Development and calibration of a high dynamic range and autonomous ocean-light instrument to measure sub-surface profiles in ice-covered waters.

The optical chain and logger (OptiCAL) is an autonomous ice-tethered observatory equipped with multiple light sensors for mapping the variation of light with depth. We describe the instrument and present an ensemble calibration for downwelling irradiance E P A R in [µm o l m -2 s -1]. Results from a long-term deployment in the Arctic Ocean demonstrate that the OptiCAL can cover the high dynamic range of under-ice light levels from July to November and produce realistic values in terms of magnitude when compared to modeled surface irradiance. Transient features of raised light levels at specific depths associated with nearby leads in the ice underline the importance of depth-resolved light measurements.

Modelling the biogeographic boundary shift of Calanus finmarchicus reveals drivers of Arctic Atlantification by subarctic zooplankton.

Biological communities in the Arctic are changing through the climate-driven encroachment of subarctic species. This "Atlantification" extends to keystone Calanoid copepods, as the small-bodied Calanus finmarchicus increases in abundance in areas where it overlaps with larger Arctic congeners. The environmental factors that are facilitating this shift, whether related to optimal conditions in temperature or seasonality, remain unclear. Assessing these drivers at an Arctic-wide scale is necessary to predict future ecosystem change and impacts. Here we have compiled range-wide occurrences of C. finmarchicus and a suite of seasonal biophysical climatologies to build a boreo-Arctic ecological niche model. The data set was divided into two eras, 1955-1984 and 1985-2017, and an optimized MaxEnt model was used to predict the seasonal distribution of the abiotic niche of C. finmarchicus in both eras. Comparing outputs between eras reveals an increase in habitat suitability at the Arctic range edge. Large and significant increases in suitability are predicted in the regions of the Greenland, Labrador, and Southern Barents Seas that have experienced reduced sea-ice cover. With the exception of the Barents Sea, these areas also show a seasonal shift in the timing of peak habitat suitability toward an earlier season. Our findings suggest that the Atlantification of Arctic zooplankton communities is accompanied by climate-driven phenology changes. Although seasonality is a critical constraint to the establishment of C. finmarchicus at Arctic latitudes, earlier sea-ice retreat and associated productivity is making these environments increasingly favorable for this subarctic species.

Can a key boreal Calanus copepod species now complete its life-cycle in the Arctic? Evidence and implications for Arctic food-webs.

The changing Arctic environment is affecting zooplankton that support its abundant wildlife. We examined how these changes are influencing a key zooplankton species, Calanus finmarchicus, principally found in the North Atlantic but expatriated to the Arctic. Close to the ice-edge in the Fram Strait, we identified areas that, since the 1980s, are increasingly favourable to C. finmarchicus. Field-sampling revealed part of the population there to be capable of amassing enough reserves to overwinter. Early developmental stages were also present in early summer, suggesting successful local recruitment. This extension to suitable C. finmarchicus habitat is most likely facilitated by the long-term retreat of the ice-edge, allowing phytoplankton to bloom earlier and for longer and through higher temperatures increasing copepod developmental rates. The increased capacity for this species to complete its life-cycle and prosper in the Fram Strait can change community structure, with large consequences to regional food-webs.

Photophysiological cycles in Arctic krill are entrained by weak midday twilight during the Polar Night.

Light plays a fundamental role in the ecology of organisms in nearly all habitats on Earth and is central for processes such as vision and the entrainment of the circadian clock. The poles represent extreme light regimes with an annual light cycle including periods of Midnight Sun and Polar Night. The Arctic Ocean extends to the North Pole, and marine light extremes reach their maximum extent in this habitat. During the Polar Night, traditional definitions of day and night and seasonal photoperiod become irrelevant since there are only "twilight" periods defined by the sun's elevation below the horizon at midday; we term this "midday twilight." Here, we characterize light across a latitudinal gradient (76.5° N to 81° N) during Polar Night in January. Our light measurements demonstrate that the classical solar diel light cycle dominant at lower latitudes is modulated during Arctic Polar Night by lunar and auroral components. We therefore question whether this particular ambient light environment is relevant to behavioral and visual processes. We reveal from acoustic field observations that the zooplankton community is undergoing diel vertical migration (DVM) behavior. Furthermore, using electroretinogram (ERG) recording under constant darkness, we show that the main migratory species, Arctic krill (Thysanoessa inermis) show endogenous increases in visual sensitivity during the subjective night. This change in sensitivity is comparable to that under exogenous dim light acclimations, although differences in speed of vision suggest separate mechanisms. We conclude that the extremely weak midday twilight experienced by krill at high latitudes during the darkest parts of the year has physiological and ecological relevance.

Autonomous Surface and Underwater Vehicles as Effective Ecosystem Monitoring and Research Platforms in the Arctic-The Glider Project.

