Parental breeding decisions and genetic quality predict social structure of independent offspring.

Across the animal kingdom, newly independent juveniles form social associations that influence later fitness, mate choice and gene flow, but little is known about the ontogeny of social environments, particularly in wild populations. Here we test whether associations among young animals form randomly or are influenced by environmental or genetic conditions established by parents. Parents' decisions determine natal birth sites, which could affect who independent young initially encounter; secondly, mate choice determines genetic condition (e.g. inbreeding) of young and the parental care they receive, which can affect sociability. However, genetic and environmental factors are confounded unless related offspring experience different natal environments. Therefore, we used a long-term genetic pedigree, breeding records and social network data from three cohorts of a songbird with high extra-pair paternity (hihi, Notiomystis cincta) to disentangle (1) how nest location and relatedness contribute to association structure once juveniles disperse away from birth sites, and (2) if juvenile and/or parental inbreeding predicts individual sociability. We detected positive spatial autocorrelation: hihi that fledged closer by were more likely to associate even after dispersing, irrespective of genetic relatedness. Juvenile inbreeding did not predict sociability, but those raised by more inbred fathers formed more, stronger, associations, which did not depend on whether that male was the genetic parent or not. These results suggest that the natal environment created by parents, rather than focal genetic condition, establishes the foundation for social associations. Overall, we highlight how social inheritance may play an important role in population dynamics and evolutionary potential in wild animals.

The evolution and ecology of multiple antipredator defences.

Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such "defence portfolios" that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies.

Polymorphism at the nestling stage and host-specific mimicry in an Australasian cuckoo-host arms race.

Decades of research have shown that the coevolutionary arms race between avian brood parasites and their hosts can promote phenotypic diversification in hosts and brood parasites. However, relatively little is known about the role of brood parasitism in promoting phenotypic diversification of nestlings. We review field data collected over four decades in Australia, New Caledonia and New Zealand to assess potential for coevolutionary interactions between the shining bronze-cuckoo (Chalcites lucidus) and its hosts, and how diversification at the nestling stage may be generating different subspecies. The shining bronze-cuckoo is a specialist parasite of a few hosts in the family Acanthizidae. It has diversified into subspecies, of which the nestlings closely mimic the respective host nestlings in each region. Additionally, some cuckoo subspecies have polymorphic nestlings. The Acanthizidae hosts have similar breeding and nesting habits and only moderately effective frontline defences against parasitism at cuckoo egg laying or at the egg stages. However, some hosts have developed highly effective defences at the nestling stage by recognising and ejecting cuckoo nestlings from the nest. As with the cuckoo nestlings, some hosts have polymorphic nestlings. The coevolutionary interactions in each region suggest different evolutionary stages of the arms race in which either the parasite or the host is currently in the lead. The presence of moderately effective defences at the egg laying and egg stages might explain why some hosts do not have defences at the nestling stage. The south-Pacific cuckoo - host systems are excellent models to explore the evolutionary mechanisms driving the diversification at the nestling stage in the coevolutionary arms race between avian brood parasites and their hosts.

Evaluating the success of functional restoration after reintroduction of a lost avian pollinator.

Conservation translocation is a common method for species recovery, for which one increasingly frequent objective is restoring lost ecological functions to promote ecosystem recovery. However, few conservation translocation programs explicitly state or monitor function as an objective, limiting the ability to test assumptions, learn from past efforts, and improve management. We evaluated whether translocations of hihi (Notiomystis cincta), a threatened New Zealand passerine, achieved their implicit objective of restoring lost pollination function. Through a pollinator-exclusion experiment, we quantified, with log response ratios (lnR), the effects of birds on fruit set and seed quality in hangehange (Geniostoma ligustrifolium), a native flowering shrub. We isolated the contributions of hihi by making comparisons across sites with and without hihi. Birds improved fruit set more at sites without hihi (lnR = 1.27) than sites with hihi (lnR = 0.50), suggesting other avian pollinators compensated for and even exceeded hihi contributions to fruit set. Although birds improved seed germination only at hihi sites (lnR = 0.22-0.41), plants at sites without hihi had germination rates similar to hihi sites because they produced 26% more filled seeds, regardless of pollination condition. Therefore, although our results showed hihi improved seed quality, they also highlighted the complexity of ecological functions. When an important species is lost, ecosystems may be able to achieve similar function through different means. Our results underscore the importance of stating and monitoring the ecological benefits of conservation translocations when functional restoration is a motivation to ensure these programs are achieving their objectives.

