Factors influencing mercury exposure in Arctic-breeding shorebirds.

Mercury (Hg) pollution remains a concern to Arctic ecosystems, due to long-range transport from southern industrial regions and melting permafrost and glaciers. The objective of this study was to identify intrinsic, extrinsic, and temporal factors influencing Hg concentrations in Arctic-breeding shorebirds and highlight regions and species at greatest risk of Hg exposure. We analyzed 1094 blood and 1384 feather samples from 12 shorebird species breeding at nine sites across the North American Arctic during 2012 and 2013. Blood Hg concentrations, which reflect Hg exposure in the local area in individual shorebirds: 1) ranged from 0.01-3.52 µg/g ww, with an overall mean of 0.30 ± 0.27 µg/g ww; 2) were influenced by species and study site, but not sampling year, with birds sampled near Utqiagvik, AK, having the highest concentrations; and 3) were influenced by foraging habitat at some sites. Feather Hg concentrations, which reflected Hg exposure from the wintering grounds: 1) ranged from 0.07-12.14 µg/g fw in individuals, with an overall mean of 1.14 ± 1.18 µg/g fw; and 2) were influenced by species and year. Most Arctic-breeding shorebirds had blood and feather Hg concentrations at levels where no adverse effects of exposure were predicted, though some individuals sampled near Utqiagvik had Hg levels that would be considered of concern. Overall, these data increase our understanding of how Hg is distributed in the various shorebird breeding areas of the Arctic, what factors predispose Arctic-breeding shorebirds to Hg exposure, and lay the foundation for future monitoring efforts.

Behavioural responses of breeding arctic sandpipers to ground-surface temperature and primary productivity.

Most birds incubate their eggs, which requires time and energy at the expense of other activities. Birds generally have two incubation strategies: biparental where both mates cooperate in incubating eggs, and uniparental where a single parent incubates. In harsh and unpredictable environments, incubation is challenging due to high energetic demands and variable resource availability. We studied the relationships between the incubation behaviour of sandpipers (genus Calidris) and two environmental variables: temperature and a proxy of primary productivity (i.e. NDVI). We investigated how these relationships vary between incubation strategies and across species among strategies. We also studied how the relationship between current temperature and incubation behaviour varies with previous day's temperature. We monitored the incubation behaviour of nine sandpiper species using thermologgers at 15 arctic sites between 2016 and 2019. We also used thermologgers to record the ground surface temperature at conspecific nest sites and extracted NDVI values from a remote sensing product. We found no relationship between either environmental variables and biparental incubation behaviour. Conversely, as ground-surface temperature increased, uniparental species decreased total duration of recesses (TDR) and mean duration of recesses (MDR), but increased number of recesses (NR). Moreover, small species showed stronger relationships with ground-surface temperature than large species. When all uniparental species were combined, an increase in NDVI was correlated with higher mean duration, total duration and number of recesses, but relationships varied widely across species. Finally, some uniparental species showed a lag effect with a higher nest attentiveness after a warm day while more recesses occurred after a cold day than was predicted based on current temperatures. We demonstrate the complex interplay between shorebird incubation strategies, incubation behaviour, and environmental conditions. Understanding how species respond to changes in their environment during incubation helps predict their future reproductive success.

Correction to: Documenting lemming population change in the Arctic: Can we detect trends?

In the original published article, some of the symbols in figure 1A were modified incorrectly during the typesetting and publication process. The correct version of the figure is provided in this correction.

Documenting lemming population change in the Arctic: Can we detect trends?

Lemmings are a key component of tundra food webs and changes in their dynamics can affect the whole ecosystem. We present a comprehensive overview of lemming monitoring and research activities, and assess recent trends in lemming abundance across the circumpolar Arctic. Since 2000, lemmings have been monitored at 49 sites of which 38 are still active. The sites were not evenly distributed with notably Russia and high Arctic Canada underrepresented. Abundance was monitored at all sites, but methods and levels of precision varied greatly. Other important attributes such as health, genetic diversity and potential drivers of population change, were often not monitored. There was no evidence that lemming populations were decreasing in general, although a negative trend was detected for low arctic populations sympatric with voles. To keep the pace of arctic change, we recommend maintaining long-term programmes while harmonizing methods, improving spatial coverage and integrating an ecosystem perspective.

Spatio-temporal responses of predators to hyperabundant geese affect risk of predation for sympatric-nesting species.

