Population Of Blue Tits Research Articles

The Genetic Response of Forest Birds to Urbanization: Variability in the Populations of Great and Blue Tits

The Genetic Response of Forest Birds to Urbanization: Variability in the Populations of Great and Blue Tits

Fine-scale genetic structure and phenotypic divergence of a passerine bird population inhabiting a continuous Mediterranean woodland.

Genetic differentiation between populations inhabiting ecologically different habitats might appear because of limited dispersal and gene flow, which may lead to patterns of phenotypic divergence and local adaptation. In this study, we use dispersal, genotypic (24 microsatellite loci) and phenotypic (body size and clutch size) data to analyse patterns of genetic structuring and phenotypic divergence in a blue tit (Cyanistes caeruleus) population inhabiting a continuous and heterogeneous woodland along a valley. The two slopes of the valley differ in their forest formations and environmental conditions. Findings showed that most blue tits reproduced within their natal slope. Accordingly, microsatellite analyses revealed that populations of blue tits established in the two slopes show subtle genetic differentiation. The two genetic populations diverged in clutch size, exceeding the level of differentiation expected based on genetic drift, hence suggesting divergent selection (or other processes promoting divergence) on this life-history trait. Our findings reveal that restricted dispersal and spatial heterogeneity may lead to genetic differentiation among bird populations at a surprisingly small scale. In this respect, it is worth highlighting that such differentiation occurs for an organism with high dispersal capacity and within a continuous woodland. Moreover, we show that small-scale ecological differences, together with limited gene flow, can result in selection favouring different phenotypes even within the same continuum population.

Removal of older males increases extra-pair siring success of yearling males.

In animals, reproductive performance typically improves over time early in life. Several ultimate and proximate mechanisms may contribute to such an age-related improvement and these mechanisms can act in a relative or in an absolute sense. Low performance of young individuals may be the consequence of a comparison or competition with older individuals (relative), or it may be due to specific traits of young individuals and be unrelated to the presence of older competitors (absolute). Here, we perform a test to disentangle whether the effect of age class (yearling or older) on male extra-pair siring success is relative or absolute. Male age is the most consistent predictor of male extra-pair siring success across bird species, yet the mechanisms underlying this pattern are not well understood. Low extra-pair siring success of yearling males may be a consequence of the presence of older ("adult") males (hypothesis 1), because adult males are more successful in intra- and intersexual interactions or because females prefer to copulate with adult males when available (relative preference). Alternatively, low extra-pair siring success of yearlings may be independent of the presence of adult males (hypothesis 2), for example, if yearling males on average invest less in extra-pair behavior or if females avoid them as extra-pair mates, independent of the availability of older males (absolute preference). To distinguish between these 2 hypotheses, we experimentally manipulated the age structure of a nest-box-breeding population of blue tits (Cyanistes caeruleus) by removing almost all adult males, and compared patterns of extra-pair paternity in the experimental year with those from the preceding 15 "control" years. Removal of adult males resulted in a substantial increase in the extra-pair siring success of yearling males compared to the "control" years, but did not affect the population-level frequency of extra-pair paternity or its spatial patterns. Our results provide clear evidence that extra-pair siring success of yearlings can increase and that it depends on the presence of older males in the population, indicating a relative effect of age on reproductive performance. These results suggest that older males outcompete yearling males in direct or indirect interactions, in sperm competition or as a result of differences in attractiveness to females.

Begging calls and mouth colouration as predictors of breeding success in blue tits

AbstractNestlings normally exhibit a mix of traits that attract parental care, such as postural and vocal begging and carotenoid‐based mouth colouration. These signals are hypothesised to be signs of nestling needs (vocal begging) and quality (mouth colouration). Therefore, we hypothesised that broods, where nestlings beg for less time and display more saturated carotenoid‐based mouth colouration, would have lower nestling mortality. We tested these predictions in two wild blue tit (Cyanistes caeruleus) populations. The breeding success (here defined as the proportion of eggs that produced fledglings) was related to nestling mouth flanges' carotenoid‐based colour saturation. This means that blue tits that raised nestlings with more coloured flanges had a higher within‐population breeding success. Time spent in vocal begging, by contrast, was not associated with breeding success. Hence, our findings reveal that some communication signals displayed by nestlings (carotenoid‐based colouration) predict breeding success, in our study mostly reflecting the proportion of eggs that hatched, while begging intensity does not, probably because the former reflects a better maternal pre‐laying condition and nestling physiological status in the mid‐term.