Evaluación del Éxito de la Restauración Funcional Posterior a la Reintroducción de un Ave Polinizadora Desaparecida Resumen La reubicación para la conservación es un método común para la recuperación de especies en el cual un objetivo cada vez más frecuente es la restauración de las funciones ecológicas que se perdieron para promover la recuperación del ecosistema. Sin embargo, pocos programas de reubicación para la conservación establecen o monitorean explícitamente a la función como un objetivo, lo que limita la posibilidad de comprobar suposiciones, aprender de esfuerzos anteriores y mejorar la gestión. Analizamos si las reubicaciones de hihi (Notiomystis cincta), un ave paseriforme amenazada de Nueva Zelanda, lograron el objetivo implícito de restaurar la desaparecida función de polinización. Mediante un experimento de exclusión del polinizador, cuantificamos con relaciones de respuesta logarítmica (lnR) los efectos de las aves sobre el conjunto de frutos y la calidad de la semilla del arbusto floral nativo Geniostoma ligustrifolium. Aislamos las contribuciones del hihi cuando comparamos entre sitios con y sin su presencia. Las aves favorecieron más al conjunto de frutos en sitios sin hihi (lnR = 1.27) que en los sitios con hihi (lnR = 0.50), lo que sugiere que otras aves polinizadoras compensaron y excedieron las contribuciones del hihi al conjunto de frutos. Aunque las aves aumentaron la germinación de semillas sólo en sitios con hihi (lnR = 0.22-0.41), las plantas en los sitios sin hihi tuvieron tasas de germinación similares a los sitios con hihi porque produjeron 26% más de semillas completas sin importar la condición de la polinización. Por lo tanto, aunque nuestros resultados mostraron mejoras en la calidad de la semilla a causa del hihi, también resaltaron la complejidad de las funciones ecológicas. Cuando desaparece una especie importante, puede que los ecosistemas logren una función similar por medio de diferentes métodos. Nuestros resultados hacen hincapié en la importancia que tiene establecer y monitorear los beneficios ecológicos de las reubicaciones para la conservación cuando la restauración es motivo para asegurar que estos programas están logrando sus objetivos.

Collaboration enhances career progression in academic science, especially for female researchers.

Collaboration and diversity are increasingly promoted in science. Yet how collaborations influence academic career progression, and whether this differs by gender, remains largely unknown. Here, we use co-authorship ego networks to quantify collaboration behaviour and career progression of a cohort of contributors to biennial International Society of Behavioral Ecology meetings (1992, 1994, 1996). Among this cohort, women were slower and less likely to become a principal investigator (PI; approximated by having at least three last-author publications) and published fewer papers over fewer years (i.e. had shorter academic careers) than men. After adjusting for publication number, women also had fewer collaborators (lower adjusted network size) and published fewer times with each co-author (lower adjusted tie strength), albeit more often with the same group of collaborators (higher adjusted clustering coefficient). Authors with stronger networks were more likely to become a PI, and those with less clustered networks did so more quickly. Women, however, showed a stronger positive relationship with adjusted network size (increased career length) and adjusted tie strength (increased likelihood to become a PI). Finally, early-career network characteristics correlated with career length. Our results suggest that large and varied collaboration networks are positively correlated with career progression, especially for women.

Hosts elevate either within-clutch consistency or between-clutch distinctiveness of egg phenotypes in defence against brood parasites.

In host-parasite arms races, hosts can evolve signatures of identity to enhance the detection of parasite mimics. In theory, signatures are most effective when within-individual variation is low ('consistency'), and between-individual variation is high ('distinctiveness'). However, empirical support for positive covariation in signature consistency and distinctiveness across species is mixed. Here, we attempt to resolve this puzzle by partitioning distinctiveness according to how it is achieved: (i) greater variation within each trait, contributing to elevated 'absolute distinctiveness' or (ii) combining phenotypic traits in unpredictable combinations ('combinatorial distinctiveness'). We tested how consistency covaries with each type of distinctiveness by measuring variation in egg colour and pattern in two African bird families (Cisticolidae and Ploceidae) that experience mimetic brood parasitism. Contrary to predictions, parasitized species, but not unparasitized species, exhibited a negative relationship between consistency and combinatorial distinctiveness. Moreover, regardless of parasitism status, consistency was negatively correlated with absolute distinctiveness across species. Together, these results suggest that (i) selection from parasites acts on how traits combine rather than absolute variation in traits, (ii) consistency and distinctiveness are alternative rather than complementary elements of signatures and (iii) mechanistic constraints may explain the negative relationship between consistency and absolute distinctiveness across species.

