An interspecific foraging association with polar bears increases foraging opportunities for avian predators in a declining Arctic seabird colony.

Interspecific foraging associations (IFAs) are biological interactions where two or more species forage in association with each other. Climate-induced reductions in Arctic sea ice have increased polar bear (Ursus maritimus) foraging in seabird colonies, which creates foraging opportunities for avian predators. We used drone video of bears foraging within a common eider (Somateria mollissima) colony on East Bay Island (Nunavut, Canada) in 2017 to investigate herring gull (Larus argentatus) foraging in association with bears. We recorded nest visitation by gulls following n = 193 eider flushing events from nests during incubation. The probability of gulls visiting eider nests increased with higher number of gulls present (ß = 0.14 ± 0.03 [SE], p < .001) and for nests previously visited by a bear (ß = 1.14 ± 0.49 [SE], p < .02). In our model examining the probability of gulls consuming eggs from nests, we failed to detect statistically significant effects for the number of gulls present (ß = 0.09 ± 0.05 [SE], p < .07) or for nests previously visited by a bear (ß = -0.92 ± 0.71 [SE], p < .19). Gulls preferred to visit nests behind bears (χ2 = 18, df = 1, p < .0001), indicating gulls are risk averse in the presence of polar bears. Our study provides novel insights on an Arctic IFA, and we present evidence that gulls capitalize on nests made available due to disturbance associated with foraging bears, as eiders disturbed off their nest allow gulls easier access to eggs. We suggest the IFA between gulls and polar bears is parasitic, as gulls are consuming terrestrial resources which would have eventually been consumed by bears. This finding has implications for estimating the energetic contribution of bird eggs to polar bear summer diets in that the total number of available clutches to consume may be reduced due to avian predators.

Co-contaminants of microplastics in two seabird species from the Canadian Arctic.

Through ingestion and subsequent egestion, Arctic seabirds can bioaccumulate microplastics at and around their colony breeding sites. While microplastics in Arctic seabirds have been well documented, it is not yet understood to what extent these particles can act as transport vehicles for plastic-associated contaminants, including legacy persistent organic pollutants (POPs), trace metals, and organic additives. We investigated the occurrence and pattern of organic and inorganic co-contaminants of microplastics in two seabird species from the Canadian Arctic - northern fulmar (Fulmarus glacialis) and black-legged kittiwake (Rissa tridactyla). We found that fulmars had higher levels of plastic contamination and emerging organic compounds (known to be plastic additives) than kittiwakes, whereas higher concentrations of legacy POPs were found in kittiwakes than the fulmars. Furthermore, fulmars, the species with the much larger foraging range (∼200 km), had higher plastic pollution and overall contaminant burdens, indicating that birds may be acting as long-range transport vectors for plastic-associated pollution. Our results suggest a potential connection between plastic additive contamination and plastic pollution burdens in the bird stomachs, highlighting the importance of treating plastic particles and plastic-associated organic additives as co-contaminants rather than separate pollution issues.

Social status modulates the behavioral and physiological consequences of a chemical pollutant in animal groups.

The social environment (i.e., the suite of social interactions that occur among individuals that can result in variation in social ranks) is a commonly overlooked aspect of biology when scientists evaluate the effects of chemical contaminants. The social environment, however, represents the arena in which individual-level performance shapes group- or population-level outcomes and may therefore mediate many of the ultimate consequences of chemicals for wildlife. Here, we evaluated the role that the social environment plays in determining the consequences of pollutant exposure. We exposed groups of juvenile brown trout (Salmo trutta) to an emerging pharmaceutical pollutant that is commonly detected in freshwaters (the benzodiazepine, oxazepam) and allowed them to form dominance hierarchies. Exposure affected dominant and subordinate fish differently, causing fish to become less aggressive at high doses and subordinate fish to become more competitively successful at low doses. These perturbations had further consequences for growth, fin damage, and survival. Exposure also modulated physiological stress in the hierarchy, and social status itself affected how much oxazepam was absorbed in tissues, potentially creating a dynamic feedback loop that further influences the asymmetric effects of exposure on differing social statuses. Many effects followed a "U-shaped" dose-response curve, highlighting the importance of nonlinear, low-dose effects. Altogether, we show that social structure in animal groups can interact with and modulate the effects of an environmental contaminant. We underscore the need to account for an organism's natural ecological context, including their social environment, in future experiments and environmental risk assessments to predict the effects of chemical contaminants on wildlife.

Polar bears are inefficient predators of seabird eggs.