Genotyping blastoderms of avian eggs.

Undeveloped eggs occur frequently in birds and are often considered infertile, and discarded. However, the majority of undeveloped eggs may in fact have been fertilised and embryos might have died at an early stage. Such eggs contain valuable information, for example about offspring sex and paternity, and level of inbreeding. Obtaining such information may also give insight into the patterns and causes of early embryo mortality. Here we describe a simple technique for removing embryo cells from the blastoderm to obtain DNA to genotype the offspring and unequivocally ascertain fertilisation status, while retaining the overlying perivitelline layer (PVL) for sperm counts over the entire membrane. We tested this method on freshly collected eggs (high-quality material), as well as on eggs from abandoned clutches and unhatched eggs (potentially deteriorated material) of blue tits (Cyanistes caeruleus). We sampled a total of 707 eggs from a wild population of blue tits, extracted DNA from the eggs' blastoderm using a Qiagen kit, and genotyped the samples with 14 polymorphic microsatellite markers, plus one sexing marker. Overall, we successfully genotyped 97% of all eggs. Our study is the most extensive dataset of genotyped undeveloped eggs to date and demonstrates that one can reliably genotype undeveloped fertile eggs as well as retain the PVL for observations of sperm and embryo cells.

Is there a role for aromatic plants in blue tit (Cyanistes caeruleus) nests? Results from a correlational and an experimental study

The utility of fresh green material in avian nests is still not fully understood. Potential explanations include the effects of plants’ volatile compounds on parasite reduction (nest protection hypothesis) or direct beneficial effects on nestling condition (drug hypothesis). We used correlative data collected during 2020 and 2021 in a Mediterranean population of blue tits (Cyanistes caeruleus) as well as experimental data (aromatic nest content manipulation) to assess the effects of aromatic plant use on nestling physiological condition and survival, nest-dwelling ectoparasitic pressure and its relationship with breeding parameters. We found that aromatic plants were disproportionally used in relation to their abundance in the environment and that their use was positively related to egg mass (but only in 2020). Blowflies and facultative parasitic mites were more frequent in nests with aromatics compared to nests without aromatics, but obligatory parasitic mites were less abundant in nests with aromatics. However, no effects of aromatic plants were observed on nestling haemoglobin levels nor erythrocyte sedimentation rate or other physiological health metrics, but the heterophil to lymphocyte ratio was higher in nests with the highest quantity of aromatics. The artificial addition of mint reduced the flea abundance, but 7-day old nestlings showed significantly lower colour saturation and brightness in the mouth flange. Nestling survival to fledging was not related to aromatic plant use. Therefore, our results partially support a beneficial effect of aromatic plants in blue tit nests because some ectoparasite groups were reduced. Immediate effects on nestling physiology or survival could not be established.Significance statementSome avian species place fresh aromatic plant material in their nests, and several non-mutually exclusive hypotheses have been proposed to explain its potential functions. In this study, we use both correlational and experimental data from a blue tit population to test two hypotheses, namely the ‘nest protection hypothesis’ and the ‘drug hypothesis’. The first one proposes that aromatic plants have direct repellent effects against ectoparasites, while the latter poses that these plant components benefit nestling condition through the stimulation of some components of the immune system. Our results suggest that some ectoparasites, such as fleas and obligatory parasitic mites, were less abundant in nests where aromatic plants were artificially or naturally added, respectively, but no relationships were detected with nestling physiology or survival, which could be partially explained by our small sample size. Our study partially supports a beneficial effect of aromatic plants in avian nests.

