This study investigated the mating behaviour and parental behaviour of reed buntings (Emberiza schoeniclus), at Rutland Water, Leicestershire. The study integrated behavioural, molecular and anatomical approaches to examine the evolutionary consequences of sperm competition in the species. The frequency of extra-pair paternity (EPP) was extremely high in the study population. 86% (50/58) of broods held at least one extra-pair offspring, whilst 55% (118/216) of young were extra-pair young (EPY). Female participation in extra-pair copulations (EPCs) was virtually ubiquitous at 97% (33/34). Most, i.e., 83% (98/118), EPY were sired by males from an adjacent territory. This pattern of extra-pair paternity was best explained by the indiscriminate female copulatory behaviour associated with the 'genetic diversity' hypothesis. DNA fingerprinting revealed that just over half the males (15/28) breeding in the core study population obtained at least one extra-pair fertilisation (EPFs), and that EPFs accounted for an average of 40% (range 0 - 100%) of a male's reproductive success. There was no relationship between a male's paternity in his own nest and the number of EPFs achieved. Neither was there any consistency in the level of paternity among broods of multiple-brooded pairs. Old males had more EPY in their own nests than young males. There was no relationship between male reproductive success and any of the male phenotypic characters tested in a multivariate analysis. An examination of the paternity protection behaviour of male reed buntings revealed that males did not attempt to expand their territory during the female's fertile period in order to reduce the risk of cuckoldry. There was some evidence of weak mate guarding in the species. It is proposed that the primary paternity guard in the species is through frequent copulations. In the study population, 27% (8/30) of males were polygynous. Males benefit from polygyny through an increased reproductive output on their territories, though not to a significant extent due to a higher level of EPY in secondary nests. Polygynous males fed at only one nest on a territory, thus secondary females incurred a potential cost in terms of increased parental effort. Male and female reed buntings exhibit morphological adaptations associated with a high degree of sperm competition. Males have larger cloacal protuberances, testis and sperm than expected for a bird of comparable size. Females have extremely long sperm storage tubules. Analysis of provisioning behaviour revealed that male feeding rate was significantly related to their level of paternity in the brood. It is proposed that males can assess the confidence of paternity and adjust their feeding rate accordingly. This indicates that there is a potential cost to females of participating in EPCs.

1. This study investigated factors affecting individual reproductive success in the wheatear Oenanthe oenanthe, a migrant passerine species, on Bardsey Island, North Wales, 1991-93 2. Behavioural and paternity data, obtained using DNA fingerprinting, indicated that males pursued a mixed reproductive strategy, i.e. they ensured their paternity during the fertile period of the pair female and pursued extra-pair copulations outwith this time. Behavioural observations were consistent with males using mate guarding to ensure their paternity. Males adjusted their intensity of guarding in response to the threat to their paternity. Males mainly pursued extra-pair copulations once their female had begun incubating. Intrusions by extra-pair males peaked in the fertile period, and later breeding territories were intruded upon more than early breeding pairs. 3. Natural levels of extra-pair paternity were relatively low: 11% of 71 offspring, occurring in 29% of 17 broods. Extra-pair fertilisations contributed little to a male's reproductive success. Not all extra-pair fathers were identified, but at two nests males within a three-territory radius were excluded as extra-pair fathers. There was no evidence that females increased their reproductive success by laying eggs in the nests of other females (intra-specific brood parasitism). Females were never observed off territory soliciting extra-pair males and rarely seen soliciting extra-pair males on territory. The majority of EPCs were resisted by the female and their co-operation appeared to be essential for males to obtain successful copulations. The presence of extra-pair young within broods indicates that females must have also pursued a mixed reproductive strategy by participating in EPCs. 4. Experimental removals of males for 24 hours during the fertile period were used to investigate the effect of the absence of the pair male on: (i) female behaviour; (ii) the behaviour of extra-pair males; and (iii) levels of extra-pair paternity. The number of intrusions and extra-pair copulations increased in the absence of the pair male. Females rejected the majority of extra-pair copulations, and there was no significant increase in extra-pair paternity resulting from these experiments: 10% of 78 offspring occurring in 38% of 16 broods. Female behaviour appeared to be the determining factor affecting the level of extra-pair paternity, although male guarding behaviours may have limited the opportunities for females to participate in extra-pair copulations by deterring intrusions. 5. Males contributed on average 50% of chick feeds, and did not adjust their investment in proportion to their paternity in the brood. There was no effect of the temporary male removals, used to simulate a male's uncertainty of paternity, on their subsequent investment. However, males which adopted broods provided 29% of chick feeds on average, but this was not in proportion to the paternity they had in the brood. This reduction in the number of chick feeds was partially compensated for by females increasing their frequency of chick feeds. 6. Territories remained relatively constant between years. There was evidence that territories varied in quality, as indicated by their consistent order of settlement between years, individuals moving to preferred areas when possible, and individuals being more faithful to preferred areas. Older males returned to the breeding ground earlier than first- year males, and were more likely to be paired than individuals breeding for the first time. This was probably due to the older males having settled on preferred territories. Territory quality had a significant effect on individual reproductive success. There were few correlates with individual quality and measures of breeding success. A male's mating status was dependent on arrival time, territory quality, breeding density and the operational sex ratio. Males which settled on preferred territories were more likely to be paired. Pairs which bred on preferred territories tended to have increased fledging success, and nestlings which fledged from preferred territories were more likely to return to the study area to breed. Female reluctance to copulate outwith the pair bond may be a result of individual reproductive success being determined more by territory quality rather than male quality.

