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In species with biparental and cooperative brood care, multiple carers cooperate by contributing costly investments to raise a shared brood. However, shared benefits and individual costs also give rise to conflict among carers conflict among carers over investment. Coordination of provisioning visits has been hypothesized to facilitate the resolution of this conflict, preventing exploitation, and ensuring collective investment in the shared brood. We used a 26-year study of long-tailed tits, Aegithalos caudatus, a facultative cooperative breeder, to investigate whether care by parents and helpers is coordinated, whether there are consistent differences in coordination between individuals and reproductive roles, and whether coordination varies with helper relatedness to breeders. Coordination takes the form of turn-taking (alternation) or feeding within a short time interval of another carer (synchrony), and both behaviors were observed to occur more than expected by chance, that is, “active” coordination. First, we found that active alternation decreased with group size, whereas active synchrony occurred at all group sizes. Second, we show that alternation was repeatable between observations at the same nest, whereas synchrony was repeatable between observations of the same individual. Active synchrony varied with reproductive status, with helpers synchronizing visits more than breeders, although active alternation did not vary with reproductive status. Finally, we found no significant effect of relatedness on either alternation or synchrony exhibited by helpers. In conclusion, we demonstrate active coordination of provisioning by carers and conclude that coordination is a socially plastic behavior depending on reproductive status and the number of carers raising the brood.

Abstract Cooperative breeding sometimes occurs when adult breeders form groups following natal dispersal and mating. In such cases, individuals typically face a choice of social partner with whom to cooperate. Selecting appropriate social partners is crucial to maximising the fitness payoffs from cooperation, but our understanding of the criteria guiding partner choice is limited. Here, we analyse helping decisions by long-tailed tits (Aegithalos caudatus), which may redirect their care to assist breeders in raising offspring following the failure of their own nests. In this species, helpers prefer to help relatives at nearby nests, but it is unclear whether other criteria that may affect helper fitness also influence helping decisions. When choosing among broods of equivalent kinship, we found that helpers did not prefer those broods that offered the greatest indirect fitness returns. Further analyses revealed that helpers did not choose nests on the basis of brood size or age, but were more likely to help broods that were closer to their own failed nests and that were already being cared for by other helpers. Both effects likely reflect the limited choice available to helpers: although individuals breed close to relatives within kin neighbourhoods, a high rate of nest predation constrains helpers’ choice of broods. In other species where cooperatively breeding groups form after natal dispersal, a greater range of options may be available and here detailed analysis of group formation will be helpful for determining the decision rules that underpin partner choice and permit stable cooperation in the face of alternative options. Significance statement Cooperative breeding occurs most frequently when offspring delay dispersal from their natal site and help to care for their younger siblings. In some species, however, individuals first disperse and then come together as adults to cooperate in rearing young. In the latter case, multiple social partners may be available — what then determines which partner is helped? We studied partner choice in long-tailed tits, which may help to feed other broods if their own brood fails. When multiple related broods were available, individuals were more likely to help those close by but showed no preference for broods offering the greatest indirect fitness returns. One explanation for this result is that helping options for most individuals are limited by high levels of nest predation, favouring a simpler decision-making process based on identifying close relatives breeding in close proximity.

The genetic structure of animal populations has considerable behavioural, ecological and evolutionary implications and may arise from various demographic traits. Here, we use observational field data and molecular genetics to determine the genetic structure of an invasive population of monk parakeets, Myiopsitta monachus, at a range of spatial scales, and investigate the demographic processes that generate the observed structure. Monk parakeets construct large nests that can house several pairs occupying separate chambers; these nests are often aggregated within nesting trees. We determined patterns of relatedness within compound nests, within nesting trees and between trees. Spatial autocorrelation analyses of pairwise genetic relatedness revealed fine-scale genetic structure with relatives of both sexes spatially clustered within, but not beyond, nesting trees. In addition, males were more related to males sharing their compound nests than to other males occupying the same nesting tree. By contrast, males and females within compound nests were not significantly more closely related than elsewhere in the same tree, and we found no evidence for inbreeding. Adults showed high breeding site fidelity between years despite considerable disturbance of nest sites. Natal dispersal was female-biased, but dispersal distances were relatively short with some natal philopatry observed in both sexes. Sibling coalitions, typically of males, were observed amongst both philopatric and dispersing birds. Our results show significant clustering of kin within compound nests and nesting trees resulting from limited and coordinated natal dispersal, with subsequent breeding site fidelity. The resulting genetic structure has implications for social behaviour in this unusual parrot species.

1. Alloparental care in cooperatively breeding species may alter breeder age-specific survival and reproduction and subsequently senescence. The helping behaviour itself might also undergo age-related change, and decisions to help in facultative cooperative breeders are likely to be affected by individual condition.\ud \ud \ud \ud 2. Helpers in long-tailed tits Aegithalos caudatus assist relatives after failing to raise their own brood, with offspring from helped nests being more likely to recruit into the breeding population.\ud \ud \ud \ud 3. Using data collected over 25 years, we examined the age trajectories of survival and reproduction in adult long-tailed tits to determine how these were affected by the presence or absence of helpers and how helper behaviour changed with age.\ud \ud \ud \ud 4. There was evidence for increased reproductive performance with breeder age, but no effect of age on the probability of survival. We found no evidence of significant senescent decline in survival or reproductive performance, although individuals accrued less inclusive fitness in their last year of life. Lifetime reproductive success was positively related to both reproductive life span and body mass. Within a season, breeders that were assisted by helpers enjoyed greater reproductive success through enhanced offspring recruitment in the following year. We found no evidence that age affected an individual's propensity to help, or the amount of indirect fitness accrued through helping.\ud \ud \ud \ud 5. We found a positive correlation between life span and multiple components of reproductive success, suggesting that individual variation in quality underpins age-related variation in fitness in this species. Helping decisions are driven by condition, and lifetime inclusive fitness of immigrants was predicted by body mass. These findings further support individual heterogeneity in quality being a major driver for fitness gains across the life course of long-tailed tits.

