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1. The social environment in which individuals live affects their fitness and in turn population dynamics as a whole. Birds with facultative cooperative breeding can live in social groups with dominants, subordinate helpers that assist with the breeding of others, and subordinate non-helpers. Helping behaviour benefits dominants through increased reproductive rates and reduced extrinsic mortality, such that cooperative breeding might have evolved in response to unpredictable, harsh conditions affecting reproduction and/or survival of the dominants. Additionally, there may be different costs and benefits to both helpers and non-helpers, depending on the time-scale. For example, early-life costs might be compensated by later-life benefits. These differential effects are rarely analysed in the same study. 2. We examined whether helping behaviour affects population persistence in a stochastic environment and whether there are direct fitness consequences of different life-history tactics adopted by helpers and non-helpers. 3. We parameterised a matrix population model describing the population dynamics of female Seychelles warblers Acrocephalus sechellensis, birds that display facultative cooperative breeding. The stochastic density-dependent model is defined by a (st)age structure that includes life-history differences between helpers and non-helpers and thus can estimate the demographic mechanisms of direct benefits of helping behaviour. 4. We found that population dynamics are strongly influenced by stochastic variation in the reproductive rates of the dominants, that helping behaviour promotes population persistence and that there are only early-life differences in the direct fitness of helpers and non-helpers. 5. Through a matrix population model, we captured multiple demographic rates simultaneously and analysed their relative importance in determining population dynamics of these cooperative breeders. Disentangling early-life versus lifetime effects of individual tactics sheds new light on the costs and benefits of helping behaviour. For example, the finding that helpers and non-helpers have similar lifetime reproductive outputs and that differences in reproductive values between the two life-history tactics arise only in early life suggests that overall, helpers and non-helpers have a similar balance of costs and benefits when analysing direct benefits. We recommend analysing the consequence of different life-history tactics, during both early life and over the lifetime, as analyses of these different time frames may produce conflicting results.

1. In many cooperatively breeding animals, a combination of ecological constraints and benefits of philopatry favours offspring taking a subordinate position on the natal territory instead of dispersing to breed independently. However, in many species individuals disperse to a subordinate position in a non-natal group ("subordinate between-group" dispersal), despite losing the kin-selected and nepotistic benefits of remaining in the natal group. It is unclear which social, genetic and ecological factors drive between-group dispersal. 2. We aim to elucidate the adaptive significance of subordinate between-group dispersal by examining which factors promote such dispersal, whether subordinates gain improved ecological and social conditions by joining a non-natal group, and whether between-group dispersal results in increased lifetime reproductive success and survival. 3. Using a long-term dataset on the cooperatively-breeding Seychelles warbler (Acrocephalus sechellensis), we investigated how a suite of proximate factors (food availability, group composition, age and sex of focal individuals, population density) promote subordinate between-group dispersal by comparing such dispersers with subordinates that dispersed to a dominant position or became floaters. We then analysed whether subordinates that moved to a dominant or non-natal subordinate position, or became floaters, gained improved conditions relative to the natal territory, and compared fitness components between the three dispersal strategies. 4. We show that individuals that joined another group as non-natal subordinates were mainly female and that, similar to floating, between-group dispersal was associated with social and demographic factors that constrained dispersal to an independent breeding position. Between-group dispersal was not driven by improved ecological or social conditions in the new territory and did not result in higher survival. Instead, between-group dispersing females often became co-breeders, obtaining maternity in the new territory, and were likely to inherit the territory in the future, leading to higher lifetime reproductive success compared to females that floated. Males never reproduced as subordinates, which may be one explanation why subordinate between-group dispersal by males is rare. 5. Our results suggest that subordinate between-group dispersal is used by females to obtain reproductive benefits when options to disperse to an independent breeding position are limited. This provides important insight into the additional strategies that individuals can use to obtain reproductive benefits.