1) This study investigated the reproductive strategies employed by European starlings (Sturnus vulgaris) nesting in two separate study areas in England and another near Wellington, New Zealand. Evidence from behavioural observations and single-locus DNA profiling was used to establish the occurrence and frequency of alternative reproductive strategies within this generally socially monogamous species. 2) In total, 8% (42/526) of nestlings were sired by an extra-pair male and 28% of broods (37/134) contained at least one extra-pair young (EPY). There were no cases of extra-pair males gaining more than 50% of the paternity within a clutch. The distribution of EPY was best explained by the 'genetic diversity' hypothesis. 19% of the males breeding within the English study areas were paired to more than one female and extra- pair paternity (EPP) was significantly more common in the broods of these polygynous males. Nevertheless polygynous males sired significantly more fledglings than males paired to a single female. EPP was equally common in the broods of primary and secondary females. 3) An examination of the possible methods of paternity protection employed by male starlings failed to find evidence for strong mate guarding. Males followed a greater proportion of female-initiated departure flights during the female's fertile period (81 - 94%). However, in each study area males actively left their females unguarded during the female's fertile period. Frequent copulation is suggested to be the primary paternity guard in starlings. 4) The frequency of intraspecific brood parasitism (IBP) was determined through regular nest inspections and DNA profiling. Double laying was detected in 9% (34/394) of all clutches monitored. The hatching success of eggs thought to be parasitic was significantly lower than that of eggs laid by host femmes. DNA profiling revealed that 2% (12/576) of chicks were parasitic and 7% (11/154) of broods contained at least one parasitic chick. A single case of communal nesting was found. I suggest that IBP is the result of nest desertion principally caused by intraspecific disturbance and is a 'best-of-a- bad-job' strategy. 5) The existence of anti-parasitism strategies such as nest guarding, egg removal, nest desertion or retaliatory parasitism was investigated. Behavioural observations revealed that the intensity of nest guarding did not increase during the laying period, the time when potential hosts are most at risk from parasitic activity. The experimental presentation of a female during the laying period failed to increase the intensity of nest guarding. An egg addition experiment did not apparently induce parasitic behaviour in host birds and the host clutch size was not significantly different to the clutch size in non-experimental nests. Egg removal was no more frequent in experimental nests than neighbouring nests or nests elsewhere in the study area. The apparent absence of anti-parasitism behaviours in starlings is discussed. 6) The influence of breeding density, colony size and nearest neighbour distance on the frequency of IBP, EPP and intraspecific disturbance was investigated. There was no evidence to suggest that the composition of colonies of different sizes or nest distribution differed relative to male age. The mean frequency of EPP among nests was significantly negatively correlated with nearest neighbour distance. The frequency of intraspecific disturbance also showed a significant negative correlation with increasing inter-nest distance. No relationship was found between the frequency of EPP and the degree of breeding synchrony. IBP was significantly less frequent in the asynchronously laid intermediate clutches. The distribution of EPP among nests is discussed against a background of frequent copulation as the principal paternity guard.