Foraging behaviour alters with social environment in a juvenile songbird.

Early independence from parents is a critical period where social information acquired vertically may become outdated, or conflict with new information. However, across natural populations, it is unclear if newly independent young persist in using information from parents, or if group-level effects of conformity override previous behaviours. Here, we test if wild juvenile hihi (Notiomystis cincta, a New Zealand passerine) retain a foraging behaviour from parents, or if they change in response to the behaviour of peers. We provided feeding stations to parents during chick-rearing to seed alternative access routes, and then tracked their offspring's behaviour. Once independent, juveniles formed mixed-treatment social groups, where they did not retain preferences from their time with parents. Instead, juvenile groups converged over time to use one access route- per group, and juveniles that moved between groups switched to copy the locally favoured option. Juvenile hihi did not copy specific individuals, even if they were more familiar with the preceding bird. Our study shows that early social experiences with parents affect initial foraging decisions, but social environments encountered later on can update transmission of arbitrary behaviours. This suggests that conformity may be widespread in animal groups, with potential cultural, ecological and evolutionary consequences.

Seeing red? Colour biases of foraging birds are context dependent.

Colours are commonly used as visual cues when measuring animals' cognitive abilities. However, animals can have innate biases towards certain colours that depend on ecological and evolutionary contexts, therefore potentially influencing their performance in experiments. For example, when foraging, the colour red can advertise profitable fruits or act as a warning signal about chemically defended prey, and an individual's propensity to take food of that colour may depend on experience, age or physical condition. Here, we investigate how these contexts influence blue tits' (Cyanistes caeruleus) and great tits' (Parus major) responses to red-coloured almond flakes. We found that juvenile birds preferred red both when it was presented simultaneously with green, and when it was presented with three alternative colours (orange, purple, green). Adult birds, however, only preferred red after a positive experience with the colour, or when it was presented with the three alternative colours. We then tested whether colour influenced avoidance learning about food unpalatability. Despite the prediction that red is a more salient warning signal than green, we found only weak evidence that birds discriminated red unpalatable almonds from a green palatable alternative more quickly than when the colours were reversed. Our results suggest that biases towards red food may depend on birds' age and previous experience, and this might influence their performance in experiments that use red stimuli. Considering the ecological relevance of colours is, therefore, important when designing experiments that involve colour cues.

Social learning within and across predator species reduces attacks on novel aposematic prey.

To make adaptive foraging decisions, predators need to gather information about the profitability of prey. As well as learning from prey encounters, recent studies show that predators can learn about prey defences by observing the negative foraging experiences of conspecifics. However, predator communities are complex. While observing heterospecifics may increase learning opportunities, we know little about how social information use varies across predator species. Social transmission of avoidance among predators also has potential consequences for defended prey. Conspicuous aposematic prey are assumed to be an easy target for naïve predators, but this cost may be reduced if multiple predators learn by observing single predation events. Heterospecific information use by predators might further benefit aposematic prey, but this remains untested. Here we test conspecific and heterospecific information use across a predator community with wild-caught blue tits (Cyanistes caeruleus) and great tits (Parus major). We used video playback to manipulate social information about novel aposematic prey and then compared birds' foraging choices in 'a small-scale novel world' that contained novel palatable and aposematic prey items. We expected that blue tits would be less likely to use social information compared to great tits. However, we found that both blue tits and great tits consumed fewer aposematic prey after observing a negative foraging experience of a demonstrator. In fact, this effect was stronger in blue tits compared to great tits. Interestingly, blue tits also learned more efficiently from watching conspecifics, whereas great tits learned similarly regardless of the demonstrator species. Together, our results indicate that social transmission about novel aposematic prey occurs in multiple predator species and across species boundaries. This supports the idea that social interactions among predators can reduce attacks on aposematic prey and therefore influence selection for prey defences.

