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 statement Many 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.

The orientation of nestboxes influences their occupation rates and the breeding success of passerine birds

Nestboxes are widely provided as nesting sites for hole-nesting birds, yet the relative contribution of nestbox characteristics and habitat quality in determining the occupancy rates and breeding success of birds remains unclear. We provided nestboxes in deciduous woodlands in England and examined if those nestboxes were erected in random orientations and whether the orientation of nestboxes and habitat quality, in the form of tree density, influenced their occupation by, and breeding success of, Blue Tits (Cyanistes caeruleus), Great Tits (Parus major) and Pied Flycatchers (Ficedula hypoleuca). We found that first, the nestboxes were erected non-randomly orientated towards the north and east, and away from the south and west. Second, the occupation rates of none of the species was related to nestbox orientation or tree density. Third, the breeding success of neither Blue Tits nor Great Tits varied with tree density but did vary with nestbox orientation. Blue Tit hatching success and fledging success was higher in nestboxes facing south than in other directions whilst in Great Tits, clutch sizes, hatching success and fledging success was higher in nestboxes facing south than nestboxes facing other directions. Our results suggest that nestbox characteristics, such as orientation, have more influence on the reproductive success of passerines than habitat quality. This further suggests that conservationists should orientate nestboxes southwards in order to maximise their benefit to birds in temperate climates during the breeding season.

Turn taking is not restricted by task specialisation but does not facilitate equality in offspring provisioning

Sexual conflict arises when two individuals invest in their common offspring because both individuals benefit when their partner invests more. Conditional cooperation is a theoretical concept that could resolve this conflict. Here, parents are thought to motivate each other to contribute to provisioning visits by following the rules of turn taking, which results in equal and efficient investment. However, parents have other tasks besides provisioning, which might hinder taking turns. To investigate restrictions by other care tasks and whether turn taking can be used to match investment, we manipulated brooding duration in female blue tits (Cyanistes caeruleus) during the early nestling phase by changing nest box temperature. As expected, females subjected to cold conditions brooded longer than females under warm conditions. Yet, contrary to our prediction, females had similar visit rates in both treatments, which suggests that females in the cold treatment invested more overall. In addition, the females’ turn taking level was higher in the more demanding cold condition (and the calculated randomised turn taking levels of females did not differ), hence females don’t seem to be restricted in their turn taking strategy by other care tasks. However, males did not seem to match the females’ turn taking levels because they did not adjust their visit rates. Thus, level of turn taking was not restricted by an other sex-specific task in females and did not facilitate a greater investment by their male partners.

Decomposing phenotypic skew and its effects on the predicted response to strong selection

The major frameworks for predicting evolutionary change assume that a phenotype's underlying genetic and environmental components are normally distributed. However, the predictions of these frameworks may no longer hold if distributions are skewed. Despite this, phenotypic skew has never been decomposed, meaning the fundamental assumptions of quantitative genetics remain untested. Here, we demonstrate that the substantial phenotypic skew in the body size of juvenile blue tits (Cyanistes caeruleus) is driven by environmental factors. Although skew had little impact on our predictions of selection response in this case, our results highlight the impact of skew on the estimation of inheritance and selection. Specifically, the non-linear parent-offspring regressions induced by skew, alongside selective disappearance, can strongly bias estimates of heritability. The ubiquity of skew and strong directional selection on juvenile body size implies that heritability is commonly overestimated, which may in part explain the discrepancy between predicted and observed trait evolution.

Nest-boxes alter the reproductive ecology of urban cavity-nesters in a species-dependent way

To mitigate the shortage of natural breeding sites in cities, nest-boxes are provided for cavity-nesters. However, these are not the breeding sites these animals originally evolved in and optimised their breeding performance to. It thus remains inconclusive if nest-boxes can provide adequate substitutes, ensuring equivalent fitness returns for breeding animals. Additionally, the majority of knowledge on the ecology of urban birds comes from nest-box populations, but no study to date directly compared fitness consequences of breeding inside nest-boxes in relation to natural-cavities in an urban context. This limits our understanding of the urban ecology of cavity-nesters and addressing its functional meaning. We investigate fitness consequences and life-history trait variation according to the nesting site type to provide a comprehensive understanding of conservation potential of nest-boxes in cities and to support/question generalisations stemming from nest-box studies on urbanization. We directly compare the reproductive performance of two small passerines, blue tits and great tits, breeding in nest-boxes as opposed to natural-cavities in a seminatural forest of a capital city using a quasi-experimental setting. We show that the effects of nest type vary between species: in blue tits, fitness proxies were negatively affected by nest-boxes (lower fledging success and fledgling numbers, longer time spent in nest and later fledging date in comparison to natural-cavities), while great tit performance appeared to be unaffected by nest type. We detected that both species breeding in nest-boxes accelerated incubation onset, but since there were no major differences in pre-hatching traits (lay date, clutch size, hatching rates) between the nest types, we attribute the fitness deterioration to post-hatching effects. Interestingly, overall breeding density of tits in urban natural-cavities was higher than observed in a primeval habitat. Nest-boxes may become an ecological trap for some species and the unaffected species can consequently outcompete them, decreasing overall biodiversity in cities. We highlight the ecological importance of old-growth tree stands, providing natural tree cavities for city-breeding animals. Due to the detected nest type-dependent variation in reproductive performance, we support the criticism regarding the unconditional extrapolation of evolutionary and ecological interpretations of nest-box studies to general populations.

Chemical analysis reveals sex differences in the preen gland secretion of breeding Blue Tits

Acoustic and visual signals are well known to play important roles in social communication in birds. Growing evidence suggests that many bird species, including species of songbirds, additionally have a well-developed sense of smell. However, we are still at the beginning of understanding the potential importance of chemical communication in the social lives of birds, for example in mate choice. The secretion of the preen gland may be an important contributor to the chemical phenotype of birds. Here, we report on a first characterisation of the chemical composition of the preen gland secretion of the Blue Tit (Cyanistes caeruleus), a common songbird which is an often used model species in animal behaviour and ecology, in particular also in studies of sexual selection and (extra-pair) mate choice. We found sex differences in the composition of the preen gland secretion in breeding Blue Tits. Females further tended to have a larger number of putative compounds in their secretions compared to males. We briefly discuss the possible implications of these findings and speculate that the chemical composition of the preen gland secretion may be a sexually selected trait in Blue Tits. Our preliminary findings warrant follow-up research into the patterns of within- and among individual variation in the chemical composition of the preen gland secretion as well as the identification of the main chemical compounds involved.