Untangling behaviours: independent expressions of female–female aggression and snake-like hissing in the blue tit (Cyanistes caeruleus)

Aggression plays a crucial role in deterring predators and securing resources to promote fitness. Nevertheless, studies focussing on female aggression remain scarce. In songbirds, aggression is prevalent during the breeding season, when same-sex individuals compete for limited resources. Additionally, females of some bird species exhibit snake-like hissing behaviour during incubation presumably to lower predation rates and improve fitness. Such behaviours may co-vary, forming a behavioural syndrome that could constrain trait expression. Here, we investigated a resident population of blue tits (Cyanistes caeruleus), to examine the repeatability and covariation of female–female aggression and hissing behaviour, aiming to determine if these constitute a behavioural syndrome. We quantified female–female aggression during simulated territorial intrusions and measured number of hissing calls in response to a simulated predator intrusion into the nest box. We found that both female–female aggression and hissing behaviour were repeatable traits, and that older females approached the intruder less. However, we found no evidence of covariation between female–female aggression and hissing behaviour. Thus, our findings suggest that female–female aggression and hissing behaviour, although both displayed in a nest defence context, are evolutionarily independent traits in the blue tit.

Age trajectories in extra-pair siring success suggest an effect of maturation or early-life experience.

Across birds, male age is the most consistent predictor of extra-pair siring success, yet little is known about age effects on paternity over the lifetime of individuals. Here, we use data from a 13-year study of a population of blue tits (Cyanistes caeruleus) to investigate how extra-pair siring success changes with age within individuals. Our results indicate that extra-pair siring success does not continuously increase with male age. Instead, siring success was related to male age in a threshold fashion, whereby yearling males were less likely to gain paternity than older males. This effect was independent of the age of the social partner, but influenced by the age of the extra-pair female: success of yearlings at siring extra-pair young (EPY) with older females was even lower. Among males that sired at least one EPY, the number of extra-pair mates and the proportion of EPY sired were unrelated to male age. We found no evidence for an influence of selective disappearance on extra-pair reproduction. Senescence, if anything, only occurs at ages blue tits rarely reach. A literature review indicates that an effect of male age on extra-pair siring success may be limited to the switch from yearling to older in many species. Thus, the generally observed age effect on male extra-pair siring success may be linked to age class rather than continuous ageing. This suggests that lack of experience or not fully completed maturation are important drivers of age patterns in extra-pair paternity.

Prevalence, Molecular Characterization, and Ecological Associations of Filarioid Helminths in a Wild Population of Blue Tits (Cyanistes caeruleus)

Filarioid nematodes (commonly known as filarial worms) are known to impact human and domestic animal health, but studies examining their ecological relevance and impacts on wildlife are still underrepresented. In the case of birds, microfilariae are typically found at low prevalence, but they may negatively affect some fitness-related traits. Here, we study the prevalence and associations of microfilariae in a wild population of blue tits (Cyanistes caeruleus) inhabiting a woodland comprising different forestry formations. In addition, we characterize the filarioid lineages through the cytochrome c oxidase subunit I (COI) gene sequence. We found a moderate prevalence of microfilariae in the blue tit population (9.4%) and that the presence of such parasites was negatively associated with host body mass. Neither forest type nor host sex influenced microfilariae presence. Phylogenetic analyses revealed the presence of five filarioid lineages clustered in the Onchocercidae family—four out of five lineages clustered in the Splendidofilaria clade, while the remaining lineage could not be clearly assigned to a genus. In addition, this is the first study examining the filarioid lineages infecting the blue tit. Our results suggest that hosts in poorer body condition, in terms of lower body mass, are more susceptible to be parasitized by filarioid nematodes and call for further genetic studies of these parasites.

Habitat-dependent breeding biology of the Blue Tit (Cyanistes caeruleus) across a continuous and heterogeneous Mediterranean woodland

