Combining personal with social information facilitates host defences and explains why cuckoos should be secretive.

Individuals often vary defences in response to local predation or parasitism risk. But how should they assess threat levels when it pays their enemies to hide? For common cuckoo hosts, assessing parasitism risk is challenging: cuckoo eggs are mimetic and adult cuckoos are secretive and resemble hawks. Here, we show that egg rejection by reed warblers depends on combining personal and social information of local risk. We presented model cuckoos or controls at a pair's own nest (personal information of an intruder) and/or on a neighbouring territory, to which they were attracted by broadcasts of alarm calls (social information). Rejection of an experimental egg was stimulated only when hosts were alerted by both social and personal information of cuckoos. However, pairs that rejected eggs were not more likely to mob a cuckoo. Therefore, while hosts can assess risk from the sight of a cuckoo, a cuckoo cannot gauge if her egg will be accepted from host mobbing. Our results reveal how hosts respond rapidly to local variation in parasitism, and why it pays cuckoos to be secretive, both to avoid alerting their targets and to limit the spread of social information in the local host neighbourhood.We thank the Natural Environment Research Council (NERC) and Society in Science – Branco Weiss for financial support.This is the final version of the article. It first appeared from Nature Publishing Group via http://dx.doi.org/10.1038/srep1987

Older and wiser? Age differences in foraging and learning by an endangered passerine

Birds use cues when foraging to help relocate food resources, but natural environments provide many potential cues and choosing which to use may depend on previous experience. Young animals have less experience of their environment compared to adults, so may be slower to learn cues or may need to sample the environment more. Whether age influences cue use and learning has, however, received little experimental testing in wild animals. Here we investigate effects of age in a wild population of hihi (Notiomystis cincta), a threatened New Zealand passerine. We manipulated bird feeders using a novel colour cue to indicate a food reward; once hihi learned its location, we rotated the feeder to determine whether the birds followed the colour or returned to the previous location. Both age groups made fewer errors over trials and learned the location of the food reward, but juveniles continued to sample unrewarding locations more than adults. Following a second rotation, more adults preferred to forage from the hole indicated by the colour cue than juveniles, despite this no longer being rewarding. Overall, juveniles spent longer in the feeder arena to reach the same proportion of foraging time as adults. Combined, these results suggest that juveniles and adults may use an “explore and exploit” foraging strategy differently, and this affects how efficiently they forage. Further work is needed to understand how juveniles may compensate for their inexperience in learning and foraging strategies.Peer reviewe

Social transmission of avoidance among predators facilitates the spread of novel prey

Warning signals are an effective defence strategy for aposematic prey, but only if they are recognized by potential predators. If predators must eat prey to associate novel warning signals with unpalatability, how can aposematic prey ever evolve? Using experiments with great tits (Parus major) as predators, we show that social transmission enhances the acquisition of avoidance by a predator population. Observing another predator’s disgust towards tasting one novel conspicuous prey item led to fewer aposematic than cryptic prey being eaten for the predator population to learn. Despite reduced personal encounters with unpalatable prey, avoidance persisted and increased over subsequent trials. Next we use a mathematical model to show that social transmission can shift the evolutionary trajectory of prey populations from fixation of crypsis to fixation of aposematism more easily than was previously thought. Therefore, social information use by predators has the potential to have evolutionary consequences across ecological communities.Peer reviewe

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.Peer reviewe

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

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.Peer reviewe

Grey Gerygone hosts are not egg rejecters, but Shining Bronze-Cuckoos lay cryptic eggs

Many brood parasites rely on mimicry to prevent the detection of their eggs by hosts, yet most Australasian cuckoo species lay darkly colored eggs while the eggs of their hosts are pale and speckled. In the dimly lit nests of their hosts, these cuckoo eggs may appear cryptic; however, it is unclear if this disguise has evolved to fool hosts or other cuckoos. Recent work suggests that in at least one species of bronze-cuckoo, cuckoos are more likely to reject conspicuous eggs than are hosts, but it remains unclear whether this is common across the species group. Here, we present field experiments on the sole host of the Shining Bronze-Cuckoo (Chalcites lucidus lucidus) in New Zealand, the Grey Gerygone (Gerygone igata; known locally as the Grey Warbler), that explored whether this host ignores cuckoo eggs because they are cryptic. Using an avian vision model, we showed that Shining Bronze-Cuckoo eggs were variable in their conspicuousness, but were more cryptic in host nests than the host's eggs. We then experimentally parasitized all available clutches with model eggs that mimicked darkly or brightly colored cuckoo eggs, or were of maximum conspicuousness (white) as determined by visual modeling. Hosts never rejected our model eggs, nor cuckoo eggs when naturally parasitized. Instead, only cuckoos rejected model eggs: In 3 out of 4 experimental nests that were subsequently parasitized, the model egg was taken and replaced by a cuckoo's egg. Together, these data and previous experiments suggest that competition among cuckoos, rather than rejection by hosts, provides a stronger selection pressure for the evolution of cryptic eggs across the genus Chalcites.Peer reviewe

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.Peer reviewe

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 differ-ently 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 ob-served 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.Peer reviewe

Enhancing dietary specialization metrics in observational studies of wild animal populations

Studies of intraspecific dietary variation can greatly enrich our view of a species' niche and role in the ecosystem, particularly when species with broad diets are found to be composed of generalist and specialist individuals. However, the current framework for quantifying dietary specialization leaves certain standards unformalized and is susceptible to overestimating specialization when there are few repeated observations per individual, as is often the case in observational studies of wild populations. Here, we use the hihi (Notiomystis cincta), a threatened New Zealand passerine, as a case study for demonstrating how existing statistical tools can be applied to strengthen the dietary specialization framework. First, we assess whether the reliability of common dietary measures can be improved through Bayesian adjustments and by using rarefaction to compare uncertainty levels of metrics calculated from different sample sizes. As diet links closely to environmental factors, we also demonstrate how adding phenological data and habitat assessments to standard protocols can help validate our dietary measures as evidence for resource selection rather than random foraging. Finally, in light of our finding that diet predicts survival in hihi, we discuss the utility of dietary specialization for elucidating broader behavioral syndromes.Peer reviewe