Trace Elements in Sea Ducks of the Alaskan Arctic Coast: Patterns of Variation Among Species, Sexes, and Ages.

Climate change and increasing industrialization in the Arctic call for the collection of reference data for assessing changes in contaminant levels. For migratory birds, measuring and interpreting changes in trace element burdens on Arctic breeding areas require insights into factors such as sex, body size, or wintering area that may modify patterns independently of local exposure. In the Alaskan Arctic, we determined levels of trace elements in liver and kidney of common eiders (Somateria mollissima) and long-tailed ducks (Clangula hyemalis) from the Prudhoe Bay oil field and of king eiders (S. spectabilis) and threatened spectacled eiders (S. fischeri) and Steller's eiders (Polystica stelleri) from near the town of Barrow. Small-bodied Steller's eiders and long-tailed ducks from different locations had similarly low levels of selenium (Se), cadmium (Cd), and copper (Cu), perhaps reflecting high mass-specific rates of metabolic depuration during long spring migrations through areas of low exposure. In larger species, Se, Cd, and Cu concentrations were higher in adults than juveniles suggesting that these elements were acquired in nonbreeding marine habitats. Adult male spectacled eiders had exceptionally high Se, Cd, and Cu compared with adult females, possibly because of depuration into eggs and longer female occupancy of nonmarine habitats. Adult female common eiders and juvenile long-tailed ducks at Prudhoe Bay had high and variable levels of Pb, potentially due to local exposure. Explanations for substantial variations in Hg levels were not apparent. Further research into reasons for differing element levels among species and sexes will help clarify the sources, pathways, and risks of exposure.

Variable wind, pack ice, and prey dispersion affect the long-term adequacy of protected areas for an Arctic sea duck.

With changing climate, delineation of protected areas for sensitive species must account for long-term variability and geographic shifts of key habitat elements. Projecting the future adequacy of protected areas requires knowing major factors that drive such changes, and how readily the animals adjust to altered resources. In the Arctic, the viability of habitats for marine birds and mammals often depends on sea ice to dissipate storm waves and provide platforms for resting. However, some wind conditions (including weak winds during extreme cold) can consolidate pack ice into cover so dense that air-breathing divers are excluded from the better feeding areas. Spectacled Eiders (Somateria fischeri) winter among leads (openings) in pack ice in areas where densities of their bivalve prey are quite high. During winter 2009, however, prevailing winds created a large region of continuous ice with inadequate leads to allow access to areas of dense preferred prey. Stable isotope and fatty acid biomarkers indicated that, under these conditions, the eiders did not diversify their diet to include abundant non-bivalve taxa but did add a smaller, less preferred, bivalve species. Consistent with a computer model of eider energy balance, the body fat of adult eiders in 2009 was 33-35% lower than on the same date (19 March) in 2001 when ice conditions allowed access to higher bivalve densities. Ice cover data suggest that the eiders were mostly excluded from areas of high bivalve density from January to March in about 30% of 14 winters from 1998 to 2011. Thus, even without change in total extent of ice, shifts in prevailing winds can alter the areal density of ice to reduce access to important habitats. Because changes in wind-driven currents can also rearrange the dispersion of prey, the potential for altered wind patterns should be an important concern in projecting effects of climate change on the adequacy of marine protected areas for diving endotherms in the Arctic.

Deposit­feeder diets in the Bering Sea: potential effects of climatic loss of sea ice­related microalgal blooms.