Mediterranean woodland environments are characterised by high spatial and temporal heterogeneity, which means the inhabiting species face a wide variety of selective pressures. Species may respond differently to habitat heterogeneity and so distinct eco-evolutionary scenarios may be responsible for the inter-habitat variability in reproductive strategies observed in certain species. The inter-forest variability of some reproductive traits in passerines has been examined by comparing forest patches or separated fragments. However, there is still little information regarding how such highly mobile animals adjust their breeding performance across continuous and heterogeneous woodlands. Here we studied the reproductive performance of a population of Blue Tits (Cyanistes caeruleus) in an area of continuous Mediterranean woodland that included two mountain slopes and four different types of forest, ranging from deciduous oak forests to perennial non-oak forests. We studied the habitat heterogeneity and inter-forest phenotypic variation in terms of reproductive performance and adult and nestling biometry, besides also exploring the effects of ectoparasites on Blue Tit reproduction. Eggs were laid earliest in deciduous Pyrenean Oak (Quercus pyrenaica) forests, while clutch size and the number of fledglings were highest in the humid Pyrenean Oak forest, which had the greatest tree coverage and most humid climate, and lowest in the coniferous Scots Pine (Pinus sylvestris) forest. There were no inter-forest differences in hatching (percentage of nests with at least one egg hatched) and fledging (percentage of nests in which at least one nestling fledged) success. Similarly, there were no inter-forest differences in adult and nestling biometry, but adults that raised more fledglings had a lower body mass, while males whose females laid larger clutches had smaller tarsi. Most ectoparasites did not affect Blue Tit reproduction, although Culicoides had a negative impact on nestling body mass. These results suggest that Blue Tits can adjust their reproductive effort to the forest where they breed even across a very small spatial scale. Different eco-evolutionary scenarios, such as phenotypic plasticity or genetic structuring and local adaptation, might explain the phenotypic differentiation in the reproductive strategies observed over small areas in woodlands.

Patterns of extra-territorial nest-box visits in a songbird suggest a role in extrapair mating.

Many animals make visits outside of their territory during the breeding period, but these are typically infrequent and difficult to observe. As a consequence, comprehensive data on extra-territorial movements at the population-level are scarce and the function of this behavior remains poorly understood. Using an automated nest-box visit tracking system in a wild blue tit population over six breeding seasons, we recorded all extra-territorial nest-box visits (n = 22 137) related to 1195 individual breeding attempts (761 unique individuals). Sixty-two percent of breeders made at least one extra-territorial visit between the onset of nest building and the day of fledging of their offspring, and individuals visited another nest-box on average on 11% of the days during this period. Visit behavior differed clearly between the sexes, with males making over three times as many extra-territorial forays as females. There was a strong overall seasonal decline in visit behavior, but this was sex dependent, with females showing a strong reduction in the number of extra-territorial visits before the onset of egg laying and males showing a strong and sudden reduction on the day their offspring hatched. The likelihood of visiting a particular nest-box declined sharply with the distance to that box, and blue tits almost exclusively visited direct neighbors. Individuals were more likely to have extrapair offspring with an individual whose box they visited, but they were not more likely to disperse to a box they had visited. Thus, our results are inconsistent with the hypothesis that extra-territorial nest-box visits serve to inform dispersal decisions, but suggest that such visits are linked to extrapair mating opportunities.

Temperature synchronizes temporal variation in laying dates across European hole-nesting passerines.

Identifying the environmental drivers of variation in fitness-related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness-related trait values (i.e., correlated temporal trait fluctuations across populations) is poorly understood. Using data from long-term monitored populations of blue tits (Cyanistes caeruleus, n = 31), great tits (Parus major, n = 35), and pied flycatchers (Ficedula hypoleuca, n = 20) across Europe, we assessed the influence of two local climatic variables (mean temperature and mean precipitation in February-May) on spatial synchrony in three fitness-related traits: laying date, clutch size, and fledgling number. We found a high degree of spatial synchrony in laying date but a lower degree in clutch size and fledgling number for each species. Temperature strongly influenced spatial synchrony in laying date for resident blue tits and great tits but not for migratory pied flycatchers. This is a relevant finding in the context of environmental impacts on populations because spatial synchrony in fitness-related trait values among populations may influence fluctuations in vital rates or population abundances. If environmentally induced spatial synchrony in fitness-related traits increases the spatial synchrony in vital rates or population abundances, this will ultimately increase the risk of extinction for populations and species. Assessing how environmental conditions influence spatiotemporal variation in trait values improves our mechanistic understanding of environmental impacts on populations.