The Arctic is undergoing rapid changes, with anthropogenic shifts in climate having important and well-documented impacts on habitat. Populations of predators and their prey are affected by changing climate and other anthropogenic factors, and these changing trophic interactions could have profound effects on breeding populations of Arctic birds. Variable abundance of lemmings (a primary prey of generalist Arctic predators) and increasing abundance of light geese (Lesser Snow and Ross' Geese; a secondary prey) could have negative consequences for numerous sympatric shorebirds (an incidental prey). Using 16 years of predator-prey observations and 13-years of shorebird nest survival data at a site near a goose colony we identify relationships among geese, lemmings, and their shared predators and then relate predator indices to shorebird risk of nest predation. During two years, we also placed time-lapse cameras and artificial shorebird nests at increasing distances from a goose colony to document spatial trends in predators and their effect on risk of predation. In the long-term data, yearly indices of light geese positively influenced indices of gulls and jaegers, and shorebird nest predation rate was negatively correlated with jaeger and fox indices. All three predator indices were highest near the goose colony and artificial nest predation probability was negatively correlated with distance from goose colony, but these effects were less apparent during the second year. Combined, these results highlight the variation in predator-mediated interactions between geese and shorebirds and outline one mechanism by which hyperabundant geese may be contributing to local or regional declines in Arctic-nesting shorebird populations.

Comment on "Global pattern of nest predation is disrupted by climate change in shorebirds".

Kubelka et al (Reports, 9 November 2018, p. 680) claim that climate change has disrupted patterns of nest predation in shorebirds. They report that predation rates have increased since the 1950s, especially in the Arctic. We describe methodological problems with their analyses and argue that there is no solid statistical support for their claims.

The strength of ecological subsidies across ecosystems: a latitudinal gradient of direct and indirect impacts on food webs.

Material and energy flows among ecosystems can directly and indirectly drive ecosystem functions. Yet, how populations of consumers respond to allochthonous inputs at a macroecological scale is still unclear. Using a meta-analysis spanning several biomes, we show that the abundance of recipient populations is 36-57% larger with increased allochthonous inputs. The strength of direct effects on the recipients of these inputs as well as the indirect effects on the consumers of these recipients (i.e. ascending indirect effects) are constant across a latitudinal gradient spanning subtropical, arid, temperate, boreal and arctic ecosystems. However, indirect effect on the in situ resources of the input recipient (i.e. descending indirect effects) decreases with latitude. Our results suggest that the influence of allochthonous inputs can vary across large-scale gradients of ecosystem productivity and may be driven by the types of trophic interactions within recipient food webs.

Unexpected diversity in socially synchronized rhythms of shorebirds.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent's incubation bout varied from 1-19 h, whereas period length-the time in which a parent's probability to incubate cycles once between its highest and lowest value-varied from 6-43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light-dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

Is it safe to nest near conspicuous neighbours? Spatial patterns in predation risk associated with the density of American Golden-Plover nests.

Predation is one of the main factors explaining nesting mortality in most bird species. Birds can avoid nest predation or reduce predation pressure by breeding at higher latitude, showing anti-predator behaviour, selecting nest sites protected from predators, and nesting in association with protective species. American Golden-Plovers (Pluvialis dominica) defend their territory by using various warning and distraction behaviours displayed at varying levels of intensity (hereafter "conspicuous behaviour"), as well as more aggressive behaviours such as aerial attacks, but only in some populations. Such antipredator behaviour has the potential to repel predators and thus benefit the neighbouring nests by decreasing their predation risk. Yet, conspicuous behaviour could also attract predators by signalling the presence of a nest. To test for the existence of a protective effect associated with the conspicuous antipredator behaviour of American Golden-Plovers, we studied the influence of proximity to plover nests on predation risk of artificial nests on Igloolik Island (Nunavut, Canada) in July 2014. We predicted that the predation risk of artificial nests would decrease with proximity to and density of plover nests. We monitored 18 plover nests and set 35 artificial nests at 30, 50, 100, 200, and 500 m from seven of those plover nests. We found that the predation risk of artificial nests increases with the density of active plover nests. We also found a significant negative effect of the distance to the nearest active protector nest on predation risk of artificial nests. Understanding how the composition and structure of shorebird communities generate spatial patterns in predation risks represents a key step to better understand the importance of these species of conservation concern in tundra food webs.

Sexing a sex-role-reversed species based on plumage: potential challenges in the red phalarope.