Animal dispersal patterns have important implications for many biological processes, but the measurement of dispersal is challenging and often requires the use of complementary approaches. In this study, we investigated the local-scale sex-biased dispersal pattern in a social bird, the black-throated tit (Aegithalos concinnus), in central China. Spatial genetic autocorrelation analyses suggested that significant fine-scale genetic structure existed in males but not in females. Mark–recapture analyses of ringed individuals also showed that female offspring were more dispersive than male offspring, supporting genetic evidence of local female-biased dispersal. These results were contrary to a previous finding of male-biased long-distance dispersal in this species that was based on analyses of gene flow across the species range in China. This implies that the species might potentially have a scale-dependent dispersal strategy, with females frequently dispersing further than males at the local level, but with a proportion of males occasionally dispersing over long distances and contributing more to gene flow at a larger geographical scale. Long-distance dispersal by male black-throated tits might be induced by competition for resources or by unfavourable environmental conditions, warranting further investigation, but our findings increase the evidence that geographical scale is an important factor to be considered when investigating animal dispersal patterns.

Riflemen/tītipounamu (Acanthisitta chloris) are kin-based cooperatively breeding birds, which appear able to recognise their relatives. Here, we investigate the potential for vocalisations to act as recognition cues in riflemen. We identified an appropriate contact call and recorded it at the nest from 19 adult riflemen. Measurements of call characteristics were individually repeatable. In addition, call similarity was significantly correlated with relatedness among all birds and among males. Thus, in principle, these contact calls contain sufficient information for individual recognition of familiar kin, and some assessment of relatedness between unfamiliar birds. To test whether riflemen responded differently to calls of kin, we broadcast calls of relatives and non-relatives as separate treatments in a playback experiment. Focal birds rarely responded aggressively or affiliatively, and their tendency to do so was unrelated to treatment. We conclude that zip calls are suitable kin recognition cues, but whether they are used as such remains unknown.

Breeding parameters for Monk Parakeets Myiopsitta monachus nesting in Barcelona, Spain, were collected for 651 nests over five breeding seasons. This invasive population has a high reproductive cap...

Inbreeding is often avoided in natural populations by passive processes such as sex-biased dispersal. But, in many social animals, opposite-sexed adult relatives are spatially clustered, generating a risk of incest and hence selection for active inbreeding avoidance. Here we show that in long-tailed tits (Aegithalos caudatus), a cooperative breeder that risks inbreeding by living alongside opposite-sex relatives, inbreeding carries fitness costs and is avoided by active kin discrimination during mate choice. First, we identified a positive association between heterozygosity and fitness, indicating that inbreeding is costly. We then compared relatedness within breeding pairs to that expected under multiple mate choice models, finding that pair relatedness is consistent with avoidance of first-order kin as partners. Finally, we show that the similarity of vocal cues offers a plausible mechanism for discrimination against first-order kin during mate choice. Long-tailed tits are known to discriminate between the calls of close kin and non-kin, and they favor first-order kin in cooperative contexts, so we conclude that long-tailed tits use the same kin discrimination rule to avoid inbreeding as they do to direct help towards kin.

Natal dispersal is a demographic trait with profound evolutionary, ecological, and behavioral consequences. However, our understanding of the adaptive value of dispersal patterns is severely hampered by the difficulty of measuring the relative fitness consequences of alternative dispersal strategies in natural populations. This is especially true in social species, in which natal philopatry allows kin selection to operate, so direct and indirect components of inclusive fitness have to be considered when evaluating selection on dispersal. Here, we use lifetime reproductive success data from a long-term study of a cooperative breeder, the long-tailed tit Aegithalos caudatus, to quantify the direct and indirect components of inclusive fitness. We show that dispersal has a negative effect on the accrual of indirect fitness, and hence inclusive fitness, by males. In contrast, the inclusive, predominantly direct, fitness of females increases with dispersal distance. We conclude that the conflicting fitness consequences of dispersal in this species result in sexually antagonistic selection on this key demographic parameter.

Kin selection is regarded as a key process in the evolution of avian cooperative breeding, and kinship influences helper decisions in many species. However, the effect of kinship on nonbreeding social organization is still poorly understood despite its potential fitness implications. Here, we investigated the origins and consequences of kin associations in nonbreeding flocks of long-tailed tits, Aegithalos caudatus, an atypical cooperative breeder where helpers are failed breeders that redirect care towards relatives living in kin neighbourhoods. We found that kinship is an important factor in initial grouping decisions; all members of a nuclear family initially joined the same flock and failed breeders chose to flock with their relatives. Flocks that merged during the nonbreeding season also contained relatives. In contrast to these findings of positive kin association, when long-tailed tits switched flocks they tended to disperse into flocks with fewer relatives, although such switches often occurred with kin. In a playback experiment, we found no evidence that aggression shown towards members of other flocks was affected by kinship, indicating that kin associations result from a preference to flock with relatives rather than a constraint on flocking with nonrelatives. Finally, using social network analysis, we show that fine-scale nonbreeding associations among individuals were positively related to kinship, and that these nonbreeding associations were reflected in helping decisions in the subsequent breeding season, in addition to the previously reported effects of kinship and proximity. We conclude that long-tailed tits prefer to associate with kin when not breeding, and suggest that by doing so they gain either nepotistic benefits within flocks or future indirect benefits during breeding.

