Spatial distribution of selenium-mercury in Arctic seabirds.

Mercury (Hg) is a metallic trace element toxic for humans and wildlife that can originate from natural and anthropic sources. Hg spatial gradients have been found in seabirds from the Arctic and other oceans, suggesting contrasting toxicity risks across regions. Selenium (Se) plays a protective role against Hg toxicity, but its spatial distribution has been much less investigated than that of Hg. From 2015 to 2017, we measured spatial co-exposure of Hg and Se in blood samples of two seabird species, the Brünnich's guillemot (Uria lomvia) and the black-legged kittiwake (Rissa tridactyla) from 17 colonies in the Arctic and subarctic regions, and we calculated their molar ratios (Se:Hg), as a measure of Hg sequestration by Se and, therefore, of Hg exposure risk. We also evaluated concentration differences between species and ocean basins (Pacific-Arctic and Atlantic-Arctic), and examined the influence of trophic ecology on Hg and Se concentrations using nitrogen and carbon stable isotopes. In the Atlantic-Arctic ocean, we found a negative west-to-east gradient of Hg and Se for guillemots, and a positive west-to-east gradient of Se for kittiwakes, suggesting that these species are better protected from Hg toxicity in the European Arctic. Differences in Se gradients between species suggest that they do not follow environmental Se spatial variations. This, together with the absence of a general pattern for isotopes influence on trace element concentrations, could be due to foraging ecology differences between species. In both oceans, the two species showed similar Hg concentrations, but guillemots showed lower Se concentrations and Se:Hg than kittiwakes, suggesting a higher Hg toxicity risk in guillemots. Within species, neither Hg, nor Se or Se:Hg differed between both oceans. Our study highlights the importance of considering Se together with Hg, along with different species and regions, when evaluating Hg toxic effects on marine predators in international monitoring programs.

Comparing non-breeding distribution and behavior of red-legged kittiwakes from two geographically distant colonies.

Knowledge of non-breeding distributions is a crucial component to seabird conservation, as conditions during the non-breeding period can play an important role in regulating seabird populations. Specifically, if seabirds from different colonies share the same wintering grounds, conditions in that shared region could have a widespread impact on multiple breeding populations. Red-legged kittiwakes (Rissa brevirostris) are endemic to the Bering Sea and may be especially susceptible to effects of climate change due to a restricted breeding range, small population size, and specialized diet. To examine whether red-legged kittiwakes from different breeding colonies overlapped in winter distribution and activity patterns, we used geolocation loggers to simultaneously track individuals from the two largest red-legged kittiwake breeding colonies in Alaska (separated by over 1000 km) during two consecutive non-breeding periods. We found that non-breeding activity patterns were generally similar between birds originating from the two colonies, but birds employed different migratory strategies during the early winter. Kittiwakes from Buldir Island in the western Aleutian Islands left the colony in September and immediately headed west, spending October through December around the Sea of Okhotsk and the Kuril Islands. In contrast, birds from St. George Island in the Pribilof Islands remained in the eastern Bering Sea or around the eastern Aleutian Islands for a couple months before traveling farther west. During late winter however, from January through March, birds from both colonies converged south of Kamchatka and east of the Kuril Islands over the Kuril-Kamchatka Trench and in the Western Subarctic Gyre before returning to their respective colonies in the spring. This late winter overlap in distributions along the Kuril-Kamchatka Trench suggests the region is a winter hotspot for red-legged kittiwakes and highlights the importance of this region for the global kittiwake population.

Seasonal variation of mercury contamination in Arctic seabirds: A pan-Arctic assessment.

Mercury (Hg) is a natural trace element found in high concentrations in top predators, including Arctic seabirds. Most current knowledge about Hg concentrations in Arctic seabirds relates to exposure during the summer breeding period when researchers can easily access seabirds at colonies. However, the few studies focused on winter have shown higher Hg concentrations during the non-breeding period than breeding period in several tissues. Hence, improving knowledge about Hg exposure during the non-breeding period is crucial to understanding the threats and risks encountered by these species year-round. We used feathers of nine migratory alcid species occurring at high latitudes to study bird Hg exposure during both the breeding and non-breeding periods. Overall, Hg concentrations during the non-breeding period were ~3 times higher than during the breeding period. In addition, spatial differences were apparent within and between the Atlantic and Pacific regions. While Hg concentrations during the non-breeding period were ~9 times and ~3 times higher than during the breeding period for the West and East Atlantic respectively, Hg concentrations in the Pacific during the non-breeding period were only ~1.7 times higher than during the breeding period. In addition, individual Hg concentrations during the non-breeding period for most of the seabird colonies were above 5 µg g-1 dry weight (dw), which is considered to be the threshold at which deleterious effects are observed, suggesting that some breeding populations might be vulnerable to non-breeding Hg exposure. Since wintering area locations, and migration routes may influence seasonal Hg concentrations, it is crucial to improve our knowledge about spatial ecotoxicology to fully understand the risks associated with Hg contamination in Arctic seabirds.

