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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 km 2 ) 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.

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.

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.

The traditional view is that the Arctic polar night is a quiescent period for marine life, but recent reports of high levels of feeding and reproduction in both pelagic and benthic taxa have challenged this. We examined the zooplankton community present in Svalbard fjords, coastal waters, and the shelf break north of Svalbard, during the polar night. We focused on the population structure of abundant copepods (Calanus finmarchicus, Calanus glacialis, Metridia longa, Oithona similis, Pseudocalanus spp., Microcalanus spp., and Microsetella norvegica) sampled using 64-µm mesh nets. Numerically, copepod nauplii (≥ 50%) and the young developmental stages of small copepods (< 2 mm prosome length as adult) dominated the samples. Three main patterns were identified: (1) large Calanus spp. were predominantly older copepodids CIV–CV, while (2) the small harpacticoid M. norvegica were adults. (3) For other species, all copepodid stages were present. Older copepodids and adults dominated populations of O. similis, Pseudocalanus spp. and M. longa. In Microcalanus spp., high proportion of young copepodids CI–CIII indicated active winter recruitment. We discuss the notion of winter as a developing and reproductive period for small copepods in light of observed age structures, presence of nauplii, and previous knowledge about the species. Lower predation risks during winter may, in part, explain why this season could be beneficial as a period for development. Winter may be a key season for development of small, omnivorous copepods in the Arctic, whereas large copepods such as Calanus spp. seems to be reliant on spring and summer for reproduction and development.

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.

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 km 2 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. Berge et al. find that the normal working-light from a ship impacts on the vertical distribution of macrozooplankton and pelagic fish communities around the ship at three stations during the Arctic Polar Night. These data suggest bias from such effects should be taken into account when performing surveys and stock assessments in the Arctic.

Seasonal patterns in mesozooplankton composition, vertical distribution, and timing of reproduction are challenging to study in the open sea due to ocean currents and mix of populations of different origins. Sill fjords, on the other hand, with restricted water exchange, are ideal locations for studying taxa- and community-specific adaptations to the prevailing environment. Here, we present re-occurring patterns in the mesozooplankton community structure in Billefjorden, Svalbard, a high Arctic sill fjord with extensive seasonal ice cover, based on monthly sampling from 2011 to 2013. The zooplankton community composition confirmed the Arctic character of this fjord. Predominantly herbivorous taxa, such as Calanus glacialis and Pseudocalanus spp., showed strong seasonal variation in abundance and depth distribution, with population minima in spring being compensated by a rapid population recovery during summer. Omnivorous taxa, such as Microcalanus spp. and copepods of the family Aetideidae , largely remained at depth throughout the year and had an extended or year-round reproductive period. Deep-dwelling omnivorous/carnivorous species peaked in abundance in winter–spring when herbivorous populations were severely depleted. Taxa with seasonally limited occurrences, i.e., meroplankton, peaked in spring and summer at the surface, but were largely absent for the rest of the year. The different life histories, with contrasting feeding modes, depth preferences, and timing of reproduction lead to reduced interspecies competition and allow for a rather high and stable abundance of mesozooplankton year-round despite the short primary production window at high latitudes.

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.

The calanoid copepod Calanus glacialis dominates the mesozooplankton biomass in the Arctic shelf seas, but its smaller North Atlantic sibling Calanus finmarchicus is expanding northwards and may potentially replace it if the climate continues to warm. Here we studied the population structure, overwintering strategies, gonad maturation and egg production of C. glacialis and C. finmarchicus over a period of 15 consecutive months in a high-Arctic fjord with sub-Arctic ocean climate and no sea ice formation in winter. The relative proportions of C. glacialis and C. finmarchicus varied throughout the study period, but with an overall dominance of C. glacialis . The overwintering population of C. glacialis was dominated by copepodite stage CIV (74%) while C. finmarchicus overwintered mainly as CV (65%), reflecting a primarily two- and one-year life cycle, respectively. Adult males and females of C. glacialis appeared as early as October with a peak during December-January, two months earlier than in C. finmarchicus , with a corresponding one-month earlier peak in recruitment for C. glacialis. While C. glacialis reproduced prior to the bloom with egg production peaking during the bloom , C. finmarchicus started egg laying during the bloom and continued to reproduce throughout the summer. Seasonal changes in the population structure suggest that C. finmarchicus born early in spring are able to develop to CV during summer and overwinter successfully, while offspring born later in the season do most likely not reach the CV overwintering stage. The ability to reproduce early and the flexibility to alter between 1- and 2-year life cycles give C. glacialis an advantage over C. finmarchicus in high-Arctic unpredictable environments with short-pulsed primary production regimes. Our data indicate that C. glacialis and C. finmarchicus occupy similar environmental niches, but different timing in reproduction reduces the competition. If sea temperatures remain within their temperature-tolerance ranges, both C. glacialis and C. finmarchicus seem to benefit from warming due to accelerating growth and higher survival of the recruits as long as C. glacialis has access to a colder refuge by descending to deeper depths.

We investigated the distribution of mesozooplankton in waters north of Svalbard (north of 78°50′N) at 38 stations in August and September of 2002, 2003 and 2004. The zooplankton community was numerically dominated by copepods (58–98% of the total abundance). Zooplankton abundance ranged from 115 individuals m−3 at the northern most location to 12,296 individuals m−3 on the shelf. Cluster analysis revealed four different groups with distinct geographic integrity that were identified by variation in species densities rather than by variation in taxonomic composition. Water temperature and salinity differed significantly between the different cluster groups indicating that part of the observed variations in species distribution relate to differences in hydrography. Numerous significant regressions between zooplankton abundance at species level and hydrographical parameters suggest that variability in water masses has measurable effects on zooplankton distribution and species composition in the study area.