Invasive species can have wide-ranging negative impacts, and an understanding of the process and success of invasions can be vital to determine management strategies, mitigate impacts and predict range expansions of such species. Monk parakeets (Myiopsitta monachus) and ring-necked parakeets (Psittacula krameri) are both widespread invasive species, but there has been little research into the genetic and social structure of these two species despite the potential links with invasion success. The aim of this study was to isolate novel microsatellite loci from the monk parakeet and characterise them in both monk and ring-necked parakeets in order to facilitate future investigations into their behaviour and population ecology. Sex-typing markers were also tested in both species. Of the 20 microsatellite loci assessed in 24 unrelated monk parakeets, 16 successfully amplified and were polymorphic displaying between 2 and 14 alleles (mean = 8.06). Expected heterozygosity ranged from 0.43 to 0.93 and observed heterozygosity ranged from 0.23 to 0.96. Nine of the 20 loci also successfully amplified and were polymorphic in the ring-necked parakeet, displaying between 2 and 10 alleles. Suitable markers to sex both species and a Z-linked microsatellite locus were identified. A multiplex marker set was validated for monk parakeets. These novel microsatellite loci will facilitate fine and broad-scale population genetic analyses of these two widespread invasive species.

1. Home ranging is a near-ubiquitous phenomenon in the animal kingdom. Understanding the behavioural mechanisms that give rise to observed home range patterns is thus an important general question, and mechanistic home range analysis (MHRA) provides the tools to address it. However, such analysis has hitherto been restricted to scent-marking territorial animals, so its potential breadth of application has not been tested.\ud \ud 2. Here, we apply MHRA to a population of long-tailed tits (Aegithalos caudatus), a non15 territorial passerine, in the non-breeding season where there is no clear “central place” near which birds need to remain. The aim is to uncover the principal movement mechanisms underlying observed home range formation.\ud \ud 3. Our foundational models consist of memory-mediated conspecific avoidance between flocks, combined with attraction to woodland. These are then modified to incorporate the effects of flock size and relatedness, to uncover the effect of these on the mechanisms of home range formation.\ud \ud 4. We found that a simple model of spatial avoidance, together with attraction to the central parts of woodland areas, accurately captures long-tailed tit home range patterns. Refining these models further, we show that the magnitude of spatial avoidance by a flock is negatively correlated to both the relative size of the flock (compared to its neighbour) and the relatedness of the flock with its neighbour.\ud \ud 5. Our study applies MHRA beyond the confines of scent-marking, territorial animals, so paves the way for much broader taxonomic application. These could potentially help uncover general properties underlying the emergence of animal space use patterns. This is also the first study to apply MHRA to questions of relatedness and flock size, thus broadening the potential possible applications of this suite of analytic techniques.

Natural selection should favor adoption of parental strategies that maximize fitness when allocating investment among offspring. In birds, begging displays often convey information of nestling need and quality, allowing parents to make adaptive food allocation decisions. We investigated how adults utilized cues likely to represent nestling competitive ability (begging position) and need (begging intensity) and a cue independent of nestling control (nestling sex) to distribute food among nestlings in a facultative cooperative breeder, the black-throated tit (Aegithalos concinnus). We found that parents reduced their efforts when helped, suggesting that parents of helped broods would have the potential to satisfy nestling needs more than unhelped parents. This suggestion was supported by the fact that nestling mass increased faster in helped than in unhelped nests. We found no effect of nestling sex on food allocation, but, as predicted, we found that adults responded differently to begging signals in relation to the presence of helpers and brood size. First, helped parents were more responsive to nestling begging intensity than parents without helpers. Second, female parents and helpers had a stronger preference for nestling begging position in large than in small broods. Third, the preference for nestling begging position was greater for unhelped than for helped female parents. These results provide evidence that carers adjust their preference for different offspring begging signals based on availability of food resources.

Natal dispersal is a demographic trait with profound evolutionary, ecological, and behavioral consequences. However, our understanding of the adaptive value of dispersal patterns is severely hampered by the difficulty of measuring the relative fitness consequences of alternative dispersal strategies in natural populations. This is especially true in social species, in which natal philopatry allows kin selection to operate, so direct and indirect components of inclusive fitness have to be considered when evaluating selection on dispersal. Here, we use lifetime reproductive success data from a long-term study of a cooperative breeder, the long-tailed tit Aegithalos caudatus, to quantify the direct and indirect components of inclusive fitness. We show that dispersal has a negative effect on the accrual of indirect fitness, and hence inclusive fitness, by males. In contrast, the inclusive, predominantly direct, fitness of females increases with dispersal distance. We conclude that the conflicting fitness consequences of dispersal in this species result in sexually antagonistic selection on this key demographic parameter.

