypothesis: Extra-group offspring (EGO) had lower first-year survival probability and lived 1.3 years less than within-group offspring (WGO). Female WGO produced more litters and offspring over their lifetime than female EGO, whereas male EGO produced more offspring than male WGO. (3) Inbreeding avoidance hypothesis: The EGP rate within a litter increased with greater average pair-wise relatedness between mothers and within-group candidate fathers. No inbreeding depression on first-year survival probability was detected, but small sample sizes limited statistical power. Socio-ecologically, at the litter level, EGP correlated negatively with the number of within-group candidate fathers, and positively with neighbouring-group candidate fathers. In conclusion, EGP in badgers may reduce inbreeding and be maintained in the population through a sex-specific antagonistic selection and indirect genetic benefits may occur when the total fitness benefits of producing extra-group sons outweigh the costs of producing extra-group daughters. These indirect genetic benefits only partially explain the evolution of promiscuity in European badgers, highlighting that evolutionary factors underlying promiscuity remain unclear.

Studies of mating systems and social organisation have been central to understanding of the evolution of social behaviour. The European badger Meles meles is a good species in which to study these processes, as its complex social system provides an opportunity to investigate how both natural and kin selection shape the evolution of mating systems and social structure. In this thesis, I use behavioural and genetic data to describe the mating system and social organisation of a high-density badger population and examine the occurrence of cooperative breeding. I genotyped 915 (85%) badgers trapped in Wytham Woods (1987–2005), 630 of which were cubs, and assigned both parents to 331 cubs with 95% confidence. This revealed a polygynandrous mating system, with up to five mothers and five fathers per social group. Mounting behaviour was also polygynandrous and I show the strongest evidence to date for multiple-paternity litters. I demonstrate, for the first time, that groups consisted of close and distant kin: approximately one third of group members were first-order kin, and overall group members had slightly lower relatedness levels than half-siblings. Within groups, adult and yearling females had higher pairwise relatedness than males, and neighbouring groups contained relatives. These findings result from the high level (42%) of extra-group paternities, 86% of which were assigned to neighbouring males. For the first time I show that females avoided inbreeding by mating with extra-group males; however, incestuous matings did occur. Promiscuous and repeated mountings were observed, which may reduce male–male aggression and infanticide, but may also promote sperm competition, genetic diversity, and / or genetic compatibility. Just under a third of adult males and females were assigned parentage each year and I quantify, for the first time, reproductive skew within badger groups. Correlations between relatedness, group productivity, and reproductive skew were not consistent with the predictions of incomplete-control models; rather, resource availability may play a role. Older and younger badgers displayed reduced annual breeding success, with male success increasing initially with experience. The Restraint, Constraint, and Selection Hypotheses did not explain the age-related breeding pattern in females. Variance in lifetime breeding success (LBS) was greater for males. Males that only bred within or only outside of their groups had half the LBS of males that did both. Females that were assigned maternity probably bred cooperatively and allonursed non-offspring, which has not been demonstrated previously. No benefit was established, however, in terms of litter size, probability of offspring breeding, or offspring lifetime breeding success, with more mothers in a group. In conclusion, badger social groups are fostered through kinship ties. Polygynandry and repeated mounting may have evolved originally to reduce male–male aggression and infanticide by males, through paternity masking. Although plural breeding occurs, group living appears to be costly. Motivation to disperse may be reduced through high-levels of extra-group paternities, which may also reduce inbreeding. Cooperative breeding among mothers may represent a low-cost behaviour with indirect benefits due to high levels of relatedness between female group-members. Badger sociality therefore represents an early stage in the evolution of social behaviour.