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

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

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

Minutes matter: brief hatching asynchrony adversely affects late-hatched hihi nestlings, but not life beyond the nest

Size hierarchies are often seen when nestlings hatch asynchronously over a period of days. Shorter hatch periods are common across passerines, however, and while these may also give rise to asymmetries, their effects are rarely considered. Regardless of hatch period, the long-term consequences for later hatched nestlings that survive to fledge is unknown for wild birds. Here we explored the timing of hatch order in a free-living population of hihi nestlings, Notiomystis cincta, and followed any effects in and out of the nest. We found that while hatching time from first- to last-hatched nestlings was often less than 24 h, last-hatched individuals grew more slowly and were lighter and smaller at fledging than older siblings. Last-hatched nestlings were also less likely to fledge. These effects were greater in larger broods. Adult body size is correlated with fledging size in hihi; however, we found no evidence that hatch order affected longevity postfledging, or lifetime reproductive success. We then explored whether carotenoid availability might buffer these stressful rearing conditions (through food supplementation of parents) but found no evidence that increased access to carotenoids for mothers and/or growing nestlings influenced incubation schedules, or the effects of hatching late. Together these results suggest that while even a very short hatch period can influence adult phenotype, hatching asynchrony is not maladaptive for hihi: when last-hatched nestlings survive to fledge they can contribute as much to their mothers' fitness as first-hatched siblings

Defences against brood parasites from a social immunity perspective

Parasitic interactions are so ubiquitous that all multicellular organisms have evolved a system of defences to reduce their costs, whether the parasites they encounter are the “classic parasites” that feed on the individual, or “brood parasites” that usurp parental care. Many parallels have been drawn between defences deployed against both types of parasite, but typically, whilst defences against classic parasites have been selected to protect survival, those against brood parasites have been selected to protect the parent’s inclusive fitness, suggesting that the selection pressures they impose are fundamentally different. However, there is another class of defences against classic parasites that have specifically been selected to protect an individual’s inclusive fitness, known as “social immunity”. Social immune responses include the anti-parasite defences typically provided for others in kin-structured groups, such as the antifungal secretions produced by termite workers to protect the brood. Defences against brood parasites, therefore, are more closely aligned with social immune responses. Much like social immunity, host defences against brood parasitism are employed by a donor (a parent) for the benefit of one or more recipients (typically kin), and as with social defences against classic parasites, defences have therefore evolved to protect the donor’s inclusive fitness, not the survival or ultimately the fitness of individual recipients This can lead to severe conflicts between the different parties, whose interests are not always aligned. Here we consider defences against brood parasitism in the light of social immunity, at different stages of parasite encounter, addressing where conflicts occur and how they might be resolved. We finish with considering how this approach could help us to address longstanding questions in our understanding of brood parasitism.Peer reviewe

Darwin's small and medium ground finches might have taste preferences, but not for human foods

Urbanization is rapidly changing ecological niches. On the inhabited Galapagos Islands, Darwin's finches consume human-introduced foods preferentially; however, it remains unclear why. Here, we presented pastry with flavour profiles typical of human foods (oily, salty and sweet) to small ground finches (Geospiza fuliginosa) and medium ground finches (Geospiza fortis) to test if latent taste preferences might drive the selection of human foods. If human food flavours were consumed more than a neutral or bitter control only at sites with human foods, then we predicted tastes were acquired after urbanization; however, if no site differences were found then this would indicate latent taste preferences. Contrary to both predictions, we found little evidence that human food flavours were preferred compared with control flavours at any site. Instead, finches showed a weak aversion to oily foods, but only at remote (no human foods present) sites. This was further supported by behavioural responses, with beak-wiping occurring more often at remote sites after finches tasted flavours associated with human foods. Our results suggest, therefore, that while Darwin's finches regularly exposed to human foods might have acquired a tolerance to human food flavours, latent taste preferences are unlikely to have played a major role in their dietary response to increased urbanization.Peer reviewe

Reed Warbler Hosts Do Not Fine-Tune Mobbing Defenses During the Breeding Season, Even When Cuckoos Are Rare