In animal societies, characteristic demographic and dispersal patterns may lead to genetic structuring of populations, generating the potential for kin selection to operate. However, even in genetically structured populations, social interactions may still require kin discrimination for cooperative behaviour to be directed towards relatives. Here, we use molecular genetics and long‐term field data to investigate genetic structure in an adult population of long‐tailed tits Aegithalos caudatus, a cooperative breeder in which helping occurs within extended kin networks, and relate this to patterns of helping with respect to kinship. Spatial autocorrelation analyses reveal fine‐scale genetic structure within our population, such that related adults of either sex are spatially clustered following natal dispersal, with relatedness among nearby males higher than that among nearby females, as predicted by observations of male‐biased philopatry. This kin structure creates opportunities for failed breeders to gain indirect fitness benefits via redirected helping, but crucially, most close neighbours of failed breeders are unrelated and help is directed towards relatives more often than expected by indiscriminate helping. These findings are consistent with the effective kin discrimination mechanism known to exist in long‐tailed tits, and support models identifying kin selection as the driver of cooperation.

In species with biparental care, there is sexual conflict over parental investment because each parent benefits when their partner bears more of the reproductive costs. Such conflict can be costly for offspring, but recent theoretical work predicts that parents can resolve sexual conflict through behavioral negotiation, specifically by alternating their trips to provision nestlings. However, this idea has received almost no empirical attention. In this study, we test the hypothesis that parents alternate their delivery of food to offspring in long-tailed tits (Aegithalos caudatus) and investigate whether this coordination of parental care is associated with greater reproductive success. We show that parents alternate provisioning trips more than would be expected by chance and that parental alternation is repeatable across multiple observation periods at a nest. More alternation is associated with increased visit synchrony and increased food delivery to nestlings. Moreover, we found that nests with more alternation were less likely to be predated, probably resulting from reduced activity around the nest when parents coordinate their provisioning behavior. Our results support the hypothesis that alternation of offspring provisioning is a behavioral adaptation to reduce the costs of sexual conflict.

Dispersal is a critical driver of gene flow, with important consequences for population genetic structure, social interactions and other biological processes. Limited dispersal may result in kin-structured populations in which kin selection may operate, but it may also increase the risk of kin competition and inbreeding. Here, we use a combination of long-term field data and molecular genetics to examine dispersal patterns and their consequences for the population genetics of a highly social bird, the sociable weaver (Philetairus socius), which exhibits cooperation at various levels of sociality from nuclear family groups to its unique communal nests. Using 20 years of data, involving capture of 6508 birds and 3151 recaptures at 48 colonies, we found that both sexes exhibit philopatry and that any dispersal occurs over relatively short distances. Dispersal is female-biased, with females dispersing earlier, further, and to less closely related destination colonies than males. Genotyping data from 30 colonies showed that this pattern of dispersal is reflected by fine-scale genetic structure for both sexes, revealed by isolation by distance in terms of genetic relatedness and significant genetic variance among colonies. Both relationships were stronger among males than females. Crucially, significant relatedness extended beyond the level of the colony for both sexes. Such fine-scale population genetic structure may have played an important role in the evolution of cooperative behaviour in this species, but it may also result in a significant inbreeding risk, against which female-biased dispersal alone is unlikely to be an effective strategy. This article is protected by copyright. All rights reserved.

Cooperative breeding typically evolves within discrete, stable groups of individuals, in which group members derive direct and/or indirect fitness benefits from cooperative behaviour. In such systems, strong selection on group discrimination should emerge. Despite this prediction, relatively few studies have investigated the mechanism of group discrimination in cooperative vertebrates, and the results of many may be confounded by ‘expectancy violation’, since test individuals from which stimuli were derived remained in the group during experimentation. Here, we used a novel experimental protocol that eliminates this confounding effect in a test of group discrimination in cooperatively breeding chestnut-crowned babblers, Pomatostomus ruficeps , using long-distance contact calls. Long-distance contact calls were found to be individually specific, but to lack obvious group level signatures. Using dual playbacks of removed group and nongroup members, we found that these calls allow effective discrimination: groups unanimously approached the speaker broadcasting calls of group members and concomitantly increased their contact call rates. Together, our results suggest that group discrimination emerges from individual recognition. Additionally, the affiliative behaviour of group members towards playbacks of removed members contrasts with the aggressive responses towards nongroup members found in all other studies. One explanation for these differences stems from our elimination of expectancy violation, but further studies are required to verify this.

Productivity is a key demographic trait that can be influenced by climate change, but there are substantial gaps in our understanding of the impact of weather on productivity and recruitment in birds. Weather is known to influence reproductive success in numerous species, although such effects have not been reported in all studies, perhaps because they are masked by high nest predation rates or buffered by density dependence. Here, we use a 19-yr study of a population of individually marked long-tailed tits Aegithalos caudatus to quantify the impacts of weather on productivity in the nest (from eggs to fledging) and subsequent recruitment, while taking nest predation rates and density dependence into account. We find that weather has negligible effects on clutch size, hatching success, brood size, probability of fledging and number of fledglings. Annual variation in nest predation rates is a strong predictor of productivity, but we find no evidence that the magnitude of nest predation is determined by weather. Recruitment was strongly associated with breeding season weather, even when taking density dependence effects into account. This contrasts with the conventional view that first year survival of temperate passerines is primarily determined by winter weather. Recruitment was reduced when March temperatures were high, perhaps caused by earlier peaks in caterpillar abundance and thus reduced food availability at the time of fledging. Recruitment increased following high May temperatures, perhaps due to an improved thermo-regulatory environment for young fledglings. These opposing effects of warm March and May temperatures highlight the importance of considering asymmetrical rates of warming in different months when predicting climate change impacts.