Climate warming in seasonally ice-covered seas is expected to reduce the extent and duration of annual sea ice. Resulting changes in sea ice related blooms of ice algae or phytoplankton may in turn alter the timing, magnitude, or quality of organic matter inputs to the sea floor. If benthic taxa rely differently on direct consumption of settling fresh microalgae for growth and reproduction, altered blooms may lead to reorganization of deposit-feeding assemblages. To assess the potential for such changes, we examined the diets of five abundant deposit-feeders (three infaunal bivalves, a polychaete, and a brittle star) with different feeding modes over the course of the spring bloom in May­June 2007 in the north-central Bering Sea (30­90 m depth). Short-term data from gut contents reflected feeding modes, with the bivalves Macoma calcarea, Ennucula tenuis, and Nuculana radiata, and the brittle star Ophiura sarsi, responding more quickly to deposition of fresh algae than did the head-down polychaete Pectinaria hyperborea. Fatty acid biomarkers also indicated rapid ingestion of settling algae by the bivalves (especially Macoma) and the brittle star, while Pectinaria continued to ingest mainly bacteria. Fatty acid biomarkers did not indicate any unique dietary importance of ice algae released from melting ice. Longer-term inference from stable isotopes suggested that fresh microalgae contributed little to overall carbon assimilated by any of these species. Instead, deposit-feeders appeared to select a consistent fraction from the pool of sediment organic matter, probably heterotrophic microbes, microbial products, and reworked phytodetritus that form a longer-term sediment "food bank." Redistribution of settled organic matter via scouring and accumulation by currents, as well as the multi-year life spans of macroinvertebrates, may further overwhelm effects of short-term variations in the timing, magnitude, and dispersion of blooms in the water column. More diet data are needed from midsummer to account for any lag in assimilation of fresh microalgae at these cold temperatures. Nevertheless, our results suggest that if annual sea ice cover is reduced, increased production of phytoplankton during longer ice-free periods could replace inputs of ice-associated microalgae to the sediment food bank used by deposit-feeders.

Organic matter sources and flows in tundra wetland food webs.

Arctic lowland tundra is often dominated by wetlands. As numbers and types of these wetlands change with climate warming, their invertebrate biomass and assemblages may also be affected. Increased influx of nutrients and dissolved organic matter (DOM) from thawing peat may alter the relative availability of organic matter (OM) sources, differentially affecting taxa with disparate dependence on those sources. In five shallow wetland types (<40 to 110 cm deep) and in littoral zones of deeper lakes (>150 cm), we used stable isotopes (δ13C, δ15N) to compare contributions of four OM sources (periphytic microalgae, cyanobacteria, macrophytes, peat) to the diets of nine macroinvertebrate taxa. Living macrophytes were not distinguishable isotopically from peat that likely contributed most DOM. Within invertebrate taxa, relative OM contributions were similar among all wetland types except deeper lakes. Physidae snails consumed substantial amounts of OM from cyanobacteria. However, for all other taxa examined, microalgae were the dominant or a major OM source (39-82%, mean 59%) in all wetland types except deeper lakes (20‒62%, mean 31%). Macrophytes and macrophyte-derived peat, likely consumed mostly indirectly as DOM-supported bacteria, ranged from 18‒61% (mean 41%) of ultimate OM sources in all wetland types except deeper lakes (38-80%, mean 69%). Invertebrate consumption of microalgal C may often have involved bacterial intermediates, or a mix of algae with bacteria consuming peat-derived OM. High production of periphyton with very low δ13C values were favored by continuous daylight illuminating shallow depths, high N and P levels, and high CO2 concentrations from bacterial respiration of peat-derived DOM. Although relative OM sources were similar across wetland types except deeper lakes, total invertebrate biomass was much higher in shallow wetlands with emergent vegetation. Impacts of warming on the availability of invertebrate prey to waterbirds will likely depend not on shifts in OM sources, but more on changes in overall number or area of shallow emergent wetlands.

Prey availability and foraging activity by tundra-nesting sea ducks: Strong preference for specific wetland types.

Wetlands in Arctic tundra support abundant breeding waterbirds. Wetland types differing in area, depth, vegetation, and invertebrate biomass density may vary in importance to birds, and in vulnerability to climate change. We studied availability and use of different wetland types by prelaying females of four species of sea ducks (Mergini) breeding on the Arctic Coastal Plain of Alaska, USA: long-tailed ducks (Clangula hyemalis) and Steller's (Polysticta stelleri), spectacled (Somateria fischeri), and king eiders (Somateria spectabilis). All four species preferred shallow vegetated wetlands versus deeper lakes. The ducks spent almost all their active time feeding, but their occurrence in different wetland types was not affected by the relative biomass density of known prey or of all invertebrates that we sampled combined. Sea ducks strongly preferred wetlands dominated by emergent and submersed Arctophila fulva over those dominated by the sedge Carex aquatilis, despite the much greater number, total area, and invertebrate biomass density of Carex wetlands. The hens depend heavily on local invertebrate prey for protein to produce eggs; thus, their preference for Arctophila wetlands likely reflects greater accessibility of prey in the near-surface canopy and detritus of Arctophila. Such shallow wetlands decreased substantially in number (-17%) and area (-30%) over 62 years before 2013 and appear highly susceptible to further declines with climate warming. Impacts on sea ducks of climate-driven changes in availability of important wetland types will depend on their adaptability in exploiting alternative wetlands.

