Pacific salmon in the Canadian Arctic highlight a range-expansion pathway for sub-Arctic fishes.

Rapid climate change is altering Arctic ecosystems at unprecedented rates. These changes in the physical environment may open new corridors for species range expansions, with substantial implications for subsistence-dependent communities and sensitive ecosystems. Over the past 20 years, rising incidental harvest of Pacific salmon by subsistence fishers has been monitored across a widening range spanning multiple land claim jurisdictions in Arctic Canada. In this study, we connect Indigenous and scientific knowledges to explore potential oceanographic mechanisms facilitating this ongoing northward expansion of Pacific salmon into the western Canadian Arctic. A regression analysis was used to reveal and characterize a two-part mechanism related to thermal and sea-ice conditions in the Chukchi and Beaufort seas that explains nearly all of the variation in the relative abundance of salmon observed within this region. The results indicate that warmer late-spring temperatures in a Chukchi Sea watch-zone and persistent, suitable summer thermal conditions in a Beaufort Sea watch-zone together create a range-expansion corridor and are associated with higher salmon occurrences in subsistence harvests. Furthermore, there is a body of knowledge to suggest that these conditions, and consequently the presence and abundance of Pacific salmon, will become more persistent in the coming decades. Our collaborative approach positions us to document, explore, and explain mechanisms driving changes in fish biodiversity that have the potential to, or are already affecting, Indigenous rights-holders in a rapidly warming Arctic.

Gill raker and pyloric caeca counts differ between Arctic char (Salvelinus alpinus) and Dolly Varden (S. malma) populations across their ranges.

Meristic characters are often used to differentiate between closely related forms, morphs, and species of fishes, and lend insight into ecology and post-glacial recolonization in taxa with complicated or contentious phylogenies, including the genus Salvelinus. Previous studies of meristics in Salvelinus have focused mostly on individual populations. We collated data from 456 populations/systems across the North American and Russian Arctic and sub-Arctic, and found that counts of pyloric caeca and gill rakers differed consistently between fish visually and/or genetically identified as Arctic char and Dolly Varden across their distributional ranges.

A new probabilistic method for quantifying n-dimensional ecological niches and niche overlap.

Considerable progress has been made in the development of statistical tools to quantify trophic relationships using stable isotope ratios, including tools that address size and overlap of isotopic niches. We build upon recent progress and propose a new probabilistic method for determining niche region and pairwise niche overlap that can be extended beyond two dimensions, provides directional estimates of niche overlap, accounts for species-specific distributions in niche space, and, unlike geometric methods, produces consistent and unique bivariate projections of multivariate data. We define the niche region (NR) as a given 95% (or user-defined a) probability region in multivariate space. Overlap is calculated as the probability that an individual from species A is found in the N(R) of species B. Uncertainty is accounted for in a Bayesian framework, and is the only aspect of the methodology that depends on sample size. Application is illustrated with three-dimensional stable isotope data, but practitioners could use any continuous indicator of ecological niche in any number of dimensions. We suggest that this represents an advance in our ability to quantify and compare ecological niches in a way that is more consistent with Hutchinson's concept of an "n-dimensional hypervolume".

Polycyclic aromatic hydrocarbon metabolites in Arctic cod (Boreogadus saida) from the Beaufort Sea and associative fish health effects.

In 2012, Arctic cod (Boreogadus saida) were collected from offshore regions of the Beaufort Sea to determine the concentrations of CYP1A1 phase I metabolites of polycyclic aromatic hydrocarbons (OH-PAHs) in liver and to correlate measured concentrations with (i) morphometric measurements that are known to be indicative of fish health and, (ii) biochemical end points of health including vitamin A/E and metabolites and hepatic deiodinase activity (DI). Four ring OH-PAHs were detected in 90% of our samples with a mean liver concentration of 1829.2 ± 159.2 ng/g (ww). Total (∑) concentrations of 5/6-membered ring OH-PAHs in liver were smaller [mean of 931.6 ± 104.3 ng/g, (ww)] and detected less frequently (75%) than the 4-ring OH-PAHs. Fish length and liver weight were both negatively correlated to ∑ concentrations of 4-ringed OH-PAHs (p < 0.001). Liver somatic index was also negatively correlated to ∑4-OH-PAHs (p < 0.05) but not for ∑5/6-OH-PAHs (p > 0.1). There was a significant positive relationship between DI and 4-ring OH-PAHs (p < 0.05) in liver, suggesting an induction of this enzyme. No such correlation was observed for the 5/6-ring OH-PAHs. Retinyl palmitate (RP) was the only vitamin that could be measured in liver ranging from 0.230 to 26.3 ug/g (ww). No associations between RP and levels of the 4- or 5/6-ringed OH-PAHs were observed. Continued baseline studies are clearly warranted to further understand effects of OH-PAHs on fish health before planned exploration activities begin in this region.