Evolutionary theory predicts that parents should invest equally in the two sexes. If one sex is more costly, a production bias is predicted in favour of the other. Two well‐studied causes of differential costs are size dimorphism, in which the larger sex should be more costly, and sex‐biased helping in cooperative breeders, in which the more helpful sex should be less costly because future helping “repays” some of its parents’ investment. We studied a bird species in which both processes should favor production of males. Female riflemen Acanthisitta chloris are larger than males, and we documented greater provisioning effort in more female‐biased broods indicating they are likely costlier to raise. Riflemen are also cooperative breeders, and males provide more help than females. Contrary to expectations, we observed no male bias in brood sex ratios, which did not differ significantly from parity. We tested whether the lack of a population‐wide pattern was a result of facultative sex allocation by individual females, but this hypothesis was not supported either. Our results show an absence of adaptive patterns despite a clear directional hypothesis derived from theory. This appears to be associated with a suboptimal female‐biased investment ratio. We conclude that predictions of adaptive sex allocation may falter because of mechanistic constraint, unrecognized costs and benefits, or weak selection.

The repayment hypothesis predicts that reproductive females in cooperative breeding systems overproduce the helping sex. Thanks to well-documented examples of this predicted sex ratio bias, repayment has been considered an important driver of variation in sex allocation patterns. Here we test this hypothesis using data on population brood sex ratios and facultative sex allocation from 28 cooperatively breeding bird species. We find that biased sex ratios of helpers do not correlate with production biases in brood sex ratios, contrary to predictions. We also test whether females facultatively produce the helping sex in response to a deficiency of help (i.e., when they have fewer or no helpers). Although this is observed in a few species, it is not a significant trend overall, with a mean effect size close to zero. We conclude that, surprisingly, repayment does not appear to be a widespread influence on sex ratios in cooperatively breeding birds. We discuss possible explanations for our results and encourage further examination of the repayment model.

Kin selection has played an important role in the evolution and maintenance of cooperative breeding behaviour in many bird species. However, although relatedness has been shown to affect the investment decisions of helpers in such systems, less is known about the role that kin discrimination plays in other contexts, such as communal roosting. Individuals that roost communally benefit from reduced overnight heat loss, but the exact benefit derived depends on an individual's position in the roost which in turn is likely to be influenced by its position in its flock's dominance hierarchy. We studied the effects of kinship and other factors (sex, age, body size and flock sex ratio) on an individual's roosting position and dominance status in captive flocks of cooperatively breeding long-tailed tits Aegithalos caudatus. We found that overall, kinship had little influence on either variable tested; kinship had no effect on a bird's position in its flock's dominance hierarchy and the effect of kinship on roosting position was dependent on the bird's size. Males were generally dominant over females and birds were more likely to occupy preferred roosting positions if they were male, old and of high status. In this context, the effect of kinship on social interactions appears to be less important than the effects of other factors, possibly due to the complex kin structure of winter flocks compared to breeding groups.

Successful urban colonization by formerly rural species represents an ideal situation in which to study adaptation to novel environments. We address this issue using candidate genes for behavioural traits that are expected to play a role in such colonization events. We identified and genotyped 16 polymorphisms in candidate genes for circadian rhythms, harm avoidance and migratory and exploratory behaviour in 12 paired urban and rural populations of the blackbird Turdus merula across the Western Palaearctic. An exonic microsatellite in the SERT gene, a candidate gene for harm avoidance behaviour, exhibited a highly significant association with habitat type in an analysis conducted across all populations. Genetic divergence at this locus was consistent in 10 of the 12 population pairs; this contrasts with previously reported stochastic genetic divergence between these populations at random markers. Our results indicate that behavioural traits related to harm avoidance and associated with the SERT polymorphism experience selection pressures during most blackbird urbanization events. These events thus appear to be influenced by homogeneous adaptive processes in addition to previously reported demographic founder events.

Structures built by animals, such as nests, mounds and burrows, are often the product of cooperative investment by more than one individual. Such structures may be viewed as a public good, since all individuals that occupy them share the benefits they provide. However, access to the benefits generated by the structure may vary among individuals and is likely to be an important determinant of social organisation. Here we use the massive, communal nests of sociable weavers Philetairus socius, to investigate whether their thermoregulatory function varies in relation to the size of communal nests, and the position of individual nest chambers within the communal structure. We then examine whether this spatial variation in thermoregulatory function predicts the social organisation of colonies. First, we show that the sociable weavers’ communal nests buffer variation in ambient temperature, and reduce temperature variability within nest chambers. The extent of this buffering effect depends significantly on the position of nest chambers within the communal structure, and on the depth to which chambers are embedded within the nest mass. We detected no effect of nest volume on thermoregulatory benefits, suggesting that there are likely to be additional, non-thermoregulatory benefits leading to communal nests. Finally, our results indicate that there may be competition for access to the benefits of the public good, since older birds occupied the chambers with the highest thermoregulatory benefits, where breeding activity was also more common. We discuss how the spatial structure of the benefits of the public good might influence social organisation in the unique communal lifestyle of sociable weavers.

Thirty-seven unique polymorphic microsatellite loci were characterised in 24 unrelated rifleman individuals (Acanthisitta chloris) sampled from a population located at Kowhai Bush, Kaikoura, New Zealand. Thirty of these loci were newly isolated from a rifleman genomic library enriched for microsatellite motifs, and seven were identified from the testing of markers of known high-cross species utility. Sequence homology to the assembled zebra finch (Taeniopygia guttata) genome was successfully used to predict the chromosome locations for 33 of these polymorphic loci. Three of the loci were found to be Z-linked based a lack of heterozygotes in females and sequence homology. These loci will be of utility for population and parentage studies in the rifleman. A proportion of these loci are expected be useful in the related species, such as the endangered rock wren (Xenicus gilviventris).