Detecting silent stressors: Trace element effects on nutritional status of declining scoter ducks of Puget Sound, USA.

White-winged scoters (Melanitta fusca; WWSC) and surf scoters (M. perspicillata; SUSC) have declined by over 60% in recent decades. Identifying contributing factors from within a mosaic of sublethal, multiple stressors is challenging. In urbanized Puget Sound, Washington, USA where scoters winter, changes in prey availability explained only a portion of local declines, suggesting that other "silent stressors" such as sublethal contaminants might play a role. Past studies of pollutant effects on scoters used Fisherian statistics that often revealed few correlates; however, novel statistical approaches could detect and provide more insights about sublethal impacts. Our objectives were to (1) relate pollutant accumulation to health of the birds, and (2) compare permutational multivariate statistics with traditional approaches in identifying sublethal health effects. We collected scoters from three locations in Puget Sound in December 2005 and March 2006, and measured cadmium (Cd), mercury (Hg), and selenium (Se) levels in livers and kidneys. To assess impacts of low contaminants levels in tissues on nutritional status (whole-body mass, lipid, and protein; and triglycerides, ß-hydroxybutyrate, and uric acid in blood), we compared statistical methods. Permutational multivariate methods use Monte Carlo techniques to assess how an integrated matrix of physiological responses in each animal respond to contaminants. Univariate regressions revealed very few and inconsistent relationships. In contrast, multivariate models showed that liver Hg and Se explained 25% of the variance in nutritional status of white-winged scoters; and in surf scoters, Cd, Hg, and Se in tissues explained 14 to 27% of nutritional status depending on site. The influence of these factors equals other aspects of habitat such as foraging conditions. Our study indicates that permutational multivariate statistics can be a powerful tool for identifying sublethal contaminant associations that, with non-contaminant stressors, can influence nutritional status and thus, contribute to population dynamics.

Modeling marine protected areas for threatened eiders in a climatically changing Bering Sea.

Delineating protected areas for sensitive species is a growing challenge as changing climate alters the geographic pattern of habitats as well as human responses to those shifts. When human impacts are expected within projected ranges of threatened species, there is often demand to demarcate the minimum habitat required to ensure the species' persistence. Because diminished or wide-ranging populations may not occupy all viable (and needed) habitat at once, one must identify thresholds of resources that will support the species even in unoccupied areas. Long-term data on the shifting mosaic of critical resources may indicate ranges of future variability. We addressed these issues for the Spectacled Eider (Somateria fischeri), a federally threatened species that winters in pack ice of the Bering Sea. Changing climate has decreased ice cover and severely reduced the eiders' benthic prey and has increased prospects for expansion of bottom trawling that may further affect prey communities. To assess long-term changes in habitats that will support eiders, we linked data on benthic prey, sea ice, and weather from 1970 to 2001 with a spatially explicit simulation model of eider energy balance that integrated field, laboratory, and remote-sensing studies. Areas estimated to have prey densities adequate for eiders in 1970-1974 did not include most areas that were viable 20 years later (1993-1994). Unless the entire area with adequate prey in 1993-1994 had been protected, the much reduced viable area in 1999-2001 might well have been excluded. During long non-foraging periods (as at night), eiders can save much energy by resting on ice vs. floating on water; thus, loss of ice cover in the future might substantially decrease the area in which prey densities are adequate to offset the eiders' energy needs. For wide-ranging benthivores such as eiders, our results emphasize that fixed protected areas based on current conditions can be too small or inflexible to subsume long-term shifts in habitat conditions. Better knowledge of patterns of natural disturbance experienced by prey communities, and appropriate allocation of human disturbance over seasons or years, may yield alternative strategies to large-scale closures that may be politically and economically problematic.

Predator size, prey size and threshold food densities of diving ducks: does a common prey base support fewer large animals?