A meta-collection of nitrogen stable isotope data measured in Arctic marine organisms from the Canadian Beaufort Sea, 1983-2013.

OBJECTIVES: Existing information on Arctic marine food web structure is fragmented. Integrating data across research programs is an important strategy for building a baseline understanding of food web structure and function in many Arctic regions. Naturally-occurring stable isotope ratios of nitrogen (δ15N) and carbon (δ13C) measured directly in the tissues of organisms are a commonly-employed method for estimating food web structure. The objective of the current dataset was to synthesize disparate δ15N, and secondarily δ13C, data in the Canadian Beaufort continental shelf region relevant to trophic and ecological studies at the local and pan-Arctic scales. DATA DESCRIPTION: The dataset presented here contains nitrogen and carbon stable isotope ratios (δ15N, δ13C) measured in marine organisms from the Canadian Beaufort continental shelf region between 1983 and 2013, gathered from 27 published and unpublished sources with associated sampling metadata. A total of 1077 entries were collected, summarizing 8859 individual organisms/samples representing 333 taxa across the Arctic food web, from top marine mammal predators to primary producers.

Mercury concentrations in landlocked Arctic char (Salvelinus alpinus) from the Canadian Arctic. Part II: influence of lake biotic and abiotic characteristics on geographic trends in 27 populations.

Among-lake variation in mercury (Hg) concentrations in landlocked Arctic char was examined in 27 char populations from remote lakes across the Canadian Arctic. A total of 520 landlocked Arctic char were collected from 27 lakes, as well as sediments and surface water from a subset of lakes in 1999, 2002, and 2005 to 2007. Size, length, age, and trophic position (delta(15)N) of individual char were determined and relationships with total Hg (THg) concentrations investigated, to identify a common covariate for adjustment using analysis of covariance (ANCOVA). A subset of 216 char from 24 populations was used for spatial comparison, after length-adjustment. The influence of trophic position and food web length and abiotic characteristics such as location, geomorphology, lake area, catchment area, catchment-to-lake area ratio of the lakes on adjusted THg concentrations in char muscle tissue were then evaluated. Arctic char from Amituk Lake (Cornwallis Island) had the highest Hg concentrations (1.31 microg/g wet wt), while Tessisoak Lake (Labrador, 0.07 microg/g wet wt) had the lowest. Concentrations of THg were positively correlated with size, delta(15)N, and age, respectively, in 88, 71, and 58% of 24 char populations. Length and delta(15)N were correlated in 67% of 24 char populations. Food chain length did not explain the differences in length-adjusted THg concentrations in char. No relationships between adjusted THg concentrations in char and latitude or longitude were found, however, THg concentrations in char showed a positive correlation with catchment-to-lake area ratio. Furthermore, we conclude that inputs from the surrounding environment may influence THg concentrations, and will ultimately affect THg concentrations in char as a result of predicted climate-driven changes that may occur in Arctic lake watersheds.

Temporal trends of mercury, cesium, potassium, selenium, and thallium in Arctic char (Salvelinus alpinus) from Lake Hazen, Nunavut, Canada: effects of trophic position, size, and age.

Arctic char (Salvelinus alpinus L.), the top predator in High Arctic lakes, often is used as a bioindicator of Hg contamination in Arctic aquatic ecosystems. The present study investigated effects of trophic position, size, and age of Arctic char in Lake Hazen, the largest lake in the Canadian High Arctic (81 degrees 50'N, 70 degrees 25'W), on Hg bioaccumulation. In addition, several essential (Se, K) and nonessential elements (Tl, Cs) in char muscle tissue were examined to compare their behavior to that of Hg. Trophic position of Arctic char was identified by stable isotope (delta15N) signature. Temporal trends of Hg from seven sampling campaigns over a 16-year period (1990-2006) were investigated for the overall data and for one trophic class. Concentrations of Hg were not correlated with age but were positively related to fork length and trophic position. Large char with greater delta15N signatures (> 12 per thousand) had larger Hg concentrations (0.09-1.63 microg/g wet wt) than small char with smaller delta15N signatures (< 12 per thousand, 0.03-0.32 microg/g wet wt), indicating that Hg concentrations increased with trophic position. Nonessential Cs and Tl showed relationships to age, length, and trophic position similar to those of Hg, indicating their potential to bioaccumulate and biomagnify. Essential Se and K did not show these relationships. Concentrations of Hg were adjusted using delta15N, leading to less within-year variability and a more consistent temporal trend. The delta15N-adjusted trend showed no decline of Hg in Arctic char from Lake Hazen (1990-2006) in the overall data set and in the small morphotype. Trends for the same period before the adjustment were not significant for the overall data set, but a slight decrease was apparent in the small morphotype. The results confirm the need to consider trophic position and fish size when monitoring temporal trends of Hg, particularly for species with different morphotypes.