How the environment influences the transmission and prevalence of disease in a population of hosts is a key aspect of disease ecology. The role that environmental factors play in host-pathogen systems has been well studied at large scales, that is, differences in pathogen pressures among separate populations of hosts or across land masses. However, despite considerable understanding of how environmental conditions vary at fine spatial scales, the effect of these parameters on host-pathogen dynamics at such scales has been largely overlooked. Here, we used a combination of molecular screening and GIS-based analysis to investigate how environmental factors determine the distribution of malaria across the landscape in a population of Berthelot's pipit (Anthus berthelotii, Bolle 1862) on the island of Tenerife (Canary Islands, Spain) using spatially explicit models that account for spatial autocorrelation. Minimum temperature of the coldest month was found to be the most important predictor of malaria infection at the landscape scale across this population. Additionally, anthropogenic factors such as distance to artificial water reservoirs and distance to poultry farms were important predictors of malaria. A model including these factors, and the interaction between distance to artificial water reservoirs and minimum temperature, best explained the distribution of malaria infection in this system. These results suggest that levels of malaria infection in this endemic species may be artificially elevated by the impact of humans. Studies such as the one described here improve our understanding of how environmental factors, and their heterogeneity, affect the distribution of pathogens within wild populations. The results demonstrate the importance of measuring fine-scale variation - and not just regional effects - to understand how environmental variation can influence wildlife diseases. Such understanding is important for predicting the future spread and impact of disease and may help inform disease management programmes as well as the conservation of specific host species.

1.Fluctuations in the quality of the habitat in which an animal lives can have major consequences for its behaviour and physiological state. In poor-quality habitat with low food availability, metabolically intensive foraging activity is likely to result in increased generation of reactive oxygen species, while scarcity of food can lead to a weakening of exogenously derived antioxidant defences. The consequent oxidant/antioxidant imbalance may lead to elevated oxidative stress. 2.Although the link between food availability and oxidative stress has been studied in the laboratory, very little is known about this relationship in the wild. Here, we investigate the association between territory quality (measured through food availability) and oxidative stress in the Seychelles warbler (Acrocephalus sechellensis). 3.Seychelles warblers are insectivorous birds that inhabit a fixed feeding territory year round. Individuals experience profound and rapid local fluctuations in territory quality within these territories, owing to changing patterns of vegetation defoliation resulting from seasonal changes in prevailing wind direction and wind-borne salt spray. 4.As expected, oxidant generation (measured as reactive oxygen metabolites; ROMs) was higher when territory quality was low, but there was no correlation between territory quality and antioxidant capacity (OXY). The negative correlation between territory quality and ROMs was significant between individuals and approached significance within individuals, indicating that the pattern resulted from individual responses to environmental variation. 5.ROMs and OXY levels within individuals were positively correlated, but the relationship between territory quality and ROMs persisted after including OXY as a covariate, implying that oxidative stress occurs in low territory quality conditions. 6.Our results indicate that the oxidative stress balance of an individual is sensitive to relatively short-term changes in territory quality, which may have consequences for the birds’ fitness.

1. Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and variation in fitness in tropical bird species.2. To investigate whether variation in survival could be explained by population size or climatic variables we used capture-recapture models in conjunction with a long-term data set from an island population of the territorial, cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). The lack of migration out of the study population means that our results are not confounded by dispersal.3. Annual survival was high, both for adults (84%) and juveniles (61%), and did not differ between the sexes. Although there was significant variation in survival between years, this variation could not be explained by overall population size or weather variables.4. For territorial species, resource competition will work mainly on a local scale. The size of a territory and number of individuals living in it will therefore be a more appropriate measure of density than overall population density. Consequently, both an index of territory quality per individual (food availability) and local density, measured as group size, were included as individual covariates in our analyses.5. Local density had a negative effect on survival; birds living in larger groups had lower survival probabilities than those living in small groups. Food availability did not affect survival.6. Our study shows that, in a territorial species, although density-dependent effects might not be detectable at the population level they can be detected at the individual territory level - the scale at which individuals compete. These results will help to provide a better understanding of the small-scale processes involved in the dynamics of a population in general, but in particular in tropical species living in relatively stable environments.