Sex-role reversal, in which males care for offspring, can occur when mate competition is stronger between females than males. Secondary sex traits and mate attracting displays in sex-role-reversed species are usually more pronounced in females than in males. The red phalarope (Phalaropus fulicarius) is a textbook example of a sex-role-reversed species. It is generally agreed that males are responsible for all incubation and parental care duties, whereas females typically desert males after having completed a clutch and may pair with new males to lay additional clutches. The breeding plumage of female red phalaropes is usually more brightly colored than male plumage, a reversed sexual dichromatism usually associated with sex-role reversal. Here, we confirm with PCR-based sexing that male red phalaropes can exhibit both the red body plumage typical of a female and the incubation behavior typical of a male. Our result, combined with previous observations of brightly colored red phalaropes incubating nests at the same arctic location (Igloolik Island, Nunavut, Canada), suggests that plumage dichromatism alone may not be sufficient to distinguish males from females in this breeding population of red phalaropes. This stresses the need for more systematic genetic sexing combined with standardized description of intersexual differences in red phalarope plumages. Determining whether such female-like plumage on males is a result of phenotypic plasticity or genetic variation could contribute to further understanding sex-role reversal strategies in the short Arctic summer.

Effects of geolocators on hatching success, return rates, breeding movements, and change in body mass in 16 species of Arctic-breeding shorebirds.

BACKGROUND: Geolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8-2.0 g total, representing 0.1-3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2-4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26-1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables. RESULTS: We detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5-5.8 % of body mass than if tags were 0.3-2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important. CONCLUSIONS: Negative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species.

Consumption of Buglossoides arvensis seed oil is safe and increases tissue long-chain n-3 fatty acid content more than flax seed oil - results of a phase I randomised clinical trial.

Enrichment of tissues with ≥20-carbon n-3 PUFA like EPA is associated with positive cardiovascular outcomes. Stearidonic acid (SDA; 18 : 4n-3) and α-linolenic acid (ALA; 18 : 3n-3) are plant-derived dietary n-3 PUFA; however, direct comparisons of their impact on tissue n-3 PUFA content are lacking. Ahiflower(®) oil extracted from Buglossoides arvensis seeds is the richest known non-genetically modified source of dietary SDA. To investigate the safety and efficacy of dietary Ahiflower oil, a parallel-group, randomised, double-blind, comparator-controlled phase I clinical trial was performed. Diets of healthy subjects (n 40) were supplemented for 28 d with 9·1 g/d of Ahiflower (46 % ALA, 20 % SDA) or flax seed oil (59 % ALA). Blood and urine chemistries, blood lipid profiles, hepatic and renal function tests and haematology were measured as safety parameters. The fatty acid composition of fasting plasma, erythrocytes, polymorphonuclear cells and mononuclear cells were measured at baseline and after 14 and 28 d of supplementation. No clinically significant changes in safety parameters were measured in either group. Tissue ALA and EPA content increased in both groups compared with baseline, but EPA accrual in plasma and in all cell types was greater in the Ahiflower group (time × treatment interactions, P ≤ 0·01). Plasma and mononuclear cell eicosatetraenoic acid (20 : 4n-3) and docosapentaenoic acid (22 : 5n-3) content also increased significantly in the Ahiflower group compared with the flax group. In conclusion, the consumption of Ahiflower oil is safe and is more effective for the enrichment of tissues with 20- and 22-carbon n-3 PUFA than flax seed oil.

Isotopic Differences between Forage Consumed by a Large Herbivore in Open, Closed, and Coastal Habitats: New Evidence from a Boreal Study System.

Documenting habitat-related patterns in foraging behaviour at the individual level and over large temporal scales remains challenging for large herbivores. Stable isotope analysis could represent a valuable tool to quantify habitat-related foraging behaviour at the scale of individuals and over large temporal scales in forest dwelling large herbivores living in coastal environments, because the carbon (δ13C) or nitrogen (δ15N) isotopic signatures of forage can differ between open and closed habitats or between terrestrial and littoral forage, respectively. Here, we examined if we could detect isotopic differences between the different assemblages of forage taxa consumed by white-tailed deer that can be found in open, closed, supralittoral, and littoral habitats. We showed that δ13C of assemblages of forage taxa were 3.0 ‰ lower in closed than in open habitats, while δ15N were 2.0 ‰ and 7.4 ‰ higher in supralittoral and littoral habitats, respectively, than in terrestrial habitats. Stable isotope analysis may represent an additional technique for ecologists interested in quantifiying the consumption of terrestrial vs. marine autotrophs. Yet, given the relative isotopic proximity and the overlap between forage from open, closed, and supralittoral habitats, the next step would be to determine the potential to estimate their contribution to herbivore diet.