Implications of climate change for northern Canada: freshwater, marine, and terrestrial ecosystems.

Climate variability and change is projected to have significant effects on the physical, chemical, and biological components of northern Canadian marine, terrestrial, and freshwater systems. As the climate continues to change, there will be consequences for biodiversity shifts and for the ranges and distribution of many species with resulting effects on availability, accessibility, and quality of resources upon which human populations rely. This will have implications for the protection and management of wildlife, fish, and fisheries resources; protected areas; and forests. The northward migration of species and the disruption and competition from invading species are already occurring and will continue to affect marine, terrestrial, and freshwater communities. Shifting environmental conditions will likely introduce new animal-transmitted diseases and redistribute some existing diseases, affecting key economic resources and some human populations. Stress on populations of iconic wildlife species, such as the polar bear, ringed seals, and whales, will continue as a result of changes in critical sea-ice habitat interactions. Where these stresses affect economically and culturally important species, they will have significant effects on people and regional economies. Further integrated, field-based monitoring and research programs, and the development of predictive models are required to allow for more detailed and comprehensive projections of change to be made, and to inform the development and implementation of appropriate adaptation, wildlife, and habitat conservation and protection strategies.

Osmolyte Adjustments as a Pressure Adaptation in Deep-Sea Chondrichthyan Fishes: An Intraspecific Test in Arctic Skates (Amblyraja hyperborea) along a Depth Gradient.

Accumulation of trimethylamine N-oxide (TMAO) by deep-sea animals is proposed to protect proteins against the destabilizing effects of high hydrostatic pressure (the piezolyte hypothesis). Chondrichthyan fishes (sharks, rays, and chimaeras) provide a unique test of this hypothesis because shallow-living species have elevated TMAO levels to counteract the destabilizing effects of high urea levels accumulated for osmoregulation. Limited interspecific studies of chondrichthyans reveal that increasing depth correlates with decreased urea and increased TMAO levels, suggesting a dynamic balance between destabilizing forces on proteins (high urea, hydrostatic pressure) and TMAO to counteract these forces. Indeed, an inability to minimize urea levels or maximize TMAO levels has been proposed to explain why chondrichthyans are absent in the vast abyssal region. An unresolved question is whether the depth-related changes in chondrichthyan osmolytes are a flexible response to depth or whether phylogenetic differences in species-specific physiological set points for osmolytes account for the differences seen with depth. Sampling Arctic skates (Amblyraja hyperborea) across a 1,015-m depth gradient in the Beaufort Sea, we measured organic osmolytes in muscle using spectrophotometry and high-performance liquid chromatography. We found that the urea-to-TMAO ratio decreased linearly with depth, with tighter correlation than that seen in interspecific studies. Minor osmolytes, including betaine, sarcosine, and some α-amino acids, also declined with depth, apparently replaced (as with urea) by TMAO (a stronger piezolyte than those solutes). These data provide the first intraspecific evidence that flexible adjustments of osmolyte combinations are a key response for deep-sea living in individual chondrichthyans, supporting the piezolyte hypothesis.

Climate impacts on arctic freshwater ecosystems and fisheries: background, rationale and approach of the Arctic Climate Impact Assessment (ACIA).

Changes in climate and ultraviolet radiation levels in the Arctic will have far-reaching impacts, affecting aquatic species at various trophic levels, the physical and chemical environment that makes up their habitat, and the processes that act on and within freshwater ecosystems. Interactions of climatic variables, such as temperature and precipitation, with freshwater ecosystems are highly complex and can propagate through the ecosystem in ways that are difficult to project. This is partly due to a poor understanding of arctic freshwater systems and their basic interrelationships with climate and other environmental variables, and partly due to a paucity of long-term freshwater monitoring sites and integrated hydro-ecological research programs in the Arctic. The papers in this special issue are an abstraction of the analyses performed by 25 international experts and their associated networks on Arctic freshwater hydrology and related aquatic ecosystems that was initially published by the Arctic Climate Impact Assessment (ACIA) in 2005 as "Chapter 8--Freshwater Ecosystems and Fisheries". The papers provide a broad overview of the general hydrological and ecological features of the various freshwater ecosystems in the Arctic, including descriptions of each ACIA region, followed by a review of historical changes in freshwater systems during the Holocene. This is followed by an assessment of the effects of climate change on broad-scale hydro-ecology; aquatic biota and ecosystem structure and function; and arctic fish and fisheries. Potential synergistic and cumulative effects are also discussed, as are the roles of ultraviolet radiation and contaminants. The nature and complexity of many of the effects are illustrated using case studies from around the circumpolar north, together with a discussion of important threshold responses (i.e., those that produce stepwise and/or nonlinear effects). The issue concludes with summary the key findings, a list of gaps in scientific understanding, and policy-related recommendations.

