Insights into the diet and feeding behavior of immature polar cod (Boreogadus saida) from the under-ice habitat of the central Arctic Ocean.

Polar cod (Boreogadus saida) is an endemic key species of the Arctic Ocean ecosystem. The ecology of this forage fish is well studied in Arctic shelf habitats where a large part of its population lives. However, knowledge about its ecology in the central Arctic Ocean (CAO), including its use of the sea-ice habitat, is hitherto very limited. To increase this knowledge, samples were collected at the under-ice surface during several expeditions to the CAO between 2012 and 2020, including the Multidisciplinary drifting Observatory for the Study of Arctic Climate (MOSAiC) expedition. The diet of immature B. saida and the taxonomic composition of their potential prey were analysed, showing that both sympagic and pelagic species were important prey items. Stomach contents included expected prey such as copepods and amphipods. Surprisingly, more rarely observed prey such as appendicularians, chaetognaths, and euphausiids were also found to be important. Comparisons of the fish stomach contents with prey distribution data suggests opportunistic feeding. However, relative prey density and catchability are important factors that determine which type of prey is ingested. Prey that ensures limited energy expenditure on hunting and feeding is often found in the stomach contents even though it is not the dominant species present in the environment. To investigate the importance of prey quality and quantity for the growth of B. saida in this area, we measured energy content of dominant prey species and used a bioenergetic model to quantify the effect of variations in diet on growth rate potential. The modeling results suggest that diet variability was largely explained by stomach fullness and, to a lesser degree, the energetic content of the prey. Our results suggest that under climate change, immature B. saida may be at least equally sensitive to a loss in the number of efficiently hunted prey than to a reduction in the prey's energy content. Consequences for the growth and survival of B. saida will not depend on prey presence alone, but also on prey catchability, digestibility, and energy content.

Global variability in seawater Mg:Ca and Sr:Ca ratios in the modern ocean.

Seawater Mg:Ca and Sr:Ca ratios are biogeochemical parameters reflecting the Earth-ocean-atmosphere dynamic exchange of elements. The ratios' dependence on the environment and organisms' biology facilitates their application in marine sciences. Here, we present a measured single-laboratory dataset, combined with previous data, to test the assumption of limited seawater Mg:Ca and Sr:Ca variability across marine environments globally. High variability was found in open-ocean upwelling and polar regions, shelves/neritic and river-influenced areas, where seawater Mg:Ca and Sr:Ca ratios range from ∼4.40 to 6.40 mmol:mol and ∼6.95 to 9.80 mmol:mol, respectively. Open-ocean seawater Mg:Ca is semiconservative (∼4.90 to 5.30 mol:mol), while Sr:Ca is more variable and nonconservative (∼7.70 to 8.80 mmol:mol); both ratios are nonconservative in coastal seas. Further, the Ca, Mg, and Sr elemental fluxes are connected to large total alkalinity deviations from International Association for the Physical Sciences of the Oceans (IAPSO) standard values. Because there is significant modern seawater Mg:Ca and Sr:Ca ratios variability across marine environments we cannot absolutely assume that fossil archives using taxa-specific proxies reflect true global seawater chemistry but rather taxa- and process-specific ecosystem variations, reflecting regional conditions. This variability could reconcile secular seawater Mg:Ca and Sr:Ca ratio reconstructions using different taxa and techniques by assuming an error of 1 to 1.50 mol:mol, and 1 to 1.90 mmol:mol, respectively. The modern ratios' variability is similar to the reconstructed rise over 20 Ma (Neogene Period), nurturing the question of seminonconservative behavior of Ca, Mg, and Sr over modern Earth geological history with an overlooked environmental effect.

Biogeographic responses of the copepod Calanus glacialis to a changing Arctic marine environment.

Dramatic changes have occurred in the Arctic Ocean over the past few decades, especially in terms of sea ice loss and ocean warming. Those environmental changes may modify the planktonic ecosystem with changes from lower to upper trophic levels. This study aimed to understand how the biogeographic distribution of a crucial endemic copepod species, Calanus glacialis, may respond to both abiotic (ocean temperature) and biotic (phytoplankton prey) drivers. A copepod individual-based model coupled to an ice-ocean-biogeochemical model was utilized to simulate temperature- and food-dependent life cycle development of C. glacialis annually from 1980 to 2014. Over the 35-year study period, the northern boundaries of modeled diapausing C. glacialis expanded poleward and the annual success rates of C. glacialis individuals attaining diapause in a circumpolar transition zone increased substantially. Those patterns could be explained by a lengthening growth season (during which time food is ample) and shortening critical development time (the period from the first feeding stage N3 to the diapausing stage C4). The biogeographic changes were further linked to large-scale oceanic processes, particularly diminishing sea ice cover, upper ocean warming, and increasing and prolonging food availability, which could have potential consequences to the entire Arctic shelf/slope marine ecosystems.