Native fauna interact differently with native and alien trees in a tropical megacity.

The negative effects of invasive alien plant species on natural ecosystems are well known. However, in rapidly growing cities, alien plants can provide native fauna with resources otherwise lost due to the biotic homogenization, which is common to urban ecosystems. Interactions of native fauna with alien flora have thus far focused largely on invertebrate pollinators in temperate cities in the northern hemisphere. Cities in tropical areas, however, are larger and are growing more rapidly, and host a variety of vertebrate pollinators. Understanding how birds and mammals interact with native and alien flora in these megacities could improve management of urban ecosystems in highly biodiverse regions while limiting invasion potential. Therefore, here we investigate whether native diurnal birds and mammals interact differently with native versus alien trees in Bengaluru, India where historical planting has led to an abundance of alien tree species. We find that tree origin alone was not an important predictor for bird species richness and abundance, but taller native trees with large floral display sizes were more species rich than alien trees of similar floral displays. As expected from their shared evolutionary history, nectarivorous birds fed from native trees more often in a manner that could facilitate pollination, but engaged in nectar theft more often with alien trees. Squirrels (the mammal observed most frequently to interact with flowers) were more likely, however, to depredate flowers of native trees. Our results suggest alien trees can be an important resource for fauna in expanding urban areas, and that nectar theft by birds could reduce the seed set of alien trees.

Maternally invested carotenoids compensate costly ectoparasitism in the hihi.

Dietary ingested carotenoid biomolecules have been linked to both improved health and immunity in nestling birds. Here, we test whether maternally invested egg carotenoids can offset the cost of parasitism in developing nestling hihi (Notiomystis cincta) from the bloodsucking mite (Ornithonyssus bursa). Our results reveal clear negative effects of parasitism on nestlings, and that maternally derived carotenoids compensate this cost, resulting in growth parameters and ultimate mass achieved being similar to nonparasitized young. Our results offer an unique example of a direct positive relationship between enhanced maternal investment of carotenoids and an ability to cope with a specific and costly parasite in young birds. As O. bursa infestations reduce population viability in hihi, our findings also highlight the importance of key nutritional resources for endangered bird populations to better cope with common parasite infestations.

Snow depth drives habitat selection by overwintering birds in built-up areas, farmlands and forests.

Aim: Abundances of animals vary according to species-specific habitat selection, but habitats are undergoing rapid change in response to anthropogenic alterations of land use and climate. The long-term decline of snowfall is one of the most dramatic abiotic changes in boreal regions, with potential to alter species communities and shape future ecosystems. However, the effects of snow cover on habitat-specific abundances remain unclear for many taxa. Here we explore whether long-term declines in snow cover affect the abundances of overwintering birds. Taxon: Fifty bird species. Location: Finland, Northern Europe. Methods: We used generalized linear mixed models to analyse citizen-led monitoring data from 196 transects over a 32-year period to assess whether abundances of birds have changed in built-up areas, farmlands and forests, and whether these covary with warming temperatures and decreasing snow. We then explored if changes in abundance can be explained by body mass, migration strategy or feeding guilds of the species. Results: Over the study period, the abundance of overwintering birds increased. This increase was most pronounced in farmlands (69.6%), where abundances were positively associated with decreasing snow depth. On the other hand, while abundances in built-up habitats (19.5%) decreased over the study period, they increased in periods of high snow depths. Finally, we found that the short-distance migration strategy explains changes in bird abundances with snow. In farmlands, ground feeding birds and heavier birds also show a positive trends in abundance with decreasing snow depths. Main conclusions: Local snow conditions are driving habitat selection of birds in the winter; birds in farmlands were most responsive to a decrease in snow depth. Changing snow depths can affect bird movements across habitats in the winter, but also influence migratory patterns and range shifts of species.

Prior experience of captivity affects behavioural responses to 'novel' environments.