There is large interspecific variation in the magnitude of population fluctuations, even among closely related species. The factors generating this variation are not well understood, primarily because of the challenges of separating the relative impact of variation in population size from fluctuations in the environment. Here, we show using demographic data from 13 bird populations that magnitudes of fluctuations in population size are mainly driven by stochastic fluctuations in the environment. Regulation towards an equilibrium population size occurs through density-dependent mortality. At small population sizes, population dynamics are primarily driven by environment-driven variation in recruitment, whereas close to the carrying capacity K, variation in population growth is more strongly influenced by density-dependent mortality of both juveniles and adults. Our results provide evidence for the hypothesis proposed by Lack that population fluctuations in birds arise from temporal variation in the difference between density-independent recruitment and density-dependent mortality during the non-breeding season., Why does population variability differ? Sæther et al. show that the magnitude of temporal variation of avian population fluctuations is mainly determined by environmental fluctuations affecting recruitment, whereas regulation of mean population size occurs through density-dependent mortality.

The population of common guillemots Uria aalge on Skomer Island, Wales has been monitored since 1963, and in the last 30 yr has increased at an almost constant rate of 5% yr−1. A previous attempt to model the population based on intrinsic demographic parameters estimated over just five years failed to explain the observed population increase, probably because the estimate of juvenile survival was too low. This raised the possibility that immigration fuelled the population increase. Here we use > 30 yr of detailed field observations to re-estimate key population parameters (productivity, adult survival and juvenile survival) in order to model the population. We show that the observed rate of increase can be explained by these intrinsic parameters, and we therefore conclude that immigration is not necessary to generate the observed population growth.

How individuals colonising novel environments overcome the diverse suite of new selection pressures is a fundamental question in ecology and evolution. Urban environments differ markedly from the rural ones that they replace and successful colonisation of urban areas may therefore require local adaptation and phenotypic/genetic divergence from ancestral populations. Such a process would be facilitated by limited dispersal to and from the novel habitat. Here we assess divergence in migratory behaviour between seven pairs of urban and rural European blackbird Turdus merula populations along a 2800 km transect across Europe. This former forest specialist is now amongst the most abundant urban birds across most of its range. We use a stable isotope approach due to the lack of sufficient ringing data from multiple urban populations, and compare hydrogen isotopic ratios of tissues grown in the breeding (feathers) and wintering areas (claws) to derive an index of long distance migratory behaviour. We find a tendency for urban blackbirds to be more sedentary than rural ones at all sites and this divergence is particularly strong at the north-eastern limit of our transect, i.e. in Estonia and Latvia. These urban populations are those that have been established most recently (from the late 1930s to 1950s) implying that urbanisation can promote rapid ecological divergence. The increased sedentary behaviour of urban birds could promote further ecological divergence between rural and urban populations, such as the earlier breeding of urban blackbirds, and in some cases may contribute to their previously documented genetic divergence.

A growing number of bird species are known to have fine‐scale genetic structure during the breeding season, with relatives breeding in close vicinity. Such genetic structure often has fitness consequences for parents, and sex ratio theory predicts that females should respond adaptively when they determine offspring sex. We examined whether or not females allocate offspring sex adaptively in response to the local genetic structures as well as other biotic and abiotic factors in a population of the vinous‐throated parrotbill Paradoxornis webbianus, a small passerine with strong flocking habit and various genetic structures among neighbouring males during the breeding season. The average brood sex ratio of hatchlings (secondary sex ratio) did not deviate from parity. In addition, the observed brood sex ratio was independent of the fine‐scale genetic structure and other factors including breeding density, clutch size, laying date, parents’ quality, and the presence of extrapair paternity. Accordingly, we reject the hypothesis of adaptive sex allocation by female parrotbills in association with local genetic structure and other factors. Instead we conclude that despite the plausible benefits of biased sex allocation, this species determines brood sex ratio via random sex allocation with equal probability of male and female offspring.

In cooperatively breeding vertebrates, investment strategies of breeders and helpers may be affected by the sex of offspring that they provision because the fitness benefits gained from caring for sons or daughters will depend upon the fitness impacts of positive (i.e. local resource enhancement) and negative (i.e. local resource competition) social interactions between relatives. We investigated the relationship between provisioning effort by breeders and helpers and the sex of nestlings that they provision, using long-term data over 16 years and detailed behavioural observation of interactions between carers and offspring in a cooperatively breeding species, the long-tailed tit, Aegithalos caudatus . We predicted preferential care for male nestlings because males are the more helpful sex in the kin-selected social system of long-tailed tits. However, we found that despite the significant size difference between male and female nestlings, there was no evidence of facultative adjustment of provisioning effort by either breeders or helpers in relation to brood sex ratio. In addition, we found no evidence for sex-biased provisioning by carers, and no bias in the feeds received by nestlings according to their sex. Thus, we conclude that the provisioning strategy of long-tailed tits does not incorporate differential investment according to sex. We suggest that the unpredictable nature of helping in the atypical cooperative breeding system of long-tailed tits makes random provisioning effort in relation to nestling sex the best option for carers.