Exploration behaviour and foraging strategies in Mediterranean blue tits

Animal behaviour is potentially a mechanism of individual diet specialization. To explore this possibility we assessed exploratory behaviour (EB) and foraging data for a population of blue tits Cyanistes caeruleus. Our results suggest that: 1) foraging differs between sexes; 2) the prey type selected by females depends on the provisioning rates of their mate, and adjustment of this prey-choice differs between EB phenotypes; and 3) foraging behaviour in males shows a trend towards linkage to their EB phenotype, with faster-exploring males obtaining marginally larger caterpillars but provisioning less frequently than slower-exploring males. Lastly, environmental variables contributed substantially to the diet composition of offspring. For this reason, we cannot suggest that EB dominates, but it may contribute to a diet specialization process in our study population.

Within-brood body size and immunological differences in Blue Tit (Cyanistes caeruleus) nestlings relative to ectoparasitism

Several ectoparasites parasitise nestlings decreasing their body condition, growth and survival. To minimise any loss of fitness due to ectoparasites, birds have developed a wide variety of defence mechanisms, potentially including hatching asynchrony. According to the Tasty Chick Hypothesis (TCH), the cost of parasitism would be reduced if ectoparasites tend to eat on less immunocompetent nestlings, typically the last-hatched chick in asynchronously hatched broods, as they are in poor body condition. Two predictions of the TCH are that immune capacity is lower in smaller nestlings than in larger ones and that parasites should provoke a more negative effect on smaller nestlings. Here, we test these predictions in a population of Blue Tits (Cyanistes caeruleus) whose broods are parasitised by Hen Fleas (Ceratophyllus gallinae) and Blowflies (Protocalliphora azurea). We recorded the presence of both ectoparasites and analysed the immunocompetence (number of leucocytes per 10,000 erythrocytes and cutaneous immune response to phytohaemagglutinin) and body condition of smaller and larger nestlings within individual broods. The leucocyte count was higher in smaller nestlings than in larger ones, whereas the cutaneous immune response did not differ between smaller and larger nestlings. Smaller nestlings, but not larger nestlings, had lower body mass when fleas were present. Blowflies, by contrast, had no detectable negative effect on nestlings. Overall, our findings provide partial support to the TCH. Lower immune capacity in smaller nestlings than in larger ones was not supported, but Hen Fleas seemed to negatively impact on smaller nestlings more than on larger ones.

Drivers of passive leadership in wild songbirds: species-level differences and spatio-temporally dependent intraspecific effects

Collective behaviors are typical for many social species and can have fitness benefits for participating individuals. To maximize the benefits obtained from group living, individuals must coordinate their behaviors to some extent. What are the mechanisms that make certain individuals more likely to initiate collective behaviors, for example, by taking a risk to initially access a resource (i.e., to act as “leaders”)? Here, we examine leading behavior in a natural population of great tits and blue tits. We use automated feeding stations to monitor the feeder visits of tagged individuals within mixed-species flocks, with a small cost (waiting < 2 s) associated with the initial unlocking of the feeder. We find that great tits, males, and individuals with high activity levels were more likely to be leading in each of their feeder visits. Using a null model approach, we demonstrate that the effects of sex and activity on passive leading behavior can be explained by patterns of spatial and temporal occurrence. In other words, these effects can be explained by the times and locations of when individuals visit rather than the actual order of arrival. Hence, an analysis of the causes of leading behavior is needed to separate the effects of different processes. We highlight the importance of understanding the mechanisms behind leading behavior and discuss directions for future experimental work to gain a better understanding of the causes of leadership in natural populations.Significance statementMany species are social and engage in collective behaviors. To benefit from group actions, individuals need to fulfill different roles. Here, we examine leading behavior during feeding events; who feeds first when birds arrive at a resource? In mixed-species flocks of passerines, great tits (the larger and more dominant species), males, and individuals with higher levels of activity lead more often than blue tits, females, and individuals with lower levels of activity. While the species effect remains even when we control for the locations and dates of individual feeder visits, the effects of sex and activity are dependent on when and where birds choose to feed.