Climate-mediated sea-ice loss is disrupting the foraging ecology of polar bears (Ursus maritimus) across much of their range. As a result, there have been increased reports of polar bears foraging on seabird eggs across parts of their range. Given that polar bears have evolved to hunt seals on ice, they may not be efficient predators of seabird eggs. We investigated polar bears' foraging performance on common eider (Somateria mollissima) eggs on Mitivik Island, Nunavut, Canada to test whether bear decision-making heuristics are consistent with expectations of optimal foraging theory. Using aerial-drones, we recorded multiple foraging bouts over 11 days, and found that as clutches were depleted to completion, bears did not exhibit foraging behaviours matched to resource density. As the season progressed, bears visited fewer nests overall, but marginally increased their visitation to nests that were already empty. Bears did not display different movement modes related to nest density, but became less selective in their choice of clutches to consume. Lastly, bears that capitalized on visual cues of flushing eider hens significantly increased the number of clutches they consumed; however, they did not use this strategy consistently or universally. The foraging behaviours exhibited by polar bears in this study suggest they are inefficient predators of seabird eggs, particularly in the context of matching behaviours to resource density.

The consequences of null model selection for predicting mortality from multiple stressors.

Many ecological systems are now exposed to multiple stressors, and ecosystem management increasingly requires consideration of the joint effects of multiple stressors on focal populations, communities and ecosystems. In the absence of empirical data, ecosystem managers could use null models based on the combination of independently acting stressors to estimate the joint effects of multiple stressors. Here, we used a simulation study and a meta-analysis to explore the consequences of null model selection for the prediction of mortality resulting from exposure to two stressors. Comparing five existing null models, we show that some null models systematically predict lower mortality rates than others, with predicted mortality rates up to 67.5% higher or 50% lower than the commonly used Simple Addition model. However, the null model predicting the highest mortality rate differed across parameter sets, and therefore there is no general 'precautionary null model' for multiple stressors. Using a multi-model framework, we re-analysed data from two earlier meta-analyses and found that 54% of the observed joint effects fell within the range of predictions from the suite of null models. Furthermore, we found that most null models systematically underestimated the observed joint effects, with only the Stressor Addition model showing a bias for overestimation. Finally, we found that the intensity of individual stressors was the strongest predictor of the magnitude of the joint effect across all null models. As a result, studies characterizing the effects of individuals stressors are still required for accurate prediction of mortality resulting from multiple stressors.

A review and meta-analysis of collaborative research prioritization studies in ecology, biodiversity conservation and environmental science.

Collaborative research prioritization (CRP) studies have become increasingly popular during the last decade. By bringing together a diverse group of stakeholders, and using a democratic process to create a list of research priorities, these methods purport to identify research topics that will better meet the needs of science users. Here, we review 41 CRP studies in the fields of ecology, biodiversity conservation and environmental science that collectively identify 2031 research priorities. We demonstrate that climate change, ecosystem services and protected areas are common terms found in the research priorities of many CRP studies, and that identified research priorities have become less unique over time. In addition, we show that there is a considerable variation in the size and composition of the groups involved in CRP studies, and that at least one aspect of the identified research priorities (lexical diversity) is related to the size of the CRP group. Although some CRP studies have been highly cited, the evidence that CRP studies have directly motivated research is weak, perhaps because most CRP studies have not directly involved organizations that fund science. We suggest that the most important impact of CRP studies may lie in their ability to connect individuals across sectors and help to build diverse communities of practice around important issues at the science-policy interface.

Evolutionary drivers of seasonal plumage colours: colour change by moult correlates with sexual selection, predation risk and seasonality across passerines.

Some birds undergo seasonal colour change by moulting twice each year, typically alternating between a cryptic, non-breeding plumage and a conspicuous, breeding plumage ('seasonal plumage colours'). We test for potential drivers of the evolution of seasonal plumage colours in all passerines (N = 5901 species, c. 60% of all birds). Seasonal plumage colours are uncommon, having appeared on multiple occasions but more frequently lost during evolution. The trait is more common in small, ground-foraging species with polygynous mating systems, no paternal care and strong sexual dichromatism, suggesting it evolved under strong sexual selection and high predation risk. Seasonal plumage colours are also more common in species predicted to have seasonal breeding schedules, such as migratory birds and those living in seasonal climates. We propose that seasonal plumage colours have evolved to resolve a trade-off between the effects of natural and sexual selection on colouration, especially in seasonal environments.

Author Correction: Direct benefits and evolutionary transitions to complex societies.

In the version of this Article originally published, references were missing from the column "Source(s) for mating and parental care system data" in Supplementary Table 1. The following references have now been added to the relevant species: Brichard 1989 has been added to Chalinochromis popelini, Chalinochromis brichardi and Julidochromis dickfeldi; Clabaut et al. 2007 to Altolamprologus calvus and Julidochromis regani; Konings 1998 to Neolamprologus hecqui; and Kuwamura 1997 to Chalinochromis popelini.