In animal societies, kin selection is a critical evolutionary process, with cooperation evolving principally among relatives living in kin-structured populations. Theoretical and empirical studies have largely focused on population viscosity--the timing or distance of dispersal--as the key factor generating kin structure. This is despite extensive theoretical broadening of the factors and processes influencing effective population size, variance in reproduction, and relatedness. Here, we explore predation mortality as a specific driver of population-level reproductive skew and variance in fecundity to show how a common and perhaps underappreciated event in organism life history can give rise to patterns of relatedness. We develop our case study around an empirically derived model where elevated relatedness arises from predation that alters the timing and nature of offspring mortality, essentially driving variance in fecundity. This leads to dramatic changes in the emergent kin structure of the surviving breeding population. Our in-silico experiments recover the theoretical predictions that when predation acts on clusters of individuals and effectively removes whole family groups (i.e., broods), rather than individuals, from the pool of potential recruits, there is a greater kin structure in the emergent adult population. We conclude that empirical attempts to understand the factors promoting kin-structured populations and the evolution of sociality should now match theoretical efforts to be more inclusive of ecological process generating life history and demographic variability. Copyright 2011, Oxford University Press.

Cooperative breeding has evolved primarily in species in which individuals are organized into family groups, and kin selection is considered to be a major force in the evolution of helping behaviour. Family groups are generally thought to form through delayed or limited dispersal, but dispersal patterns vary considerably both between species and in different populations of the same species, and the relationship between dispersal, kinship and cooperation is poorly understood. In this study, we combined long-term observational and genetic data to compare the patterns of demography, kinship and helping in two populations of long-tailed tits, Aegithalos caudatus, a species that exhibits kin-biased helping by failed breeders but not delayed dispersal. Both populations had the same annual breeding success, but philopatric recruitment rates were significantly higher in one population, especially for females. This led to a correspondingly higher proportion of individuals having at least one close relative in that population. Surprisingly, however, there was no difference in the pattern of helping behaviour between the two populations in terms of helper prevalence, helper sex or the relatedness between helpers and the breeders they assisted. We discuss possible explanations and highlight the limitations of population-level analyses for understanding the relationship between demography and cooperation.

In cooperatively breeding species, the fitness consequences of producing sons or daughters depend upon the fitness impacts of positive (repayment hypothesis) and negative (local competition hypothesis) social interactions among relatives. In this study, we examine brood sex allocation in relation to the predictions of both the repayment and the local competition hypotheses in the cooperatively breeding long-tailed tit Aegithalos caudatus. At the population level, we found that annual brood sex ratio was negatively related to the number of male survivors across years, as predicted by the local competition hypothesis. At an individual level, in contrast to predictions of the repayment hypothesis, there was no evidence for facultative control of brood sex ratio. However, immigrant females produced a greater proportion of sons than resident females, a result consistent with both hypotheses. We conclude that female long-tailed tits make adaptive decisions about brood sex allocation.

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 x 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.

Dispersal is a key factor affecting the dynamics and genetic structure of populations. Patterns of natal dispersal vary widely depending on the selection pressures that each species encounters, but the underlying factors determining this variation are poorly understood. We studied the pattern of natal dispersal in a small passerine, the vinous-throated parrotbill, Paradoxornis webbianus, using both direct ringing data and indirect genetic methods. The social structure of this species is characterized by group living without apparent territorial behaviour. The dispersal rate of juveniles from their natal groups was moderate (about 40%) and dispersal distance was short, most individuals moving to neighbouring flocks. Sibling groups that included both sexes usually dispersed or were philopatric together so that they were found in the same winter flock where pairing occurs. Thus, the avoidance of inbreeding or intrasexual competition among siblings appears not to affect natal dispersal in this species. Instead, our results suggest that the benefits of dispersing in groups and/or selection of kin structure within social groups via kin-selected reproductive benefits may promote the observed dispersal pattern: sibling associations without separation of the sexes. This behaviour may have important implications for the kin structure of populations and the evolution of sociality.

Previous studies have shown that a female dunnock Prunella modularis increases her reproductive success on average by copulating with more than one male resident on her territory and thereby obtaining extra help in raising offspring. Here we document behavior by females that affects which males copulate with them. During her period of receptivity to copulation, a female in a territory shared by two males often left the dominant (or alpha) male, which guarded her most of the time, and approached the subordinate (or beta) male when he sang. A female's responses to individual males thus tend to increase her own reproductive success by increasing her chances for copulation with both males sharing her territory. Playbacks of tape-recorded songs in the field showed that females approached only songs of resident males, not neighbors. They can therefore discriminate individual males by their songs alone, a capability not previously established for female songbirds. Despite intensive guarding of females by males, mating success among male dunnocks depends in part on female choice.

Animal societies of varying complexity have been the favoured testing ground for inclusive fitness theory, and there is now abundant evidence that kin selection has played a critical role in the evolution of cooperative behaviour. One of the key theoretical and empirical findings underlying this conclusion is that cooperative systems have a degree of kin structure, often the product of delayed dispersal, that facilitates interactions with relatives. However, recent population genetic studies have revealed that many non-cooperative animals also have kin-structured populations, providing more cryptic opportunities for kin selection to operate. In this article, I first review the evidence that kin structure is widespread among non-cooperative vertebrates, and then consider the various contexts in which kin selection may occur in such taxa, including: leks, brood parasitism, creches, breeding associations, territoriality and population dynamics, foraging and predator deterrence. I describe the evidence that kin-selected benefits arise from interacting with kin in each of these contexts, notwithstanding the potential costs of kin competition and inbreeding. I conclude that as the tools required to determine population genetic structure are readily available, measurement of kin structure and the potential for kin selection on a routine basis is likely to reveal that this process has been an important driver of evolutionary adaptation in many non-cooperative as well as cooperative species.