Effective ocean management requires integrated and sustainable ocean observing systems enabling us to map and understand ecosystem properties and the effects of human activities. Autonomous subsurface and surface vehicles, here collectively referred to as "gliders", are part of such ocean observing systems providing high spatiotemporal resolution. In this paper, we present some of the results achieved through the project "Unmanned ocean vehicles, a flexible and cost-efficient offshore monitoring and data management approach-GLIDER". In this project, three autonomous surface and underwater vehicles were deployed along the Lofoten-Vesterålen (LoVe) shelf-slope-oceanic system, in Arctic Norway. The aim of this effort was to test whether gliders equipped with novel sensors could effectively perform ecosystem surveys by recording physical, biogeochemical, and biological data simultaneously. From March to September 2018, a period of high biological activity in the area, the gliders were able to record a set of environmental parameters, including temperature, salinity, and oxygen, map the spatiotemporal distribution of zooplankton, and record cetacean vocalizations and anthropogenic noise. A subset of these parameters was effectively employed in near-real-time data assimilative ocean circulation models, improving their local predictive skills. The results presented here demonstrate that autonomous gliders can be effective long-term, remote, noninvasive ecosystem monitoring and research platforms capable of operating in high-latitude marine ecosystems. Accordingly, these platforms can record high-quality baseline environmental data in areas where extractive activities are planned and provide much-needed information for operational and management purposes.

Pelagic organisms avoid white, blue, and red artificial light from scientific instruments.

In situ observations of pelagic fish and zooplankton with optical instruments usually rely on external light sources. However, artificial light may attract or repulse marine organisms, which results in biased measurements. It is often assumed that most pelagic organisms do not perceive the red part of the visible spectrum and that red light can be used for underwater optical measurements of biological processes. Using hull-mounted echosounders above an acoustic probe or a baited video camera, each equipped with light sources of different colours (white, blue and red), we demonstrate that pelagic organisms in Arctic and temperate regions strongly avoid artificial light, including visible red light (575-700 nm), from instruments lowered in the water column. The density of organisms decreased by up to 99% when exposed to artificial light and the distance of avoidance varied from 23 to 94 m from the light source, depending on colours, irradiance levels and, possibly, species communities. We conclude that observations from optical and acoustic instruments, including baited cameras, using light sources with broad spectral composition in the 400-700 nm wavelengths do not capture the real state of the ecosystem and that they cannot be used alone for reliable abundance estimates or behavioural studies.

Seasonal variability in non-consumptive mortality of Arctic zooplankton.

Recent observations from high-latitude marine ecosystems indicate that non-consumptive mortality may be particularly high in Arctic zooplankton during the polar night. Here we have estimated the contribution of dead organisms to the mesozooplankton community in the high Arctic (Svalbard 78-81oN) during the polar night (January), in spring (May) and in late summer (end of August). To identify in situ dead organisms, we used Neutral Red Stain. The dead zooplankton fraction consisted mainly of copepods, while the contribution of dead non-copepods was low in all seasons. The absolute abundance of dead copepods varied little between seasons; however, the relative contribution of dead copepods was highest in January with 11-35% of the copepods classified as dead, in contrast to 2-12% in spring and summer. Furthermore, there were species-specific differences: copepods of the genus Calanus contributed more to the dead fraction of the copepod community during the polar night compared to spring and summer, leading to a higher "dead" biomass in winter. We conclude that non-consumptive winter mortality is considerable in calanoid copepods in the Arctic and an important but so far neglected component of the passive carbon flux, providing carbon in larger portions for higher trophic level consumers during the low-productive winter.

Artificial light during the polar night disrupts Arctic fish and zooplankton behaviour down to 200 m depth.

For organisms that remain active in one of the last undisturbed and pristine dark environments on the planet-the Arctic Polar Night-the moon, stars and aurora borealis may provide important cues to guide distribution and behaviours, including predator-prey interactions. With a changing climate and increased human activities in the Arctic, such natural light sources will in many places be masked by the much stronger illumination from artificial light. Here we show that normal working-light from a ship may disrupt fish and zooplankton behaviour down to at least 200 m depth across an area of >0.125 km2 around the ship. Both the quantitative and qualitative nature of the disturbance differed between the examined regions. We conclude that biological surveys in the dark from illuminated ships may introduce biases on biological sampling, bioacoustic surveys, and possibly stock assessments of commercial and non-commercial species.

Pelagic occurrences of the ice amphipod Apherusa glacialis throughout the Arctic.