Extreme mortality and reproductive failure of common murres resulting from the northeast Pacific marine heatwave of 2014-2016.

About 62,000 dead or dying common murres (Uria aalge), the trophically dominant fish-eating seabird of the North Pacific, washed ashore between summer 2015 and spring 2016 on beaches from California to Alaska. Most birds were severely emaciated and, so far, no evidence for anything other than starvation was found to explain this mass mortality. Three-quarters of murres were found in the Gulf of Alaska and the remainder along the West Coast. Studies show that only a fraction of birds that die at sea typically wash ashore, and we estimate that total mortality approached 1 million birds. About two-thirds of murres killed were adults, a substantial blow to breeding populations. Additionally, 22 complete reproductive failures were observed at multiple colonies region-wide during (2015) and after (2016-2017) the mass mortality event. Die-offs and breeding failures occur sporadically in murres, but the magnitude, duration and spatial extent of this die-off, associated with multi-colony and multi-year reproductive failures, is unprecedented and astonishing. These events co-occurred with the most powerful marine heatwave on record that persisted through 2014-2016 and created an enormous volume of ocean water (the "Blob") from California to Alaska with temperatures that exceeded average by 2-3 standard deviations. Other studies indicate that this prolonged heatwave reduced phytoplankton biomass and restructured zooplankton communities in favor of lower-calorie species, while it simultaneously increased metabolically driven food demands of ectothermic forage fish. In response, forage fish quality and quantity diminished. Similarly, large ectothermic groundfish were thought to have increased their demand for forage fish, resulting in greater top-predator demands for diminished forage fish resources. We hypothesize that these bottom-up and top-down forces created an "ectothermic vise" on forage species leading to their system-wide scarcity and resulting in mass mortality of murres and many other fish, bird and mammal species in the region during 2014-2017.

Unusual mortality of Tufted puffins (Fratercula cirrhata) in the eastern Bering Sea.

Mass mortality events are increasing in frequency and magnitude, potentially linked with ongoing climate change. In October 2016 through January 2017, St. Paul Island, Bering Sea, Alaska, experienced a mortality event of alcids (family: Alcidae), with over 350 carcasses recovered. Almost three-quarters of the carcasses were unscavenged, a rate much higher than in baseline surveys (17%), suggesting ongoing deposition and elevated mortality around St Paul over a 2-3 month period. Based on the observation that carcasses were not observed on the neighboring island of St. George, we bounded the at-sea distribution of moribund birds, and estimated all species mortality at 3,150 to 8,800 birds. The event was particularly anomalous given the late fall/winter timing when low numbers of beached birds are typical. In addition, the predominance of Tufted puffins (Fratercula cirrhata, 79% of carcass finds) and Crested auklets (Aethia cristatella, 11% of carcass finds) was unusual, as these species are nearly absent from long-term baseline surveys. Collected specimens were severely emaciated, suggesting starvation as the ultimate cause of mortality. The majority (95%, N = 245) of Tufted puffins were adults regrowing flight feathers, indicating a potential contribution of molt stress. Immediately prior to this event, shifts in zooplankton community composition and in forage fish distribution and energy density were documented in the eastern Bering Sea following a period of elevated sea surface temperatures, evidence cumulatively suggestive of a bottom-up shift in seabird prey availability. We posit that shifts in prey composition and/or distribution, combined with the onset of molt, resulted in this mortality event.

An assessment of climate change vulnerability for Important Bird Areas in the Bering Sea and Aleutian Arc.

Recently available downscaled ocean climate models for the Bering Sea and Aleutian Arc offer the opportunity to assess climate vulnerability for upper trophic level consumers such as marine birds. We analyzed seasonal and annual spatial projections from three climate models for two physical climate variables (seawater temperature and sea ice) and three forage variables (large copepods, euphausiids, and benthic infauna), comparing projected conditions from a recent time period (2003-2012) to a future time period (2030-2039). We focused the analyses on core areas within globally significant Important Bird Areas, and developed indices of the magnitude of projected change and vulnerability agreement among models. All three climate models indicated a high degree of change for seawater temperature warming (highest in the central and eastern Aleutian Islands) and ice loss (most significant in the eastern Bering Sea) across scales, and we found those changes to be significant for every species and virtually every core area assessed. There was low model agreement for the forage variables; while the majority of core areas were identified as climate vulnerable by one or more models (72% for large copepods, 73% for euphausiids, and 94% for benthic infauna), very few were agreed upon by all three models (only 6% of euphausiid-forager core areas). Based on the magnitude-agreement score, euphausiid biomass decline affected core areas for fulmars, gulls, and auklets, especially along the outer shelf and Aleutian Islands. Benthic biomass decline affected eiders along the inner shelf, and large copepod decline was significant for storm-petrels and auklets in the western Aleutians. Overall, 12% of core areas indicated climate vulnerability for all variables assessed. Modeling and interpreting biological parameters to project future dynamics remains complex; the strong signal for projected physical changes raised concerns about lagged responses such as distribution shifts, breeding failures, mortality events, and population declines.