Direct benefits and evolutionary transitions to complex societies.

The selective forces that drive the evolution of cooperation have been intensely debated. Evolutionary transitions to cooperative breeding, a complex form of cooperation, have been hypothesized to be linked to low degrees of promiscuity, which increases intragroup relatedness and the indirect (that is, kin selected) benefits of helping. However, ecological factors also promote cooperative breeding, and may be more important than relatedness in some contexts. Identifying the key evolutionary drivers of cooperative breeding therefore requires an integrated assessment of these hypotheses. Here we show, using a phylogenetic framework that explicitly evaluates mating behaviours and ecological factors, that evolutionary transitions to cooperative breeding in cichlid fishes were not associated with social monogamy. Instead, group living, biparental care and diet type directly favoured the evolution of cooperative breeding. Our results suggest that cichlid fishes exhibit an alternative path to the evolution of complex societies compared to other previously studied vertebrates, and these transitions are driven primarily by direct fitness benefits.

Direct benefits and evolutionary transitions to complex societies.

The selective forces that drive the evolution of cooperation have been intensely debated. Evolutionary transitions to cooperative breeding, a complex form of cooperation, have been hypothesized to be linked to low degrees of promiscuity, which increases intragroup relatedness and the indirect (that is, kin selected) benefits of helping. However, ecological factors also promote cooperative breeding, and may be more important than relatedness in some contexts. Identifying the key evolutionary drivers of cooperative breeding therefore requires an integrated assessment of these hypotheses. Here we show, using a phylogenetic framework that explicitly evaluates mating behaviours and ecological factors, that evolutionary transitions to cooperative breeding in cichlid fishes were not associated with social monogamy. Instead, group living, biparental care and diet type directly favoured the evolution of cooperative breeding. Our results suggest that cichlid fishes exhibit an alternative path to the evolution of complex societies compared to other previously studied vertebrates, and these transitions are driven primarily by direct fitness benefits.

Increasing nest predation will be insufficient to maintain polar bear body condition in the face of sea ice loss.

Climate change can influence interspecific interactions by differentially affecting species-specific phenology. In seasonal ice environments, there is evidence that polar bear predation of Arctic bird eggs is increasing because of earlier sea ice breakup, which forces polar bears into nearshore terrestrial environments where Arctic birds are nesting. Because polar bears can consume a large number of nests before becoming satiated, and because they can swim between island colonies, they could have dramatic influences on seabird and sea duck reproductive success. However, it is unclear whether nest foraging can provide an energetic benefit to polar bear populations, especially given the capacity of bird populations to redistribute in response to increasing predation pressure. In this study, we develop a spatially explicit agent-based model of the predator-prey relationship between polar bears and common eiders, a common and culturally important bird species for northern peoples. Our model is composed of two types of agents (polar bear agents and common eider hen agents) whose movements and decision heuristics are based on species-specific bioenergetic and behavioral ecological principles, and are influenced by historical and extrapolated sea ice conditions. Our model reproduces empirical findings that polar bear predation of bird nests is increasing and predicts an accelerating relationship between advancing ice breakup dates and the number of nests depredated. Despite increases in nest predation, our model predicts that polar bear body condition during the ice-free period will continue to decline. Finally, our model predicts that common eider nests will become more dispersed and will move closer to the mainland in response to increasing predation, possibly increasing their exposure to land-based predators and influencing the livelihood of local people that collect eider eggs and down. These results show that predator-prey interactions can have nonlinear responses to changes in climate and provides important predictions of ecological change in Arctic ecosystems.

Effectiveness of baseline corticosterone as a monitoring tool for fitness: a meta-analysis in seabirds.

Many ecosystems have experienced anthropogenically induced changes in biodiversity, yet predicting these patterns has been difficult. Recently, individual behavioural and physiological measures have been proposed as more rapid links between environmental variation and fitness compared to demographics. Glucocorticoid hormones have received much attention given that they mediate energetic demands, metabolism, and foraging behaviour. However, it is currently unclear whether glucocorticoids can reliably predict environmental and fitness-related traits and whether they may be useful in specific groups of taxa. In particular, seabirds are a well-studied avian group often employed as biomonitoring tools for environmental change given their wide distribution and reliance on large oceanic patterns. Despite the increase in studies attempting to link variation in baseline corticosterone (the primary avian glucocorticoid) to variation in fitness-related traits in seabirds, there has been no comprehensive review of the relationship in this taxon. We present a phylogenetically controlled systematic review and meta-analysis of correlative and experimental studies examining baseline corticosterone as a predictor of fitness-related traits relevant to predicting seabird population health. Our results suggest that, while variation in baseline corticosterone may be a useful predictor of larger-scale environmental traits such as overall food availability and fitness-related traits such as reproductive success, this hormone may not be sensitive enough to detect variation in body condition, foraging effort, and breeding effort. Overall, our results support recent work suggesting that the use of baseline glucocorticoids as conservation biomarkers is complex and highly context dependent, and we suggest caution in their use and interpretation as simplified, direct biomarkers of fitness.

