In many cooperatively breeding birds, kin selection has an important role in the evolution and maintenance of social behaviour, and 'helpers' can maximize indirect fitness gains by preferentially allocating care to close relatives. Although there is evidence for kin-biased helping behaviour in several species, the mechanism of kin recognition underlying this behaviour is poorly understood. Vocalizations are the most commonly used cues in avian recognition systems, but the effectiveness of vocal signals as reliable recognition cues must depend on how they are acquired. However, there have been no experimental studies of the development of vocal recognition cues in cooperative birds; indeed, the ontogeny of all bird vocalizations other than song is poorly known in any species. Here, we show that cooperatively breeding long-tailed tits (Aegithalos caudatus) can discriminate between kin and non-kin according to the individual-specific characteristics of contact calls, and show experimentally that individuals learn these calls from provisioning adults during the nestling period. Finally, we show that the pattern of cooperative behaviour in this species is consistent with the use of recognition cues learned through association.

Summary1. The aim of this study was to investigate the structure and thermoregulatory function of nests of the Long-Tailed Tit, Aegithalos caudatus . 2. The feather lining of Long-Tailed Tit nests represents a major portion (41%) of the total nest mass. 3. The mass of feathers varied among nests and declined through the breeding season, but there was no seasonal loss of nest insulation quality because of increasing ambient temperatures. 4. In an experiment to investigate the seasonal decline in the feather mass of nests, feathers were added to nests at an early stage of the lining phase of nest construction. Nest structure and insulating properties were then examined following nest completion. 5. The total mass of feathers in treatment and control nests did not differ significantly and there was no significant difference in their nest insulation quality. 6. Our results demonstrate that Long-Tailed Tits adjust their nest-building behaviour according to the nest's thermal environment. Moreover, nest structure appears to be adjusted to prevailing environmental conditions rather than being a function of feather availability or time constraints.

Urban areas are expanding rapidly, but a few native species have successfully colonized them. The processes underlying such colonization events are poorly understood. Using the blackbird Turdus merula, a former forest specialist that is now one of the most common urban birds in its range, we provide the first assessment of two contrasting urban colonization models. First, that urbanization occurred independently. Second, that following initial urbanization, urban-adapted individuals colonized other urban areas in a leapfrog manner. Previous analyses of spatial patterns in the timing of blackbird urbanization, and experimental introductions of urban and rural blackbirds to uncolonized cities, suggest that the leapfrog model is likely to apply. We found that, across the western Palaearctic, urban blackbird populations contain less genetic diversity than rural ones, urban populations are more strongly differentiated from each other than from rural populations and assignment tests support a rural source population for most urban individuals. In combination, these results provide much stronger support for the independent urbanization model than the leapfrog one. If the former model predominates, colonization of multiple urban centres will be particularly difficult when urbanization requires genetic adaptations, having implications for urban species diversity.

Despite increasing interest in urban ecology most attention has focussed on describing changes in assemblage composition and structure along urbanisation gradients, whilst relatively little research has focussed on the mechanisms behind these changes. Ecological theory predicts that alterations in biotic interactions are particularly likely to arise, especially with regard to disease risk. Here, we report on differences in prevalence of avian malaria and tick infection and intensity in 11 paired urban and rural blackbird Turdus merula populations from across the western Palearctic. We find large and consistent reductions in tick prevalence and intensity in urban areas. There are also large reductions in the prevalence of avian malaria in many, but not all, urban areas. The proportion of first year birds in urban populations is significantly lower than that in rural ones, and across the more natural rural sites southerly populations contain fewer first years than northern ones. These patterns are expected to arise if survival rates are higher in urban areas, and are negatively correlated with latitude.

