There is an urgent need to thoroughly review and comprehend the effects of urbanization on wildlife in order to understand both the ecological implications of increasing urbanization and how to mitigate its threat to biodiversity globally. We examined patterns in comparative productivity of urban and non-urban passerine birds, using published estimates from paired comparisons, and by reviewing and developing explanations in terms of resources, competitors, predators and other specifically urban environmental factors. The most consistent patterns were for earlier lay dates, lower clutch size, lower nestling weight and lower productivity per nesting attempt in urban landscapes; these were supported by a formal meta-analysis. Nest failure rates did not show consistent patterns across the species considered. We suggest that food availability is a key driver of differences in passerine demography between landscapes. In urban habitats, human-provided food may improve adult condition over winter, leading to earlier lay dates and, in some species, to higher survival and higher breeding densities, but paucity of natural food may lead to lower productivity per nesting attempt. We demonstrate that additional comparative research is needed on a wider range of species, on the effects of natural and human-provided food availability, and on the differences in survival and dispersal between urban and non-urban populations. Importantly, better-targeted research and monitoring is needed in areas that are at greatest threat from urbanization, especially in the developing world.

The ecological constraints hypothesis is widely accepted as an explanation for the evolution of delayed dispersal in cooperatively breeding birds. Intraspecific studies offer the strongest support. Observational studies have demonstrated a positive association between the severity of ecological constraints and the prevalence of cooperation, and experimental studies in which constraints on independent breeding were relaxed resulted in helpers moving to adopt the vacant breeding opportunities. However, this hypothesis has proved less successful in explaining why cooperative breeding has evolved in some species or lineages but not in others. Comparative studies have failed to identify ecological factors that differ consistently between cooperative and noncooperative species. The life history hypothesis, which emphasizes the role of life history traits in the evolution of cooperative breeding, offers a solution to this difficulty. A recent analysis showed that low adult mortality and low dispersal predisposed certain lineages to show cooperative behaviour, given the right ecological conditions. This represents an important advance, not least by offering an explanation for the patchy phylogenetic distribution of cooperative breeding. We discuss the complementary nature of these two hypotheses and suggest that rather than regarding life history traits as predisposing and ecological factors as facilitating cooperation, they are more likely to act in concert. While acknowledging that different cooperative systems may be a consequence of different selective pressures, we suggest that to identify the key differences between cooperative and noncooperative species, a broad constraints hypothesis that incorporates ecological and life history traits in a single measure of 'turnover of breeding opportunities' may provide the most promising avenue for future comparative studies.

Urban development is increasing across the globe. This poses a major threat to biodiversity, which is often relatively poor in towns and cities. Despite much interest in identifying species' traits that can predict their responses to environmental degradation this approach has seldom been used to assess which species are particularly vulnerable to urban development. Here we explore this issue, exploiting one of the best available datasets on species' responses to towns and cities in a highly urbanized region, comprising avian densities across approximately 3000 British urban and rural 1 km  1 km grid cells. We find that the manner in which species' responses to urbanization is measured has a marked influence on the nature of associations between these responses and species' ecological and life history traits. We advocate that future studies should use continuous indices of responses that take relative urban and rural densities into account, rather than using urban densities in isolation, or a binary response recording the presence/ absence of a species in towns and cities. Contrary to previous studies we find that urban development does not select against avian long-distance migrants and insectivores, or species with limited annual fecundity and dispersal capacity. There was no evidence that behavioural flexibility, as measured by relative brain size, influenced species' responses to urban environments. In Britain, generalist species, as measured by niche position rather than breadth, are favoured by urban development as are, albeit to a lesser extent, those that feed on plant material and nest above the ground. Our results suggest that avian biodiversity in towns and cities in urbanizing regions will be promoted by providing additional resources that are currently scarce in urban areas, and developing suitable environments for ground-nesting species.

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The evolution of cooperation among animals has posed a major problem for evolutionary biologists, and despite decades of research into avian cooperative breeding systems, many questions about the evolution of their societies remain unresolved. A review of the kin structure of avian societies shows that a large majority live in kin-based groups. This is consistent with the proposed evolutionary routes to cooperative breeding via delayed dispersal leading to family formation, or limited dispersal leading to kin neighbourhoods. Hypotheses proposed to explain the evolution of cooperative breeding systems have focused on the role of population viscosity, induced by ecological/demographic constraints or benefits of philopatry, in generating this kin structure. However, comparative analyses have failed to generate robust predictions about the nature of those constraints, nor differentiated between the viscosity of social and non-social populations, except at a coarse level. I consider deficiencies in our understanding of how avian dispersal strategies differ between social and nonsocial species, and suggest that research has focused too narrowly on population viscosity and that a broader perspective that encompasses life history and demographic processes may provide fresh insights into the evolution of avian societies.

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An individual's optimal investment in young depends partly on the number of individuals caring for the same brood. In cooperative breeders, the investment strategy of parents with helpers is variable. When parents maintain the same effort regardless of helper number, helper care is additive. When parents fully compensate for the care of helpers by decreasing their own effort, total care does not increase. A study of long-tailed tits Aegithalos caudatus showed that both parental strategies may occur within a species, depending on the number of helpers. A comparative analysis of 27 cooperative breeders was conducted to test the predictions of a graphical model that care is additive when nestling starvation is frequent and parents exhibit compensatory reductions in care when starvation is rare. Both predictions were supported. In this interspecific comparison, a species' mean group size was not associated with compensatory responses by parents. There was some evidence that males and females had different investment rules. Males tended to show compensatory reductions in care when adult survival rate was low. In contrast, while both sexes showed compensation when nestling starvation was infrequent, this association was significant only for females.

The widespread belief that kin selection is necessary for the evolution of cooperative breeding in vertebrates has recently been questioned. These doubts have primarily arisen because of the paucity of unequivocal evidence for kin preferences in cooperative behaviour. Using the cooperative breeding system of long-tailed tits (Aegithalos caudatus) in which kin and non-kin breed within each social unit and helpers are failed breeders, we investigated whether helpers preferentially direct their care towards kin following breeding failure. First, using observational data, we show that not all failed breeders actually become helpers, but that those that do help usually do so at the nest of a close relative. Second, we confirm the importance of kinship for helping in this species by conducting a choice experiment. We show that potential helpers do not become helpers in the absence of close kin and, when given a choice between helping equidistant broods belonging to kin and non-kin within the same social unit, virtually all helped at the nest of kin. This study provides strong evidence that kinship plays an essential role in the maintenance of cooperative breeding in this species.