Identifying drivers of spatio-temporal variation in survival in four blue tit populations

In a context of rapid climate change, the influence of large-scale and local climate on population demography is increasingly scrutinized, yet studies are usually focused on one population. Demographic parameters, including survival, can vary significantly across populations of a given species, depending on global or local climatic fluctuations but also on many other population-specific parameters such as breeding density, habitat naturalness, predation or parasitism. Such ecological differences between populations could lead to different paces-of-life (POL), whereby populations where individuals display higher reproductive investment and bolder behaviours would have lower survival probabilities. We use here long-term (19 to 38 years) monitoring datasets from four Mediterranean populations of blue tits (Cyanistes caeruleus) to investigate the effects of sex, age class, large-scale and local climate temporal variation and population breeding density, on adult survival, using Capture-Mark-Recapture modelling. Environment heterogeneity in these four populations (two in evergreen and two in deciduous forests) has been linked to strong multi-trait phenotypic variation, suggesting blue tits in deciduous forests display faster POL compared to their conspecifics in evergreen habitats. The present results show heterogeneity in average survival probabilities across the four populations, with, as predicted, lower survival in the fast blue tits occupying deciduous habitats. Interestingly, the year-to-year variation in survival probabilities was synchronous among populations. This suggests that regional environmental conditions could drive survival fluctuations across populations. However, breeding densities were not correlated across populations, and we found no evidence that adult survival is correlated with either large-scale or local, climate temporal variation in these four blue tit populations. Finally, two of the focal populations displayed a linear temporal decrease in adult survival over the study period, calling for further investigation to explain this decline. Overall, this multi-site study shows that blue tit parental survival from one spring to the next can vary substantially across years, in a synchronous way across populations, yet the climate indices we used are not correlated with the temporal variation. This calls for further investigations in other potential drivers such as resource (in particular insect) abundance, predation or parasitism.

Phenotypic plasticity drives phenological changes in a Mediterranean blue tit population.

Earlier phenology induced by climate change, such as the passerines' breeding time, is observed in many natural populations. Understanding the nature of such changes is key to predict the responses of wild populations to climate change. Genetic changes have been rarely investigated for laying date, though it has been shown to be heritable and under directional selection, suggesting that the trait could evolve. In a Corsican blue tit population, the birds' laying date has significantly advanced over 40years, and we here determine whether this response is of plastic or evolutionary origin, by comparing the predictions of the breeder's and the Robertson-Price (STS) equations, to the observed genetic changes. We compare the results obtained for two fitness proxies (fledgling and recruitment success), using models accounting for their zero inflation. Because the trait appears heritable and under directional selection, the breeder's equation predicts that genetic changes could drive a significant part of the phenological change observed. We, however, found that fitness proxies and laying date are not genetically correlated. The STS, therefore, predicts no evolution of the breeding time, predicting correctly the absence of trend in breeding values. Our results also emphasize that when investigating selection on a plastic trait under fluctuating selection, part of the fitness-trait phenotypic covariance can be due to within individual covariance. In the case of repeated measurements, splitting within and between individual covariance can shift our perspective on the actual intensity of selection over multiple selection episodes, shedding light on the potential for the trait to evolve.

Urbanisation weakens selection on the timing of breeding and clutch size in blue tits but not in great tits

Urbanisation is a globally occurring phenomenon and is predicted to continue increasing rapidly. Urban ecosystems present novel environments and challenges which species must acclimate or adapt to. These novel challenges alter existing or create new selection pressures on behaviours which provide an opportunity to investigate eco-evolutionary responses to contemporary environmental change. We used 7 years of breeding data from urban and forest populations of blue and great tits to understand whether selection for timing of breeding or clutch size differed between the two habitats and species. We found that urban great tits laid eggs earlier than their forest counterparts, but there was no evidence of a difference in selection for earlier breeding. Blue tits, however, did not differ in timing of egg laying between the two habitats, but selection for earlier laying was weaker in the urban environment. Both species laid smaller clutches in the urban site and had positive selection for larger clutch sizes which did not differ in strength for the great tits but did for blue tits, with weaker selection in the urban population. Our results suggest that food availability for nestlings may be constraining urban birds, and that the temporal cues females use to time breeding correctly, such as tree budburst and food availability, may be absent or reduced in urban areas due to lower caterpillar availability. These results have implications for our understanding of the adaptation of wild animals to city life.Significance statementUrbanisation is expanding rapidly and changing the environment many species live in. A key challenge is to understand how species adapt to the urban environment, why some species can adapt, why others cannot and what we can do to ensure that cities are ecologically sustainable and biodiversity rich. Here we show that the strength of natural selection for early breeding and larger clutch size is weaker in urban than non-urban blue tits, likely due to reduced and irregular availability of natural insect food in urban areas. This effect was not found in great tits. Thus, urbanisation can alter the selection pressures wild animals are exposed to, but this effect may differ between species, even when closely related. This has implications for our understanding of how species adapt to urban life.