Urban areas occupy a large and growing proportion of the earth. Such sites exhibit distinctive characteristics relative to adjacent rural habitats, and many species have colonised and now successfully exploit urban habitats. The change in selection pressures as a result of urbanisation has led to trait divergence in some urban populations relative to their rural counterparts, but studies have generally been local in scale and the generality of differentiation thus remains unknown. The European blackbird Turdus merula is one of the commonest urban bird species in the Western Palearctic, but populations vary substantially in the length of time they have been urbanised. Here we investigate patterns of morphological variation in European blackbirds occupying 11 paired urban and rural habitats across much of the urbanised range of this species and spanning 258 of latitude. First, we assessed the extent to which urban and rural blackbirds are differentiated morphologically and the consistency of any differentiation across the range. Paired urban and rural Blackbird populations frequently exhibited significant morphological differences, but the magnitude and direction of differentiation was site dependent. We then investigated whether the nature of latitudinal gradients in bodysize differed between urban and rural populations, as predicted by differences in the climatic regimes of urban and rural areas. Blackbird body-size exhibited strong latitudinal gradients, but their form did not differ significantly between urban and rural habitats. The latitudinal gradient in body size may be a consequence of Seebohm's rule, that more migratory populations occurring at high latitudes have longer wings. We conclude that while there can be substantial morphological variation between adjacent urban and rural bird populations, such differentiation may not apply across a species' range. Locality specific differentiation of urban and rural blackbirds may arise if the selection pressures acting on blackbird morphology vary in an inconsistent manner between urban and rural habitats. Alternatively, phenotypic divergence could arise in a stochastic manner depending on the morphological traits of colonists, through founder effects.

Long-tailed tits Aegithalos caudatus are cooperative breeders in which helpers exhibit a kin preference in their cooperative behaviour. We investigated the mechanism through which this preference is achieved by ¢rst conducting an experiment for testing whether breeders could recognize the calls of their relatives while controlling for spatial e¡ects. We found that there were signi¢cant di¡erences in the responses of breeders to the vocalizations of kin and non-kin, suggesting that vocal cues may be used for kin recognition. We conducted a second experiment in order to investigate whether recognition is achieved on the basis of relatedness per se or through association. Nestlings were cross-fostered between unrelated broods in order to create broods composed of true and foster siblings. In subsequent years, survivors from experimental broods did not discriminate between true and fostered siblings when making helping decisions, indicating that recognition is learned and not genetically determined. We discuss the e¡ectiveness of learning through association as an indirect cue to kinship.

For organisms with temperature-dependent sex determination (TSD), skewed offspring sex ratios are common. However, climate warming poses the unique threat of producing extreme sex ratio biases that could ultimately lead to population extinctions. In marine turtles, highly female-skewed hatchling sex ratios already occur and predicted increases in global temperatures are expected to exacerbate this trend, unless species can adapt. However, it is not known whether offspring sex ratios persist into adulthood, or whether variation in male mating success intensifies the impact of a shortage of males on effective population size. Here, we use parentage analysis to show that in a rookery of the endangered green turtle (Chelonia mydas), despite an offspring sex ratio of 95 per cent females, there were at least 1.4 reproductive males to every breeding female. Our results suggest that male reproductive intervals may be shorter than the 2 -4 years typical for females, and/or that males move between aggregations of receptive females, an inference supported by our satellite tracking, which shows that male turtles may visit multiple rookeries. We suggest that male mating patterns have the potential to buffer the disruptive effects of climate change on marine turtle populations, many of which are already seriously threatened.

Somatic growth is an integrated, individual-based response to environmental conditions, especially in ectotherms. Growth dynamics of large, mobile animals are particularly useful as bio-indicators of environmental change at regional scales. We assembled growth rate data from throughout the West Atlantic for green turtles, Chelonia mydas, which are long-lived, highly migratory, primarily herbivorous mega-consumers that may migrate over hundreds to thousands of kilometers. Our dataset, the largest ever compiled for sea turtles, has 9690 growth increments from 30 sites from Bermuda to Uruguay from 1973 to 2015. Using generalized additive mixed models, we evaluated covariates that could affect growth rates; body size, diet, and year have significant effects on growth. Growth increases in early years until 1999, then declines by 26% to 2015. The temporal (year) effect is of particular interest because two carnivorous species of sea turtles-hawksbills, Eretmochelys imbricata, and loggerheads, Caretta caretta-exhibited similar significant declines in growth rates starting in 1997 in the West Atlantic, based on previous studies. These synchronous declines in productivity among three sea turtle species across a trophic spectrum provide strong evidence that an ecological regime shift (ERS) in the Atlantic is driving growth dynamics. The ERS resulted from a synergy of the 1997/1998 El Niño Southern Oscillation (ENSO)-the strongest on record-combined with an unprecedented warming rate over the last two to three decades. Further support is provided by the strong correlations between annualized mean growth rates of green turtles and both sea surface temperatures (SST) in the West Atlantic for years of declining growth rates (r = -.94) and the Multivariate ENSO Index (MEI) for all years (r = .74). Granger-causality analysis also supports the latter finding. We discuss multiple stressors that could reinforce and prolong the effect of the ERS. This study demonstrates the importance of region-wide collaborations.