General features of the arctic relevant to climate change in freshwater ecosystems.

Large variations exist in the size, abundance and biota of the two principal categories of freshwater ecosystems, lotic (flowing water; e.g., rivers, streams, deltas and estuaries) and lentic (standing water; lakes, ponds and wetlands) found across the circumpolar Arctic. Arctic climate, many components of which exhibit strong variations along latitudinal gradients, directly affects a range of physical, chemical and biological processes in these aquatic systems. Furthermore, arctic climate creates additional indirect ecological effects through the control of terrestrial hydrologic systems and processes, particularly those associated with cryospheric components such as permafrost, freshwater ice and snow accumulation/ablation. The ecological structure and function of arctic freshwater systems are also controlled by external processes and conditions, particularly those in the headwaters of the major arctic rivers and in the adjacent marine environment. The movement of physical, chemical and biotic components through the interlinked lentic and lotic freshwater systems are major determinants of arctic freshwater ecology.

Historical changes in arctic freshwater ecosystems.

Various types of ecosystem-based climate proxies have been used to assess past arctic change. Although lotic records are relatively poor because of the constant reworking of riverine material, high-quality lentic data have been assembled back to the end of the Pleistocene and deglaciation of the circumpolar Arctic. In general, climatic variations in the Holocene, partly due to changes in the shrinking effect of glacier coverage, produced significant temporal and spatial variations in arctic hydrology and freshwater ecosystems. Of particular note were the vast expansions of northern peatlands during major protracted periods of wetting. More recent lake biota and sedimentiological data reflect the general warming trend that has occurred over the last one to two centuries and indicate major changes to freshwater characteristics such as ice-cover duration and thermal stratification. Such data provide an excellent baseline against which future effects of climate change can be both projected and measured.

Climate change effects on hydroecology of arctic freshwater ecosystems.

In general, the arctic freshwater-terrestrial system will warm more rapidly than the global average, particularly during the autumn and winter season. The decline or loss of many cryospheric components and a shift from a nival to an increasingly pluvial system will produce numerous physical effects on freshwater ecosystems. Of particular note will be reductions in the dominance of the spring freshet and changes in the intensity of river-ice breakup. Increased evaporation/evapotranspiration due to longer ice-free seasons, higher air/water temperatures and greater transpiring vegetation along with increase infiltration because of permafrost thaw will decrease surface water levels and coverage. Loss of ice and permafrost, increased water temperatures and vegetation shifts will alter water chemistry, the general result being an increase in lotic and lentic productivity. Changes in ice and water flow/levels will lead to regime-specific increases and decreases in habitat availability/quality across the circumpolar Arctic.

Climate change effects on aquatic biota, ecosystem structure and function.

Climate change is projected to cause significant alterations to aquatic biogeochemical processes, (including carbon dynamics), aquatic food web structure, dynamics and biodiversity, primary and secondary production; and, affect the range, distribution and habitat quality/quantity of aquatic mammals and waterfowl. Projected enhanced permafrost thawing is very likely to increase nutrient, sediment, and carbon loadings to aquatic systems, resulting in both positive and negative effects on freshwater chemistry. Nutrient and carbon enrichment will enhance nutrient cycling and productivity, and alter the generation and consumption of carbon-based trace gases. Consequently, the status of aquatic ecosystems as carbon sinks or sources is very likely to change. Climate change will also very likely affect the biodiversity of freshwater ecosystems across most of the Arctic. The magnitude, extent, and duration of the impacts and responses will be system- and location-dependent. Projected effects on aquatic mammals and waterfowl include altered migration routes and timing; a possible increase in the incidence of mortality and decreased growth and productivity from disease and/or parasites; and, probable changes in habitat suitability and timing of availability.

General effects of climate change on Arctic fishes and fish populations.