Forage-mediated density and climate effects on body mass in a temperate herbivore: a mechanistic approach.

The interplay between density and climate in shaping the dynamics of herbivore populations is widely acknowledged, and current research is fueled by the identification of mechanisms underlying their effects on individuals and populations. We assessed whether forage availability mediated the effects of density and winter climate on body mass of white-tailed deer (Odocoileus virginianus) yearlings by experimentally reducing deer density to 7.5 and 15 deer/km2 during eight growing seasons, and by using causal (graphical) hierarchical models and Bayesian hierarchical modeling to assess relationships. The abundance of preferred forage decreased with deer density and varied quadratically (positive parabola) with winter North Atlantic Oscillation (NAO), whereas the fall mass of yearlings increased with forage abundance and spring mass. Fall mass did not differ between experimentally reduced deer densities, yet experimental yearlings were 30% heavier than yearlings harvested at ambient densities. Hence, forage abundance simultaneously mediated the effects of density and climate on fall body mass, which was also influenced by carry-over effects of spring body mass. Our findings increase our ability to anticipate how temperate large herbivores will respond to ongoing changes in intrinsic (e.g., large-herbivore density) and extrinsic (e.g., climate) factors.

A new way of assessing foraging behaviour at the individual level using faeces marking and satellite telemetry.

Heterogeneity in foraging behaviour can profoundly influence ecological processes shaping populations. To scale-up from individual foraging behaviour to processes occurring at the population scale, one needs to sample foraging behaviour at the individual level, and over large temporal scales or during critical seasons known to influence life-history traits. We developed an innovative technique to monitor foraging behaviour at the individual level in secretive species, a technique that can be ultimately used to investigate the links between foraging behaviour and life-history traits. First, the technique used a novel approach, namely the combination of telemetry tracking and biomarking of faeces with food dyes to locate fresh signs of presence left by individuals equipped with GPS collars. Second, the technique is based on the simultaneous or successive sampling of life-history traits and individual foraging behaviour, using tracks with high probabilities of recovery of dyed faeces. We first describe our methodological approach, using a case study of a large herbivore, and then provide recommendations and guidelines for its use. Sampling single snow tracks of individuals equipped with a GPS collar was a reliable way to assess individual winter foraging behaviour in a white-tailed deer (Odocoileus virginianus Zimmermann) population. During that period, the probability of recovery of dyed faeces within the range of the collar precision was very high for single snow tracks of equipped deer (97%). Our approach is well suited to study individual foraging behaviour, and could ultimately be used to investigate the interplay between intra-population heterogeneity in foraging behaviour, life-history traits, and demographic processes.

Benefiting from a migratory prey: spatio-temporal patterns in allochthonous subsidization of an Arctic predator.

1. Flows of nutrients and energy across ecosystem boundaries have the potential to subsidize consumer populations and modify the dynamics of food webs, but how spatio-temporal variations in autochthonous and allochthonous resources affect consumers' subsidization remains largely unexplored. 2. We studied spatio-temporal patterns in the allochthonous subsidization of a predator living in a relatively simple ecosystem. We worked on Bylot Island (Nunavut, Canada), where arctic foxes (Vulpes lagopus L.) feed preferentially on lemmings (Lemmus trimucronatus and Dicrostonyx groenlandicus Traill), and alternatively on colonial greater snow geese (Anser caerulescens atlanticus L.). Geese migrate annually from their wintering grounds (where they feed on farmlands and marshes) to the Canadian Arctic, thus generating a strong flow of nutrients and energy across ecosystem boundaries. 3. We examined the influence of spatial variations in availability of geese on the diet of fox cubs (2003-2005) and on fox reproductive output (1996-2005) during different phases of the lemming cycle. 4. Using stable isotope analysis and a simple statistical routine developed to analyse the outputs of a multisource mixing model (SIAR), we showed that the contribution of geese to the diet of arctic fox cubs decreased with distance from the goose colony. 5. The probability that a den was used for reproduction by foxes decreased with distance from the subsidized goose colony and increased with lemming abundance. When lemmings were highly abundant, the effect of distance from the colony disappeared. The goose colony thus generated a spatial patterning of reproduction probability of foxes, while the lemming cycle generated a strong temporal variation of reproduction probability of foxes. 6. This study shows how the input of energy owing to the large-scale migration of prey affects the functional and reproductive responses of an opportunistic consumer, and how this input is spatially and temporally modulated through the foraging behaviour of the consumer. Thus, perspectives of both landscape and foraging ecology are needed to fully resolve the effects of subsidies on animal demographic processes and population dynamics.