Extra‐pair paternity in Blue Tits (Cyanistes caeruleus) depends on the combination of social partners' age

There is growing evidence that engaging in extra‐pair copulations may be a strategy by which females can modify their initial mate choice if they are constrained by primary choice of the social mate. Several factors such as genetic similarity and adult phenotypic traits can affect extra‐pair paternity (EPP) patterns, but the relative importance of these factors may vary among species. Moreover, interactive effects of male and female characteristics have rarely been considered. Here, we aimed to study how multiple parameters characterizing a breeding pair (i.e. genetic similarity between mates, partners’ age and laying date) predict the occurrence of EPP at the brood level. The study uses 4 years of data from a wild population of Blue Tits Cyanistes caeruleus. Contrary to predictions of the inbreeding avoidance hypothesis, we did not find a positive relationship between the occurrence of EPP and the relatedness of social mates. We also found that the probability of EPP was unrelated to laying date. However, EPP was predicted by an interaction of social partners' ages. Specifically, EPP was less likely when old females were paired with old males in comparison to old females paired to young males. Our study suggests that the occurrence of EPP may be the result of behavioural interactions in which both male and female age are important for determining the outcome. Our results confirm the importance of considering the interactions of male and female characteristics in studies investigating EPP patterns.

Male aggressiveness and risk-taking during reproduction are repeatable but not correlated in a wild bird population

The existence of among-individual variation in behaviour within populations is poorly understood. Recent theory suggests that fine-scale individual differences in investment into current versus future reproduction may lead to a ‘slow-fast’-pace-of-life continuum, also referred to as the ‘pace-of-life-syndrome’ (POLS) hypothesis. According to this idea, individuals are predicted to differ in their level of risk-taking, which may drive among-individual variation and covariation of behaviours. Consistent individual differences in aggression, an ecologically relevant and potentially risky behaviour, have been reported across the animal kingdom. Here we test whether such individual differences in aggression are a manifestation of underlying differences in risk-taking. In a wild blue tit (Cyanistes caeruleus) population, we used standard behavioural tests to investigate if male territorial aggressiveness and risk-taking during breeding are positively related. At the start of breeding, we simulated conspecific territorial intrusions to obtain repeated measures of male aggressiveness. Subsequently, we measured male risk-taking as their latency to resume brood provisioning after presenting two different predators at their nest: human and sparrowhawk, a common predator of adult songbirds. First, we found substantial repeatability for male aggressiveness (R = 0.56 ± 0.08 SE). Second, while males took longer to resume provisioning after presentation of a sparrowhawk mount as compared to a human observer, risk-taking was repeatable across these two predator contexts (R = 0.51 ± 0.13 SE). Finally, we found no evidence for a correlation between male aggressiveness and risk-taking, thereby providing little support to a main prediction of the POLS hypothesis.Significance statementConsistent, and often correlated, individual differences in basal behaviours, such as aggression, exploration and sociability, are found across the animal kingdom. Why individuals consistently differ in their behaviour is poorly understood, as behavioural traits would seem inherently flexible. The ‘pace-of-life syndrome’ (POLS) hypothesis proposes observed behavioural variation to reflect differences in risk-taking associated with individual reproductive strategies. We tested this idea in a wild blue tit population by investigating whether individual males that were more aggressive toward territorial intruders also took more risk when provisioning their nestlings under a threat of predation. While we found consistent individual differences in both aggressiveness and risk-taking, these behaviours were not significantly correlated. Therefore, our study demonstrates among-individual variation in ecologically relevant behaviours in wild blue tits but provides little support for the POLS hypothesis.