Information ecology theory predicts that prior experience influences current behaviour, even if the information is acquired under a different context. However, when individuals are tested to quantify personality, cognition, or stress, we usually assume that the novelty of the test is consistent among individuals. Surprisingly, this 'gambit of prior experience' has rarely been explored. Therefore, here we make use of a wild population of great tits (Parus major) to test if prior experience of handling and captivity influences common measures of exploration (open field tests in two novel contexts: room and cage arenas), social response (simulated using a mirror), and behavioural stress (breathing rate). We found that birds with prior experience of captivity (caught previously for unrelated learning and foraging experiments) were more exploratory, but this depended on age: exploration and captivity experience (in terms of both absolute binary experience and the length of time spent in captivity) were associated more strongly in young (first-winter) birds than in adults. However, there was no association of prior experience of captivity with social response and breathing rate, and nor did the measures of exploration correlate. Together our results suggest that re-testing of individuals requires careful consideration, particularly for younger birds, and previous experiences can carry over and affect behaviours differently.

Sense and sensitivity: responsiveness to offspring signals varies with the parents' potential to breed again.

How sensitive should parents be to the demands of their young? Offspring are under selection to seek more investment than is optimal for parents to supply, which makes parents vulnerable to losing future fitness by responding to manipulative displays. Yet, parents cannot afford to ignore begging and risk allocating resources inefficiently. Here, we show that parents may solve this problem by adjusting their sensitivity to begging behaviour in relation to their own likelihood of breeding again, a factor largely neglected in previous analyses of parent-offspring interactions. In two carotenoid-supplementation experiments on a New Zealand passerine, the hihi Notiomystis cincta, we supplemented adults to enhance their propensity to breed again, and supplemented entire broods to increase their mouth colour, thus enhancing their solicitation display. We found that adults that attempted two breeding attempts a season were largely insensitive to the experimentally carotenoid-rich gapes of their brood, whereas those that bred just once responded by increasing their rate of provisioning at the nest. Our results show that parents can strategically vary their sensitivity to begging in relation to their future reproductive potential. By restricting opportunities for offspring to influence provisioning decisions, parents greatly limit the potential for offspring to win parent-offspring conflict.

Demographic consequences of adult sex ratio in a reintroduced hihi population.

1. Male-biased adult sex ratios are frequently observed in free-ranging populations and are known to cause changes in mating behaviours including increased male harassment of females, which can cause injury to females and/or alter female behaviour during breeding. 2. Although we can explain why such behaviours may evolve and have studied their impacts on individuals when it does, we know very little about the demographic consequences of harassment caused by changes in adult sex ratio. 3. Using a 12-year longitudinal data set of a free-living and endangered New Zealand passerine, the hihi (Notiomystis cincta), we show that a changing adult sex ratio has little or no effect on adult female survival or the number of fledglings produced per female. This is despite clear evidence of male harassment of breeding females when the sex ratio was male biased (up to three males per female). 4. The length of the study and major fluctuations in sex ratio observed made it possible to obtain narrow confidence or credible intervals for effect sizes, showing that any effect of sex ratio on demographic rates were small. 5. Our results provide rare empirical evidence for the demographic consequences of biased adult sex ratios in the wild and particularly in a conservation context.

Social transmission in the wild can reduce predation pressure on novel prey signals.

Social transmission of information is taxonomically widespread and could have profound effects on the ecological and evolutionary dynamics of animal communities. Demonstrating this in the wild, however, has been challenging. Here we show by field experiment that social transmission among predators can shape how selection acts on prey defences. Using artificial prey and a novel approach in statistical analyses of social networks, we find that blue tit (Cyanistes caeruleus) and great tit (Parus major) predators learn about prey defences by watching others. This shifts population preferences rapidly to match changes in prey profitability, and reduces predation pressure from naïve predators. Our results may help resolve how costly prey defences are maintained despite influxes of naïve juvenile predators, and suggest that accounting for social transmission is essential if we are to understand coevolutionary processes.

A Chromosome-Level Genome Assembly of the Reed Warbler (Acrocephalus scirpaceus).

The reed warbler (Acrocephalus scirpaceus) is a long-distance migrant passerine with a wide distribution across Eurasia. This species has fascinated researchers for decades, especially its role as host of a brood parasite, and its capacity for rapid phenotypic change in the face of climate change. Currently, it is expanding its range northwards in Europe, and is altering its migratory behavior in certain areas. Thus, there is great potential to discover signs of recent evolution and its impact on the genomic composition of the reed warbler. Here, we present a high-quality reference genome for the reed warbler, based on PacBio, 10×, and Hi-C sequencing. The genome has an assembly size of 1,075,083,815 bp with a scaffold N50 of 74,438,198 bp and a contig N50 of 12,742,779 bp. BUSCO analysis using aves_odb10 as a model showed that 95.7% of BUSCO genes were complete. We found unequivocal evidence of two separate macrochromosomal fusions in the reed warbler genome, in addition to the previously identified fusion between chromosome Z and a part of chromosome 4A in the Sylvioidea superfamily. We annotated 14,645 protein-coding genes, and a BUSCO analysis of the protein sequences indicated 97.5% completeness. This reference genome will serve as an important resource, and will provide new insights into the genomic effects of evolutionary drivers such as coevolution, range expansion, and adaptations to climate change, as well as chromosomal rearrangements in birds.