Pushing the limit: Resilience of an Arctic copepod to environmental fluctuations.

Life history strategies such as multiyear life cycles, resting stages, and capital breeding allow species to inhabit regions with extreme and fluctuating environmental conditions. One example is the zooplankton species Calanus hyperboreus, whose life history is considered an adaptation to the short and unpredictable growth season in the central Arctic Ocean. This copepod is commonly described as a true Arctic endemic; however, by statistically analyzing compiled observational data, we show that abundances are relatively low and later stages and adults dominate in the central Arctic Ocean basins, indicating expatriation. Combining data analyses with individual-based modeling and energy requirement estimation, we further demonstrate that while C. hyperboreus can reach higher abundances in areas with greater food availability outside the central Arctic basins, the species' resilience to environmental fluctuations enables the life cycle to be completed in the central Arctic basins. Specifically, the energy level required to reach the first overwintering stage-a prerequisite for successful local production-is likely met in some-but not all-years. This fine balance between success and failure indicates that C. hyperboreus functions as a peripheral population in the central Arctic basins and its abundance will likely increase in areas with improved growth conditions in response to climate change. By illustrating a key Arctic species' resilience to extreme and fluctuating environmental conditions, the results of this study have implications for projections of future biogeography and food web dynamics in the Arctic Ocean, a region experiencing rapid warming and sea ice loss.

Alexandrium on the Alaskan Beaufort Sea shelf: Impact of upwelling in a warming Arctic.

The harmful algal genus Alexandrium has characteristically been found in temperate and subtropical regions; however recent evidence suggests global warming may be expanding its range into high latitude waters. Alexandrium cysts have previously been documented in the Chukchi Sea and we hypothesize that Alexandrium may be expanding further into the Arctic due to distribution by the Beaufort shelfbreak jet. Here we document the presence of Alexandrium catenella along the Alaskan Beaufort Sea shelf, marking an expansion of its known range. The observations of A. catenella were made using three different methods: FlowCAM imaging, 18S eukaryotic sequencing, and real-time quantitative PCR. Four occupations of a shelf/slope transect spanned the evolution of a strong wind-driven upwelling event over a 5-day period. A nearby mooring provided the physical context for the event, revealing that enhanced easterly winds reversed the Beaufort shelfbreak jet to the west and induced upwelling of colder, denser water onto the outer shelf. A. catenella sequences dominated the surface phytoplankton community at the onset of the upwelling event. This signal vanished during and after the event, likely due to a combination of alongstream advection, cross-stream advection, and wind mixing. These results suggest contrasting physical processes that are both subject to global warming amplification, delivery of warm waters via the Beaufort shelfbreak jet and upwelling, may control the proliferation of this potential harmful alga into the Arctic.

Lingering Chukchi Sea sea ice and Chukchi Sea mean winds influence population age structure of euphausiids (krill) found in the bowhead whale feeding hotspot near Pt. Barrow, Alaska.

Interannual variability in euphausiid (krill) abundance and population structure and associations of those measures with environmental drivers were investigated in an 11-year study conducted in late August-early September 2005-2015 in offshelf waters (bottom depth > 40 m) in Barrow Canyon and the Beaufort Sea just downstream of Distributed Biological Observatory site 5 (DBO5) near Pt. Barrow, Alaska. Statistically-significant positive correlations were observed among krill population structure (proportion of juveniles and adults), the volume of Late Season Melt Water (LMW), and late-spring Chukchi Sea sea ice extent. High proportions of juvenile and adult krill were seen in years with larger volumes of LMW and greater spring sea ice extents (2006, 2009, 2012-2014) while the converse, high proportions of furcilia, were seen in years with smaller volumes of LMW and lower spring sea ice extent (2005, 2007, 2010, 2011, 2015). These different life stage, sea ice and water mass regimes represent integrated advective responses to mean fall and/or spring Chukchi Sea winds, driven by prevailing atmospheric pressure distributions in the two sets of years. In years with high proportions of juveniles and adults, late-spring and preceding-fall winds were weak and variable while in years with high proportions of furcilia, late-spring and preceding-fall winds were strong, easterly and consistent. The interaction of krill life history with yearly differences in the northward transports of krill and water masses along with sea ice retreat determines the population structure of late-summer krill populations in the DBO5 region near Pt. Barrow. Years with higher proportions of mature krill may provide larger prey to the Pt. Barrow area bowhead whale prey hotspot. The characteristics of prey near Pt. Barrow is dependent on krill abundance and size, large-scale environmental forcing, and interannual variability in recruitment success of krill in the Bering Sea.