Apherusa glacialis is a common, sea ice-associated amphipod found throughout the Arctic Ocean and has long been considered permanently associated with the sea ice habitat. However, pelagic occurrences of A. glacialis have also been reported. It was recently suggested that A. glacialis overwinters at depth within the Atlantic-water inflow near Svalbard, to avoid being exported out of the Arctic Ocean through the Fram Strait. This study collated pelagic occurrence records over a 71-year period and found that A. glacialis was consistently found away from its presumed sea ice habitat on a pan-Arctic scale, in different depths and water masses. In the Svalbard region, A. glacialis was found in Atlantic Water both in winter and summer. Additionally, we analyzed A. glacialis size distributions throughout the year, collected mostly from sea ice, in order to elucidate potential life cycle strategies. The majority of young-of-the-year A. glacialis was found in the sea ice habitat during spring, supporting previous findings. Data on size distributions and sex ratios suggest a semelparous lifestyle. A synchronous seasonal vertical migration was not evident, but our data imply a more complex life history than previously assumed. We provide evidence that A. glacialis can no longer be regarded as an autochthonous sympagic species.

Remote sensing of zooplankton swarms.

Zooplankton provide the key link between primary production and higher levels of the marine food web and they play an important role in mediating carbon sequestration in the ocean. All commercially harvested fish species depend on zooplankton populations. However, spatio-temporal distributions of zooplankton are notoriously difficult to quantify from ships. We know that zooplankton can form large aggregations that visibly change the color of the sea, but the scale and mechanisms producing these features are poorly known. Here we show that large surface patches (>1000 km2) of the red colored copepod Calanus finmarchicus can be identified from satellite observations of ocean color. Such observations provide the most comprehensive view of the distribution of a zooplankton species to date, and alter our understanding of the behavior of this key zooplankton species. Moreover, our findings suggest that high concentrations of astaxanthin-rich zooplankton can degrade the performance of standard blue-green reflectance ratio algorithms in operational use for retrieving chlorophyll concentrations from ocean color remote sensing.

Seasonal ecology in ice-covered Arctic seas - Considerations for spill response decision making.

Due to retreating sea ice and predictions of undiscovered oil and gas resources, increased activity in Arctic shelf sea areas associated with shipping and oil and gas exploration is expected. Such activities may accidentally lead to oil spills in partly ice-covered ocean areas, which raises issues related to oil spill response. Net Environmental Benefit Analysis (NEBA) is the process that the response community uses to identify which combination of response strategies minimises the impact to environment and people. The vulnerability of Valued Ecosystem Components (VEC's) to oil pollution depends on their sensitivity to oil and the likelihood that they will be exposed to oil. As such, NEBA requires a good ecological knowledge base on biodiversity, species' distributions in time and space, and timing of ecological events. Biological resources found at interfaces (e.g., air/water, ice/water or water/coastline) are in general vulnerable because that is where oil can accumulate. Here, we summarize recent information about the seasonal, physical and ecological processes in Arctic waters and evaluate the importance these processes when considering in oil spill response decision making through NEBA. In spring-time, many boreal species conduct a lateral migration northwards in response to sea ice retraction and increased production associated with the spring bloom. However, many Arctic species, including fish, seabirds and marine mammals, are present in upper water layers in the Arctic throughout the year, and recent research has demonstrated that bioactivity during the Arctic winter is higher than previously assumed. Information on the seasonal presence/absence of less resilient VEC's such as marine mammals and sea birds in combination with the presence/absence of sea ice seems to be especially crucial to consider in a NEBA. In addition, quantification of the potential impact of different, realistic spill sizes on the energy cascade following the spring bloom at the ice-edge would provide important information for assessing ecosystem effects.

Use of an Autonomous Surface Vehicle reveals small-scale diel vertical migrations of zooplankton and susceptibility to light pollution under low solar irradiance.

Light is a major cue for nearly all life on Earth. However, most of our knowledge concerning the importance of light is based on organisms' response to light during daytime, including the dusk and dawn phase. When it is dark, light is most often considered as pollution, with increasing appreciation of its negative ecological effects. Using an Autonomous Surface Vehicle fitted with a hyperspectral irradiance sensor and an acoustic profiler, we detected and quantified the behavior of zooplankton in an unpolluted light environment in the high Arctic polar night and compared the results with that from a light-polluted environment close to our research vessels. First, in environments free of light pollution, the zooplankton community is intimately connected to the ambient light regime and performs synchronized diel vertical migrations in the upper 30 m despite the sun never rising above the horizon. Second, the vast majority of the pelagic community exhibits a strong light-escape response in the presence of artificial light, observed down to 100 m. We conclude that artificial light from traditional sampling platforms affects the zooplankton community to a degree where it is impossible to examine its abundance and natural rhythms within the upper 100 m. This study underscores the need to adjust sampling platforms, particularly in dim-light conditions, to capture relevant physical and biological data for ecological studies. It also highlights a previously unchartered susceptibility to light pollution in a region destined to see significant changes in light climate due to a reduced ice cover and an increased anthropogenic activity.

