Borealization of nearshore fishes on an interior Arctic shelf over multiple decades.

Borealization is a type of community reorganization where Arctic specialists are replaced by species with more boreal distributions in response to climatic warming. The process of borealization is often exemplified by the northward range expansions and subsequent proliferation of boreal species on the Pacific and Atlantic inflow Arctic shelves (i.e., Bering/Chukchi and Barents seas, respectively). But the circumpolar nearshore distribution of Arctic-boreal fishes that predates recent warming suggests borealization is possible beyond inflow shelves. To examine this question, we revisited two nearshore lagoons in the eastern Alaska Beaufort Sea (Kaktovik and Jago lagoons, Arctic National Wildlife Refuge, Alaska, USA), a High Arctic interior shelf. We compared summer fish species assemblage, catch rate, and size distribution among three periods that spanned a 30-year record (baseline conditions, 1988-1991; moderate sea ice decline, 2003-2005; rapid sea ice decline, 2017-2019). Fish assemblages differed among periods in both lagoons, consistent with borealization. Among Arctic specialists, a clear decline in fourhorn sculpin (Myoxocephalus quadricornis, Kanayuq in Iñupiaq) occurred in both lagoons with 86%-90% lower catch rates compared with the baseline period. Among the Arctic-boreal species, a dramatic 18- to 19-fold increase in saffron cod (Eleginus gracilis, Uugaq) occurred in both lagoons. Fish size (length) distributions demonstrated increases in the proportion of larger fish for most species examined, consistent with increasing survival and addition of age-classes. These field data illustrate borealization of an Arctic nearshore fish community during a period of rapid warming. Our results agree with predictions that Arctic-boreal fishes (e.g., saffron cod) are well positioned to exploit the changing Arctic ecosystem. Another Arctic-boreal species, Dolly Varden (Salvelinus malma, Iqalukpik), appear to have already responded to warming by shifting from Arctic nearshore to shelf waters. More broadly, our findings suggest that areas of borealization could be widespread in the circumpolar nearshore.

Investigating effects of climate-induced changes in water temperature and diet on mercury concentrations in an Arctic freshwater forage fish.

The amount of mercury (Hg) in Arctic lake food webs is, and will continue to be, affected by rapid, ongoing climate change. At warmer temperatures, fish require more energy to sustain growth; changes in their metabolic rates and consuming prey with potentially higher Hg concentrations could result in increased Hg accumulation. To examine the potential implications of climate warming on forage fish Hg accumulation in Arctic lakes, we quantified growth and Hg accumulation in Ninespine Stickleback Pungitius pungitius under different temperature and diet scenarios using bioenergetics models. Four scenarios were considered that examined the role of climate, diet, climate × diet, and climate × diet × elevated prey Hg. As expected, annual fish growth increased with warmer temperatures, but growth rates and Hg accumulation were largely diet dependent. Compared to current growth rates of 0.3 g⋅y-1, fish growth increased at least 200% for fish consuming energy-dense benthic prey and decreased at least 40% for fish consuming pelagic prey. Compared to baseline levels, the Hg burden per kilocalorie of Ninespine Stickleback declined up to 43% with benthic consumption - indicating strong somatic growth dilution - but no more than 4% with pelagic consumption; elevated prey Hg concentrations led to moderate Hg declines in benthic-foraging fish and Hg increases in pelagic-foraging fish. Bioenergetics models demonstrated the complex interaction of water temperature, growth, prey proportions, and prey Hg concentrations that respond to climate change. Further work is needed to resolve mechanisms and rates linking climate change to Hg availability and uptake in Arctic freshwater systems.

A manipulative thermal challenge protocol for adult salmonids in remote field settings.

Manipulative experiments provide stronger evidence for identifying cause-and-effect relationships than correlative studies, but protocols for implementing temperature manipulations are lacking for large species in remote settings. We developed an experimental protocol for holding adult Chinook salmon (Oncorhynchus tshawytscha) and exposing them to elevated temperature treatments. The goal of the experimental protocol was to validate heat stress biomarkers by increasing river water temperature from ambient (~14°C) to a treatment temperature of 18°C or 21°C and then maintain the treatment temperature over 4 hours within a range of ±1.0°C. Our protocol resulted in a mean rate of temperature rise of 3.71°C h-1 (SD = 1.31) to treatment temperatures and mean holding temperatures of 18.0°C (SD = 0.2) and 21.0°C (SD = 0.2) in the low- and high-heat treatments, respectively. Our work demonstrated that manipulative experiments with large, mobile study species can be successfully developed in remote locations to examine thermal stress.

Transcriptomic response to elevated water temperatures in adult migrating Yukon River Chinook salmon (Oncorhynchus tshawytscha).

Chinook salmon (Oncorhynchus tshawytscha) declines are widespread and may be attributed, at least in part, to warming river temperatures. Water temperatures in the Yukon River and tributaries often exceed 18°C, a threshold commonly associated with heat stress and elevated mortality in Pacific salmon. Untangling the complex web of direct and indirect physiological effects of heat stress on salmon is difficult in a natural setting with innumerable system challenges but is necessary to increase our understanding of both lethal and sublethal impacts of heat stress on populations. The goal of this study was to characterize the cellular stress response in multiple Chinook salmon tissues after acute elevated temperature challenges. We conducted a controlled 4-hour temperature exposure (control, 18°C and 21°C) experiment on the bank of the Yukon River followed by gene expression (GE) profiling using a 3'-Tag-RNA-Seq protocol. The full transcriptome was analysed for 22 Chinook salmon in muscle, gill and liver tissue. Both the 21°C and 18°C treatments induced greater activity in genes associated with protein folding (e.g. HSP70, HSP90 mRNA) processes in all tissues. Global GE patterns indicate that transcriptomic responses to heat stress were highly tissue-specific, underscoring the importance of analyzing multiple tissues for determination of physiological effect. Primary superclusters (i.e. groupings of loosely related terms) of altered biological processes were identified in each tissue type, including regulation of DNA damage response (gill), regulation by host of viral transcription (liver) and regulation of the force of heart contraction (muscle) in the 21°C treatment. This study provides insight into mechanisms potentially affecting adult Chinook salmon as they encounter warm water during their spawning migration in the Yukon River and suggests that both basic and more specialized cellular functions may be disrupted.