Hosts of brood parasitic cuckoos often employ mobbing attacks to defend their nests and, when mobbing is costly, hosts are predicted to adjust their mobbing to match parasitism risk. While evidence exists for fine-tuned plasticity, it remains unclear why mobbing does not track larger seasonal changes in parasitism risk. Here we test a possible explanation from parental investment theory: parents should defend their current brood more intensively as the opportunity to replace it declines (re-nesting potential), and therefore “counteract” any apparent seasonal decline to match parasitism risk. We take advantage of mobbing experiments conducted at two sites where reed warblers (Acrocephalus scirpaceus) experience (in Italy), or do not experience (in Finland), brood parasitism. We predicted that mobbing of cuckoos should be higher overall in Italy, but remain constant over the season as in other parasitised sites, whereas in Finland where cuckoos do not pose a local threat, we predicted that mobbing should be low at the beginning of the season but increase as re-nesting potential declined. However, while cuckoos were more likely to be mobbed in Italy, we found little evidence that mobbing changed over the season at either the parasitized or non-parasitized sites. This suggests that re-nesting potential has either little influence on mobbing behavior, or that its effects are obscured by other seasonal differences in ecology or experience of hosts

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.Peer reviewe

Cancer incidence in British vegetarians

Background: Few prospective studies have examined cancer incidence among vegetarians. Methods: We studied 61 566 British men and women, comprising 32 403 meat eaters, 8562 non-meat eaters who did eat fish ('fish eaters') and 20 601 vegetarians. After an average follow-up of 12.2 years, there were 3350 incident cancers of which 2204 were among meat eaters, 317 among fish eaters and 829 among vegetarians. Relative risks (RRs) were estimated by Cox regression, stratified by sex and recruitment protocol and adjusted for age, smoking, alcohol, body mass index, physical activity level and, for women only, parity and oral contraceptive use. Results: There was significant heterogeneity in cancer risk between groups for the following four cancer sites: stomach cancer, RRs (compared with meat eaters) of 0.29 (95% CI: 0.07–1.20) in fish eaters and 0.36 (0.16–0.78) in vegetarians, P for heterogeneity=0.007; ovarian cancer, RRs of 0.37 (0.18–0.77) in fish eaters and 0.69 (0.45–1.07) in vegetarians, P for heterogeneity=0.007; bladder cancer, RRs of 0.81 (0.36–1.81) in fish eaters and 0.47 (0.25–0.89) in vegetarians, P for heterogeneity=0.05; and cancers of the lymphatic and haematopoietic tissues, RRs of 0.85 (0.56–1.29) in fish eaters and 0.55 (0.39–0.78) in vegetarians, P for heterogeneity=0.002. The RRs for all malignant neoplasms were 0.82 (0.73–0.93) in fish eaters and 0.88 (0.81–0.96) in vegetarians (P for heterogeneity=0.001). Conclusion: The incidence of some cancers may be lower in fish eaters and vegetarians than in meat eaters

Analysing age structure, residency and relatedness uncovers social network structure in aggregations of young birds

Animal sociality arises from the cumulative effects of both individual social decisions and environmental factors. While juveniles' social interactions with parents prior to independence shape later life sociality, in most bird and mammal species at least one sex undergoes an early life dispersal before first-year reproduction. The social associations from this period could also have implications for later life yet are rarely characterized. Here, we derived predictions from available examples of juvenile groups in the literature (mobile ‘flocks’, spatially stable ‘gangs’ or adult-associated ‘crèches’) and then used three cohorts of juvenile hihi, Notiomystis cincta, a threatened New Zealand passerine, to demonstrate how multistate modelling and social network analysis approaches can be used to characterize group type based on residency, movement, relatedness and social associations. At sites where hihi congregated, we found that juveniles were resighted at a higher frequency than adults and associated predominantly with unrelated juveniles rather than siblings or parents. Movement between group sites occurred, but associations developed predominantly within the sites. We suggest therefore that juvenile hihi social structure is most similar to a ‘gang’, a group structure in which juveniles congregate without adults at predictable sites. Such gangs have previously only been described formally in ravens, Corvus corax. By combining spatial and social network analyses, our study demonstrates how social group structures can be described and therefore facilitate broader comparisons and discussion about the form and function of juvenile groups across taxa