1. Allometry predicts that a given habitat area or common prey biomass supports fewer numbers of larger than smaller predators; however, birds from related taxa or the same feeding guild often deviate from this pattern. In particular, foraging costs of birds may differ among locomotor modes, while intake rates vary with accessibility, handling times and energy content of different-sized prey. Such mechanisms might affect threshold prey densities needed for energy balance, and thus relative numbers of different-sized predators in habitats with varying prey patches. 2. We compared the foraging profitability (energy gain minus cost) of two diving ducks: smaller lesser scaup (Aythya affinis, 450-1090 g) and larger white-winged scoters (Melanitta fusca, 950-1800 g). Calculations were based on past measurements of dive costs with respirometry, and of intake rates of a common bivalve prey ranging in size, energy content and burial depth in sediments. 3. For scaup feeding on small prey <12 mm long, all clams buried deeper than 5 cm were unprofitable at realistic prey densities. For clams buried in the top 5 cm, the profitability threshold decreased from 216 to 34 clams m(-2) as energy content increased from 50 to 300 J clam(-1). 4. For larger scoters feeding on larger prey 18-24 mm long, foraging was profitable for clams buried deeper than 5 cm, with a threshold density of 147 m(-2) for clams containing 380 J clam(-1). For clams <5 cm deep, the threshold density decreased from 86 to 36 clams m(-2) as energy content increased from 380 to 850 J clam(-1). If scoters decreased dive costs by swimming with wings as well as feet (not an option for scaup), threshold prey densities were 11-12% lower. 5. Our results show that threshold densities of total prey numbers for different-sized ducks depend on prey size structure and depth in the sediments. Thus, heterogeneity in disturbance regimes and prey population dynamics can create a mosaic of patches favouring large or small predators. Whether a given area or total prey biomass will support greater numbers of larger or smaller predators will vary with these effects.

Effects of air and water temperatures on resting metabolism of auklets and other diving birds.

For small aquatic endotherms, heat loss while floating on water can be a dominant energy cost, and requires accurate estimation in energetics models for different species. We measured resting metabolic rate (RMR) in air and on water for a small diving bird, the Cassin's auklet (Ptychoramphus aleuticus), and compared these results to published data for other diving birds of diverse taxa and sizes. For 8 Cassin's auklets (~165 g), the lower critical temperature was higher on water (21 °C) than in air (16 °C). Lowest values of RMR (W kg⁻¹) averaged 19% higher on water (12.14 ± 3.14 SD) than in air (10.22 ± 1.43). At lower temperatures, RMR averaged 25% higher on water than in air, increasing with similar slope. RMR was higher on water than in air for alcids, cormorants, and small penguins but not for diving ducks, which appear exceptionally resistant to heat loss in water. Changes in RMR (W) with body mass either in air or on water were mostly linear over the 5- to 20-fold body mass ranges of alcids, diving ducks, and penguins, while cormorants showed no relationship of RMR with mass. The often large energetic effects of time spent floating on water can differ substantially among major taxa of diving birds, so that relevant estimates are critical to understanding their patterns of daily energy use.

Swim speeds and stroke patterns in wing-propelled divers: a comparison among alcids and a penguin.

In diving birds, the volume and resulting buoyancy of air spaces changes with dive depth, and hydrodynamic drag varies with swim speed. These factors are important in the dive patterns and locomotion of alcids that use their wings both for aerial flight and underwater swimming and of penguins that use their wings only for swimming. Using small data-loggers on free-ranging birds diving to 20-30 m depth, we measured depth at 1 Hz and surge and heave accelerations at 32-64 Hz of four species of alcids (0.6-1.0 kg mass) and the smallest penguin species (1.2 kg). Low- and high-frequency components of the fluctuation of acceleration yielded estimates of body angles and stroke frequencies, respectively. Swim speed was estimated from body angle and rate of depth change. Brünnich's (Uria lomvia) and common (Uria aalge) guillemots descended almost vertically, whereas descent of razorbills (Alca torda), rhinoceros auklets (Cerorhinca monocerata) and little penguins (Eudyptula minor) was more oblique. For all species, swim speed during descent was within a relatively narrow range. Above depths of 20-30 m, where they were all positively buoyant, all species ascended without wing stroking. During descent, little penguins made forward accelerations on both the upstroke and downstroke regardless of dive depth. By contrast, descending alcids produced forward accelerations on both upstroke and downstroke at depths of <10 m but mainly on the downstroke at greater depths; this change seemed to correspond to the decrease of buoyancy with increasing depth. The magnitude of surge (forward) acceleration during downstrokes was smaller, and that during upstrokes greater, in little penguins than in alcids. This pattern presumably reflected the proportionally greater mass of upstroke muscles in penguins compared with alcids and may allow little penguins to swim at less variable instantaneous speeds.