The signal detection problem of aposematic prey revisited: integrating prior social and personal experience.

Ever since Alfred R. Wallace suggested brightly coloured, toxic insects warn predators about their unprofitability, evolutionary biologists have searched for an explanation of how these aposematic prey evolve and are maintained in natural populations. Understanding how predators learn about this widespread prey defence is fundamental to addressing the problem, yet individuals differ in their foraging decisions and the predominant application of associative learning theory largely ignores predators' foraging context. Here we revisit the suggestion made 15 years ago that signal detection theory provides a useful framework to model predator learning by emphasizing the integration of prior information into predation decisions. Using multiple experiments where we modified the availability of social information using video playback, we show that personal information (sampling aposematic prey) improves how predators (great tits, Parus major) discriminate between novel aposematic and cryptic prey. However, this relationship was not linear and beyond a certain point personal encounters with aposematic prey were no longer informative for prey discrimination. Social information about prey unpalatability reduced attacks on aposematic prey across learning trials, but it did not influence the relationship between personal sampling and discrimination. Our results suggest therefore that acquiring social information does not influence the value of personal information, but more experiments are needed to manipulate pay-offs and disentangle whether information sources affect response thresholds or change discrimination. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

The effect of social information from live demonstrators compared to video playback on blue tit foraging decisions.

Video playback provides a promising method to study social interactions, and the number of video playback experiments has been growing in recent years. Using videos has advantages over live individuals as it increases the repeatability of demonstrations, and enables researchers to manipulate the features of the presented stimulus. How observers respond to video playback might, however, differ among species, and the efficacy of video playback should be validated by investigating if individuals' responses to videos are comparable to their responses to live demonstrators. Here, we use a novel foraging task to compare blue tits' (Cyanistes caeruleus) responses to social information from a live conspecific vs video playback. Birds first received social information about the location of food, and were then presented with a three-choice foraging task where they could search for food from locations marked with different symbols (cross, square, plain white). Two control groups saw only a foraging tray with similar symbols but no information about the location of food. We predicted that socially educated birds would prefer the same location where a demonstrator had foraged, but we found no evidence that birds copied a demonstrator's choice, regardless of how social information was presented. Social information, however, had an influence on blue tits' foraging choices, as socially educated birds seemed to form a stronger preference for a square symbol (against two other options, cross and plain white) than the control birds. Our results suggest that blue tits respond to video playback of a conspecific similarly as to a live bird, but how they use this social information in their foraging decisions, remains unclear.

The coevolutionary biology of brood parasitism: a call for integration.

Obligate brood-parasitic cheats have fascinated natural historians since ancient times. Passing on the costs of parental care to others occurs widely in birds, insects and fish, and often exerts selection pressure on hosts that in turn evolve defences. Brood parasites have therefore provided an illuminating system for researching coevolution. Nevertheless, much remains unknown about how ecology and evolutionary history constrain or facilitate brood parasitism, or the mechanisms that shape or respond to selection. In this special issue, we bring together examples from across the animal kingdom to illustrate the diverse ways in which recent research is addressing these gaps. This special issue also considers how research on brood parasitism may benefit from, and in turn inform, related fields such as social evolution and immunity. Here, we argue that progress in our understanding of coevolution would benefit from the increased integration of ideas across taxonomic boundaries and across Tinbergen's Four Questions: mechanism, ontogeny, function and phylogeny of brood parasitism. We also encourage renewed vigour in uncovering the natural history of the majority of the world's brood parasites that remain little-known. Indeed, it seems very likely that some of nature's brood parasites remain entirely unknown, because otherwise we are left with a puzzle: if parental care is so costly, why is brood parasitism not more common? This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.