Pan-Arctic Depth Distribution of Diapausing Calanus Copepods.

Diapause at depth is considered an integral part of the life cycle of Calanus copepods, but few studies have focused on the Arctic species Calanus glacialis and Calanus hyperboreus. By analyzing a large set of pan-arctic observational data compiled from multiple sources, we show that Arctic Calanus has a broad depth distribution in winter, indicating that diapause at depth is a facultative strategy. Both species' vertical distributions tend to deepen in winter and to be deeper and broader with increasing bottom depth, while individuals are aggregated closer to the sea floor in shallow areas. These results indicate that Arctic Calanus species pursue a relatively deep diapause habitat but are topographically blocked on the shelves. Interspecific differences in depth distribution during diapause suggest the importance of predation. The larger C. hyperboreus has a deeper diapause depth than C. glacialis, potentially to alleviate predation pressure or as a result of predation loss near the surface. Moreover, the mean depth of C. hyperboreus in winter is negatively associated with latitude, indicating a shoaling of the diapause population in the central Arctic Ocean where predation pressure is lower. Our results suggest a complex diapause behavior by Arctic Calanus, with implications for our view of the species' roles in Arctic ecosystems.

The great 2012 Arctic Ocean summer cyclone enhanced biological productivity on the shelves.

[1] A coupled biophysical model is used to examine the impact of the great Arctic cyclone of early August 2012 on the marine planktonic ecosystem in the Pacific sector of the Arctic Ocean (PSA). Model results indicate that the cyclone influences the marine planktonic ecosystem by enhancing productivity on the shelves of the Chukchi, East Siberian, and Laptev seas during the storm. Although the cyclone's passage in the PSA lasted only a few days, the simulated biological effects on the shelves last 1 month or longer. At some locations on the shelves, primary productivity (PP) increases by up to 90% and phytoplankton biomass by up to 40% in the wake of the cyclone. The increase in zooplankton biomass is up to 18% on 31 August and remains 10% on 15 September, more than 1 month after the storm. In the central PSA, however, model simulations indicate a decrease in PP and plankton biomass. The biological gain on the shelves and loss in the central PSA are linked to two factors. (1) The cyclone enhances mixing in the upper ocean, which increases nutrient availability in the surface waters of the shelves; enhanced mixing in the central PSA does not increase productivity because nutrients there are mostly depleted through summer draw down by the time of the cyclone's passage. (2) The cyclone also induces divergence, resulting from the cyclone's low-pressure system that drives cyclonic sea ice and upper ocean circulation, which transports more plankton biomass onto the shelves from the central PSA. The simulated biological gain on the shelves is greater than the loss in the central PSA, and therefore, the production on average over the entire PSA is increased by the cyclone. Because the gain on the shelves is offset by the loss in the central PSA, the average increase over the entire PSA is moderate and lasts only about 10 days. The generally positive impact of cyclones on the marine ecosystem in the Arctic, particularly on the shelves, is likely to grow with increasing summer cyclone activity if the Arctic continues to warm and the ice cover continues to shrink.

Improved parameterization of Antarctic krill target strength models.

There are historical discrepancies between empirical observations of Antarctic krill target strength and predictions using theoretical scattering models. These differences are addressed through improved understanding of key model parameters. The scattering process was modeled using the distorted-wave Born approximation, representing the shape of the animal as a bent and tapered cylinder. Recently published length-based regressions were used to constrain the sound speed and density contrasts between the animal and the surrounding seawater, rather than the earlier approach of using single values for all lengths. To constrain the parameter governing the orientation of the animal relative to the incident acoustic wave, direct measurements of the orientation of krill in situ were made with a video plankton recorder. In contrast to previous indirect and aquarium-based observations, krill were observed to orient themselves mostly horizontally. Averaging predicted scattering over the measured distribution of orientations resulted in predictions of target strength consistent with in situ measurements of target strength of large krill (mean length 40-43 mm) at four frequencies (43-420 kHz), but smaller than expected under the semi-empirical model traditionally used to estimate krill target strength.