Genetics redraws pelagic biogeography of Calanus.

Planktonic copepods of the genus Calanus play a central role in North Atlantic/Arctic marine food webs. Here, using molecular markers, we redrew the distributional ranges of Calanus species inhabiting the North Atlantic and Arctic Oceans and revealed much wider and more broadly overlapping distributions than previously described. The Arctic shelf species, C. glacialis, dominated the zooplankton assemblage of many Norwegian fjords, where only C. finmarchicus has been reported previously. In these fjords, high occurrences of the Arctic species C. hyperboreus were also found. Molecular markers revealed that the most common method of species identification, prosome length, cannot reliably discriminate the species in Norwegian fjords. Differences in degree of genetic differentiation among fjord populations of the two species suggested that C. glacialis is a more permanent resident of the fjords than C. finmarchicus We found no evidence of hybridization between the species. Our results indicate a critical need for the wider use of molecular markers to reliably identify and discriminate these morphologically similar copepod species, which serve as important indicators of climate responses.

Unexpected Levels of Biological Activity during the Polar Night Offer New Perspectives on a Warming Arctic.

The current understanding of Arctic ecosystems is deeply rooted in the classical view of a bottom-up controlled system with strong physical forcing and seasonality in primary-production regimes. Consequently, the Arctic polar night is commonly disregarded as a time of year when biological activities are reduced to a minimum due to a reduced food supply. Here, based upon a multidisciplinary ecosystem-scale study from the polar night at 79°N, we present an entirely different view. Instead of an ecosystem that has entered a resting state, we document a system with high activity levels and biological interactions across most trophic levels. In some habitats, biological diversity and presence of juvenile stages were elevated in winter months compared to the more productive and sunlit periods. Ultimately, our results suggest a different perspective regarding ecosystem function that will be of importance for future environmental management and decision making, especially at a time when Arctic regions are experiencing accelerated environmental change [1].

Is Ambient Light during the High Arctic Polar Night Sufficient to Act as a Visual Cue for Zooplankton?

The light regime is an ecologically important factor in pelagic habitats, influencing a range of biological processes. However, the availability and importance of light to these processes in high Arctic zooplankton communities during periods of 'complete' darkness (polar night) are poorly studied. Here we characterized the ambient light regime throughout the diel cycle during the high Arctic polar night, and ask whether visual systems of Arctic zooplankton can detect the low levels of irradiance available at this time. To this end, light measurements with a purpose-built irradiance sensor and coupled all-sky digital photographs were used to characterize diel skylight irradiance patterns over 24 hours at 79°N in January 2014 and 2015. Subsequent skylight spectral irradiance and in-water optical property measurements were used to model the underwater light field as a function of depth, which was then weighted by the electrophysiologically determined visual spectral sensitivity of a dominant high Arctic zooplankter, Thysanoessa inermis. Irradiance in air ranged between 1-1.5 x 10-5 µmol photons m-2 s-1 (400-700 nm) in clear weather conditions at noon and with the moon below the horizon, hence values reflect only solar illumination. Radiative transfer modelling generated underwater light fields with peak transmission at blue-green wavelengths, with a 465 nm transmission maximum in shallow water shifting to 485 nm with depth. To the eye of a zooplankter, light from the surface to 75 m exhibits a maximum at 485 nm, with longer wavelengths (>600 nm) being of little visual significance. Our data are the first quantitative characterisation, including absolute intensities, spectral composition and photoperiod of biologically relevant solar ambient light in the high Arctic during the polar night, and indicate that some species of Arctic zooplankton are able to detect and utilize ambient light down to 20-30m depth during the Arctic polar night.

Floating ice-algal aggregates below melting arctic sea ice.

During two consecutive cruises to the Eastern Central Arctic in late summer 2012, we observed floating algal aggregates in the melt-water layer below and between melting ice floes of first-year pack ice. The macroscopic (1-15 cm in diameter) aggregates had a mucous consistency and were dominated by typical ice-associated pennate diatoms embedded within the mucous matrix. Aggregates maintained buoyancy and accumulated just above a strong pycnocline that separated meltwater and seawater layers. We were able, for the first time, to obtain quantitative abundance and biomass estimates of these aggregates. Although their biomass and production on a square metre basis was small compared to ice-algal blooms, the floating ice-algal aggregates supported high levels of biological activity on the scale of the individual aggregate. In addition they constituted a food source for the ice-associated fauna as revealed by pigments indicative of zooplankton grazing, high abundance of naked ciliates, and ice amphipods associated with them. During the Arctic melt season, these floating aggregates likely play an important ecological role in an otherwise impoverished near-surface sea ice environment. Our findings provide important observations and measurements of a unique aggregate-based habitat during the 2012 record sea ice minimum year.