Helpers in cooperatively breeding species may gain direct fitness benefits that increase their survival probability and/or reproductive success. However, survival and productivity may be influenced by many other factors, including variation in dispersal, nepotistic interactions, or individual condition. High helper survival relative to nonhelpers has been reported in the long-tailed tit Aegithalos caudatus, a cooperative breeder where helpers are failed breeders that redirect care toward kin. Using capturemark-recapture analysis of a long-term data set, we confirm this result and show that it is not attributable to differential dispersal. Then, using only males with first-order relatives (to control for any effects of nepotism), we investigated the survival of 3 groups of failed breeders (i.e., potential helpers): survival of helpers was highest (61%) but, contrary to predictions based on direct benefits, survival of 2 categories of nonhelpers differed; those with helping opportunities had the lowest survival rate (24%), whereas those without helping opportunities had intermediate survival (52%). We suggest that the groups varied in condition; helpers are in good condition, males with helping opportunities who did not help are in poor condition, and nonhelpers without helping opportunities comprise a mixture of birds in good and poor condition. This conclusion was supported by differences in the timing of breeding (a proxy for condition) between groups: helpers bred earliest and nonhelpers with helping opportunities bred latest. Furthermore, we found no evidence that helpers gained any future reproductive benefits. We suggest that condition rather than benefits accrued as a direct result of helping influenced helper survival. Key words: condition, direct fitness, helper, long-tailed tit, survival. [Behav Ecol]

The amount of food delivered by parents to their chicks is affected by various life history traits as well as environmental and social factors, and this investment ultimately determines the current and future fitness of parents and their offspring. We studied parental provisioning behaviour in the Vinous-throated Parrotbill Paradoxornis webbianus, a species with an unusual social system that is characterised by flock-living, weak territoriality and variable nesting dispersion. Parental provisioning rate had a positive influence on chick mass gain, suggesting that provisioning rate is an effective measure of parental investment in this species. Males and females fed nestlings at approximately the same rate, and no other carers were observed at nests. Parents coordinated provisioning rates so that they mostly fed chicks synchronously. However, the extent to which parents coordinated provisioning was associated with their social environment, synchrony being positively related to local breeding density and negatively to nearest neighbour distance. The rate at which parents provisioned nestlings showed the same relationships with social measures, being greatest at higher density and when neighbours were closer. Visit rate was also related to chick age, but not to brood size, brood sex ratio, extra pair paternity, laying date, temperature, parents’ body characters, time of day or year. We conclude that a breeding pairs’ social environment plays an important role in determining parental investment, probably through its effects on the opportunities that parents have for foraging with conspecifics.

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 non-social 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.

An emerging consequence of global climate change is its potential effect on the timing of seasonal biological events. Analysis of long-term datasets reveals a high degree of plasticity in the nature of phenological responses both within and among species, and understanding these differ- ences is central to understanding the mechanisms and implications of climate-related change. We investigated factors influencing timing of breeding (median laying date) in a colonial nesting seabird, the common guillemot Uria aalge, over 23 breeding seasons between 1973 and 2008. There was a trend for earlier laying over this period, and earlier laying was associated with higher average breed- ing success. Multiple regression models (with de-trended explanatory variables to control for linear trends over time) indicate that the timing of breeding is positively correlated with a wide-scale cli- matic driver, the winter North Atlantic Oscillation (WNAO), and negatively correlated with popula- tion size: guillemots lay later in years with high WNAO indices and earlier in years with larger pop- ulations. Responses to environmental conditions are probably related to indirect effects on timing or abundance of food availability, direct effects of weather or both. The mechanism(s) leading to a pos- sible relationship between laying date and population size are less clear. They may be related to Allee-type effects associated with social stimulation, improved foraging efficiency or a density- dependent increase in breeding site quality. Given the correlative nature of these results, we are cau- tious about the role of non-climatic (intrinsic) factors, but we cannot exclude that they play a role alongside climatic (extrinsic) factors in influencing reproductive phenology.

The adaptive significance of flocking behaviour has been intensively studied, especially among birds, but the demography and genetic structure of winter flocks is poorly understood, despite their importance for many biological processes. For three successive winters, we studied the demography and genetic structure of winter flocks in a small passerine, the vinous-throated parrotbill Paradoxornis webbianus. Our objectives were to determine the match between observed demography and the genetic structure of winter flocks, and to investigate the consequences of kin structure for the risk of inbreeding during the breeding season. The size of five main study flocks ranged from 60 to 120 individuals. The sex ratio of each flock did not deviate from parity and survival rates of adults were moderately low (36 and 17% in 2 years). Adults showed strong fidelity to a winter flock not only within a season but also between years. As expected from observational data, we found significant genetic differentiation and isolation by distance among males and females in winter flocks at a fine spatial scale. Likewise, relatedness among individuals within a winter flock was significantly greater than between flocks. Despite these demographic and genetic circumstances, the inbreeding rate was relatively low (4%). We conclude that fine-scale genetic structure occurs in the winter flocks of birds as a result of demographic processes without physical barriers. This genetic structure introduced a risk of inbreeding, but our evidence on whether individuals avoided incest was equivocal.