Projected shifts in climate forcing variables such as temperature and precipitation are of great relevance to arctic freshwater ecosystems and biota. These will result in many direct and indirect effects upon the ecosystems and fish present therein. Shifts projected for fish populations will range from positive to negative in overall effect, differ among species and also among populations within species depending upon their biology and tolerances, and will be integrated by the fish within their local aquascapes. This results in a wide range of future possibilities for arctic freshwater and diadromous fishes. Owing to a dearth of basic knowledge regarding fish biology and habitat interactions in the north, complicated by scaling issues and uncertainty in future climate projections, only qualitative scenarios can be developed in most cases. This limits preparedness to meet challenges of climate change in the Arctic with respect to fish and fisheries.

An overview of effects of climate change on selected arctic freshwater and anadromous fishes.

Arctic freshwater and diadromous fish species will respond to the various effects of climate change in many ways. For wide-ranging species, many of which are key components of northern aquatic ecosystems and fisheries, there is a large range of possible responses due to inter- and intra-specific variation, differences in the effects of climate drivers within ACIA regions, and differences in drivers among regions. All this diversity, coupled with limited understanding of fish responses to climate parameters generally, permits enumeration only of a range of possible responses which are developed here for selected important fishes. Accordingly, in-depth examination is required of possible effects within species within ACIA regions, as well as comparative studies across regions. Two particularly important species (Arctic char and Atlantic salmon) are examined as case studies to provide background for such studies.

Effects of ultraviolet radiation and contaminant-related stressors on arctic freshwater ecosystems.

Climate change is likely to act as a multiple stressor, leading to cumulative and/or synergistic impacts on aquatic systems. Projected increases in temperature and corresponding alterations in precipitation regimes will enhance contaminant influxes to aquatic systems, and independently increase the susceptibility of aquatic organisms to contaminant exposure and effects. The consequences for the biota will in most cases be additive (cumulative) and multiplicative (synergistic). The overall result will be higher contaminant loads and biomagnification in aquatic ecosystems. Changes in stratospheric ozone and corresponding ultraviolet radiation regimes are also expected to produce cumulative and/or synergistic effects on aquatic ecosystem structure and function. Reduced ice cover is likely to have a much greater effect on underwater UV radiation exposure than the projected levels of stratospheric ozone depletion. A major increase in UV radiation levels will cause enhanced damage to organisms (biomolecular, cellular, and physiological damage, and alterations in species composition). Allocations of energy and resources by aquatic biota to UV radiation protection will increase, probably decreasing trophic-level productivity. Elemental fluxes will increase via photochemical pathways.

Effects of climate change and UV radiation on fisheries for arctic freshwater and anadromous species.

Fisheries for arctic freshwater and diadromous fish species contribute significantly to northern economies. Climate change, and to a lesser extent increased ultraviolet radiation, effects in freshwaters will have profound effects on fisheries from three perspectives: quantity of fish available, quality of fish available, and success of the fishers. Accordingly, substantive adaptation will very likely be required to conduct fisheries sustainably in the future as these effects take hold. A shift to flexible and rapidly responsive 'adaptive management' of commercial fisheries will be necessary; local land- and resource-use patterns for subsistence fisheries will change; and, the nature, management and place for many recreational fisheries will change. Overall, given the complexity and uncertainty associated with climate change and related effects on arctic freshwaters and their biota, a much more conservative approach to all aspects of fishery management will be required to ensure ecosystems and key fished species retain sufficient resiliency and capacity to meet future changes.

Biological and life-history factors affecting total mercury concentrations in Arctic charr from Heintzelman Lake, Ellesmere Island, Nunavut.

A snapshot sample of Arctic charr (Salvelinus alpinus) from Heintzelman Lake (81°42'N, 66°56'W), Ellesmere Island, Canada was used to elucidate the biological and life-history factors potentially influencing individual total mercury (THg) concentrations. Migratory history was significant, with anadromous fish having a lower mean THg concentration (64 µg/kg ww) than the non-anadromous Arctic charr (117 µg/kg ww). The increase in individual THg concentration with age was shown to be independent of length-at-age when large and small individuals within the same age groups were compared. Similarly, the diets of individual Arctic charr were comparable regardless of size, and there was no apparent ontogenetic shift in diet that could explain differences in length-at-age or THg concentration among fast- and slow-growing groups of fish (i.e., fish of the same age but differing sizes). Maturity state was also not related to THg concentration, but appears to be related to differences in length-at-age, with slow-growing fish allocating more energy to reproduction than fast-growing conspecifics. The differences in THg concentration among individual Arctic charr were best explained by fish age. We suggest that the increase in mercury concentration with age can be altered by a shift in diet (e.g., to piscivory) or habitat (e.g., anadromy), but is otherwise unaffected by changes in size or length-at-age.