The effects of life history and sexual selection on male and female plumage colouration.

Classical sexual selection theory provides a well-supported conceptual framework for understanding the evolution and signalling function of male ornaments. It predicts that males obtain greater fitness benefits than females through multiple mating because sperm are cheaper to produce than eggs. Sexual selection should therefore lead to the evolution of male-biased secondary sexual characters. However, females of many species are also highly ornamented. The view that this is due to a correlated genetic response to selection on males was widely accepted as an explanation for female ornamentation for over 100 years and current theoretical and empirical evidence suggests that genetic constraints can limit sex-specific trait evolution. Alternatively, female ornamentation can be the outcome of direct selection for signalling needs. Since few studies have explored interspecific patterns of both male and female elaboration, our understanding of the evolution of animal ornamentation remains incomplete, especially over broad taxonomic scales. Here we use a new method to quantify plumage colour of all ~6,000 species of passerine birds to determine the main evolutionary drivers of ornamental colouration in both sexes. We found that conspecific male and female colour elaboration are strongly correlated, suggesting that evolutionary changes in one sex are constrained by changes in the other sex. Both sexes are more ornamented in larger species and in species living in tropical environments. Ornamentation in females (but not males) is increased in cooperative breeders--species in which female-female competition for reproductive opportunities and other resources related to breeding may be high. Finally, strong sexual selection on males has antagonistic effects, causing an increase in male colouration but a considerably more pronounced reduction in female ornamentation. Our results indicate that although there may be genetic constraints to sexually independent colour evolution, both female and male ornamentation are strongly and often differentially related to morphological, social and life-history variables.

Manipulating the appearance of a badge of status causes changes in true badge expression.

Signals of dominance and fighting ability (i.e. status signals) are found in a wide range of taxa and are used to settle disputes between competitive rivals. Most previous research has considered status-signal phenotype as an attribute of the individual; however, it is more likely that signal expression is an emergent property that also incorporates aspects of the social environment. Furthermore, because an individual's signal phenotype is likely to influence its social interactions, the relationships between status signals, social environment and individual quality are probably much more complex than previously appreciated. Here, we explore the dynamic relationship between social interactions and signal expression in a previously undescribed status signal, the frontal shield of the pukeko (Porphyrio porphyrio melanotus: Aves). We demonstrate that frontal shield size is a strong predictor of dominance status within social groups, even after controlling for potentially confounding variables. Then, we evaluate the relationship between social interactions and signal expression by testing whether manipulating apparent shield size influences (i) dominance interactions and (ii) future signal expression. By showing that decreasing apparent shield size causes both an increase in the amount of aggression received and a decrease in an individual's true shield size, we provide the first evidence of dynamic feedback between signal expression and social interactions. Our study provides important insight into the role of receiver-dependent (i.e. social) costs in maintaining signal honesty and demonstrates a unique approach to studying status signalling applicable to future studies on dynamic morphological signals.

Behavioral and physiological responses of a wild teleost fish to cortisol and androgen manipulation during parental care.

Proximate mediators of reproductive behaviors in vertebrates have a long history of study. In fishes, relatively few studies have focused on hormonal control of parental care, despite a comprehensive background on the general physiology of fishes, and the frequent occurrence of parental care behaviors. Studies on this taxon have repeatedly found that the relationships between androgens and paternal care do not follow the predictions made in the avian and mammalian literature. Glucocorticoids may also have a role in mediating parental behaviors, possibly through their role as regulators of metabolism. As such, we investigated the role of 11-ketotestosterone (11-KT) and cortisol in mediating parental effort of male smallmouth bass (Micropterus dolomieu) by manipulating hormone titers in wild fish. In smallmouth bass, males spawn annually with a single female and defend a single brood for up to 30 days. Treatment of parental fish with cyproterone acetate (CYA; an androgen receptor antagonist) resulted in a decrease in nest defense in response to a simulated brood predator; however, no changes in nest success, nest tending or biochemical indicators of nutritional status were detected. Treatment with exogenous cortisol did not change parental behavior, but did increase the rate of nest failure, possibly owing to the energetic cost of chronically elevated cortisol concentrations. We discuss these findings in the context of resource-driven trade-offs and highlight life history as an important factor controlling parental effort in species with costly parental care behaviors.