Circumpolar dynamics of a marine top-predator track ocean warming rates.

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long-lived, wide-ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black-legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea-surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large-scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom-up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large-scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.

Foraging responses of black-legged kittiwakes to prolonged food-shortages around colonies on the Bering Sea shelf.

We hypothesized that changes in southeastern Bering Sea foraging conditions for black-legged kittiwakes (Rissa tridactyla) have caused shifts in habitat use with direct implications for population trends. To test this, we compared at-sea distribution, breeding performance, and nutritional stress of kittiwakes in three years (2008-2010) at two sites in the Pribilof Islands, where the population has either declined (St. Paul) or remained stable (St. George). Foraging conditions were assessed from changes in (1) bird diets, (2) the biomass and distribution of juvenile pollock (Theragra chalcogramma) in 2008 and 2009, and (3) eddy kinetic energy (EKE; considered to be a proxy for oceanic prey availability). In years when biomass of juvenile pollock was low and patchily distributed in shelf regions, kittiwake diets included little or no neritic prey and a much higher occurrence of oceanic prey (e.g. myctophids). Birds from both islands foraged on the nearby shelves, or made substantially longer-distance trips overnight to the basin. Here, feeding was more nocturnal and crepuscular than on the shelf, and often occurred near anticyclonic, or inside cyclonic eddies. As expected from colony location, birds from St. Paul used neritic waters more frequently, whereas birds from St. George typically foraged in oceanic waters. Despite these distinctive foraging patterns, there were no significant differences between colonies in chick feeding rates or fledging success. High EKE in 2010 coincided with a 63% increase in use of the basin by birds from St. Paul compared with 2008 when EKE was low. Nonetheless, adult nutritional stress, which was relatively high across years at both colonies, peaked in birds from St. Paul in 2010. Diminishing food resources in nearby shelf habitats may have contributed to kittiwake population declines at St Paul, possibly driven by increased adult mortality or breeding desertion due to high foraging effort and nutritional stress.

Evaluating the power of surface attendance counts to detect long-term trends in populations of crevice-nesting auklets.

Power analyses are essential when developing a long-term monitoring program for a target species whose observation is logistically challenging and expensive. These analyses can be complicated when the observations have a complex variance structure reflecting many factors. Crevice-nesting seabirds such as least and crested auklets Aethia pusilla and Aethia cristatella illustrate both this need and these challenges. They are ecosystem indicators for the Bering Sea, a system expected to undergo large changes. Unfortunately, they are difficult to monitor as colonies occur on remote, hard to access islands in the Aleutians and Bering Sea, and nests occur in crevices underground, preventing direct observation. Current monitoring consists of breeding-season counts of auklets standing on the surface of sample plots in the colony; logically, a substantial decline in nesting population guarantees an eventual substantial decline in surface attendants. Yet, it remains debatable whether these highly variable counts can be used to statistically detect biologically relevant declines in the attending population let alone the nesting population. Subsequently, existing monitoring programs vary widely in survey design, effort levels, and daily summary statistics. The power of different survey designs was assessed by simulating observations from a state model developed from 11 years of observations using mixed-effects models and zero-inflated Poisson-lognormal regression. The analyses illustrate the process required for any monitoring program whose observations are described inadequately by standard statistical models. State model development revealed survey design refinements that reduce sampling variation. For least auklets, current sampling efforts provided 90% power to detect annual declines of 11% ("Critically Endangered" using IUCN Red List criteria), 4.5% ("Endangered"), or 2.4% ("Vulnerable") in two, four, or six generations, respectively; crested auklets took a few years longer. Power was more sensitive to number of days than number of plots. Results appear robust across a range of bird densities, providing guidance for monitoring other colonies or crevice-nesting species with similar life history strategies. Research should now focus on illuminating the relationship between the attending and nesting populations. Given the frequency of complicated variance structures and zero counts in ecological data, the general statistical models used here should prove widely applicable.