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Global climate change, and associated habitat destruction and degradation, comprise 2 of the most important drivers of declines in biodiversity, and understanding their impact upon threatened taxa is an important objective for conservation initiatives. The farmland and migratory bird assemblages of Europe have both declined dramatically over the last century, and improving our understanding of the effects of climate and land use change upon them is of import when implementing successful conservation interventions. We focused upon the European Roller (Coracias garrulus), a migratory farmland species which has experienced large-scale population declines and range contractions across Europe, investigating the effects of habitat and climate change upon the species’ breeding biology and distribution. We modelled the effects of climatic conditions at key phenological points in the species’ annual cycle upon breeding parameters across its Mediterranean range and found that temperature comprised the key climatic correlate of clutch size and fledging success. This suggests that changing temperatures will lead to declines in Roller productivity across the Mediterranean. We produced a Habitat Suitability Model for the species on Cyprus and confirmed that the island’s Protected Area network incorporates a higher proportion of high value Roller habitat than external areas. We also identified high value areas currently external to the Protected Area network. We used novel Global Positioning System technology to quantify Roller home ranges in Cyprus and to identify optimal and avoided foraging habitat types. The importance of traditional farmland and grassland habitats for Rollers was confirmed, as was the negative impact of urban habitat upon Roller productivity. Nest site selection was also investigated for Rollers in Cyprus and Portugal, with our results indicating that selection occurs at the cavity level rather than landscape scale. Rollers in both populations preferred nest sites with smaller entrances, potentially due to decreased predation risk. High levels of nest site competition were identified in Portugal, and evidence of nest site limitation in both the Cypriot and Portuguese Roller populations was observed. Key overall recommendations from the study comprise the promotion of traditional farmland and grassland habitats within European agri-environment schemes, and the extension/initialisation of artificial nest site schemes for the Roller in Cyprus and Portugal.

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Grey-necked Picathartes, Picathartes oreas, is among 349 species of African birds that are globally threatened with extinction. With the current deforestation and fragmentation of its habitat, this study was undertaken to assess population status and habitat requirements in one such forest fragment, the Mbam Minkom Mountain Forest in southern Cameroon. 90 breeding sites with 207 nests and 24 potential breeding sites were found. From 34 nests with signs of breeding activity, the entire population was estimated to be 68 breeding individuals. Populations were highest in the north west region but had declined from 40 breeding individuals in 2003 to 20 in 2007.

In cooperatively breeding species, 'helpers' assist in raising non-descendant offspring. In many cooperative species, delayed dispersal of offspring provides the permissive conditions for kin-directed helping and helpers gain indirect fitness benefits by enhancing productiviry of non-descendant kin. However, kin-directed cooperation may also arise via other routes, where offspring disperse before becoming helpers, but limited dispersal and demographic processes allow kin association within extended networks of relatives. In such systems, kin discrimination may be important for individuals to maximise fitness by directing care towards relatives and to avoid the risk of inbreeding. In this thesis I investigated the cooperative breeding system of the rifleman Acanthisitta chloris in a study population at Kaikoura, New Zealand. Routes to helping were diverse and flexible, and helpers were not constrained by age, sex or breeding status. In contrast to previous reports, I showed riflemen helpers were closely related to the recipients of their care. Furtherrnore, adult helpers were associated with increased offspring recruitment and thus, could gain substantial indirect fitness benefits. The social and genetic background against which helping operates was also investigated. Both sexes showed localised natal dispersal resulting in positive fine-scale genetic structure. Whilst this may promote sociality in the rifleman, it also generated a significant risk of inbreeding risk. However, the relatedness of pairs was lower than expected under biologically realistic models of random mating, suggesting active inbreeding avoidance through mate choice. The proximate mechanism of kin recognition was investigated but r found no evidence of active kin recognition using experimental playbacks of vocalisations. Overall, this thesis shows that cooperation riflemen occurs among relatives living in kin networks that arise through limited dispersal by both sexes. The increased productivity of helped broods indicates that kin selection has played an important role in the evolution of helping in the rifleman.

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