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Author's accepted version (postprint).This is an Accepted Manuscript of an article published by Wiley in Molecular Ecology Resources on 30/06/2020.Available online: https://onlinelibrary.wiley.com/doi/epdf/10.1111/1755-0998.13218acceptedVersio

No evidence for hybridization between Calanus finmarchicus and Calanus glacialis in a subarctic area of sympatry

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The complete mitochondrial genome of the copepod Calanus glacialis

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Seasonal Enzyme Activities of Sympatric Calanus glacialis and C. finmarchicus in the High-Arctic

In the Arctic shelf seas, the mesozooplankton biomass is dominated by the arctic copepod Calanus glacialis, but its boreal congeneric C. finmarchicus is expanding northwards. Even though it is already there, C. finmarchicus may not be able to truly establish itself in the Arctic seas and potentially replace C. glacialis. We compared metabolic and digestive enzyme activities of sympatric C. glacialis and C. finmarchicus from Isfjorden, Svalbard and off-shelf north of Svalbard. The seasonal regulation of anabolic and catabolic enzyme activities was generally similar for the two species, but with some interspecific differences corresponding to their ontogeny. Wake-up from overwintering started earlier in adults of C. glacialis than in C. finmarchicus, while the onset of dormancy started early in the overwintering stages of both species. Furthermore, C. glacialis showed an earlier and higher mobilization of lipase enzyme activities, indicating higher efficiency in assimilating dietary lipids compared to C. finmarchicus. Similar population sizes and population structures for C. finmarchicus off-shelf north of Svalbard and in Isfjorden support a similar origin. Still, C. finmarchicus was able to match regulation of enzyme activities to the bloom even though the bloom peaked approximately a month later off-shelf north of Svalbard, indicating that food availability is an important signal for the final step of termination of diapause. Even though the two species largely follow the same patterns of metabolic enzyme activities, the more efficient lipid anabolism of C. glacialis may give it an advantage over C. finmarchicus in high-Arctic unpredictable environments with short-pulsed primary production regimes.publishedVersio

Photoperiodism and overwintering in boreal and sub-Arctic Calanus finmarchicus populations

The copepod Calanus finmarchicus, a key species in the North Atlantic, generally spends the non-productive season by descending into deep waters and entering diapause, a physiological state characterized by reduced metabolism and arrested development. In the open ocean, overwintering depths are below 600 m, where temperature and light conditions are favourable to initiate diapause. However, C. finmarchicus has also been reported diapausing in areas with shallow water depth such as fjords, coastal waters and shelf seas. In these environments, the temperature and light conditions are different, and it has been hypothesized that under such conditions C. finmarchicus may remain active throughout winter. Here, we investigated changes in the swimming activity of C. finmarchicus from shallow fjords in the eastern North Atlantic during overwintering in response to ambient photoperiod. We conducted monthly experiments with populations from 2 fjords from different latitudes (sub-Arctic Ramfjord, 69°N and boreal Loch Etive, 56°N), measuring the locomotor activity of individual C. finmarchicus stage CVs exposed to a natural light:dark cycle. At both locations, peaks in activity in response to the light cycle were observed to shift from nocturnal during the early overwintering phase to diurnal during mid and late overwintering phase, with a minimal intensity observed during the mid-overwintering phase. In Ramfjord, activity and rhythmicity were generally lower than in Loch Etive. We conclude that C. finmarchicus remains active throughout its overwintering period when in shallow (<200 m) locations but down-regulates its locomotor activity during the main overwintering phase, which we describe as a winter resting state as distinct from classical diapause

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 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.publishedVersio

The β-oxidation pathway is downregulated during diapause termination in Calanus copepods

Calanus copepods are keystone species in marine ecosystems, mainly due to their high lipid content, which is a nutritious food source for e.g. juvenile fish. Accumulated lipids are catabolized to meet energy requirements during dormancy (diapause), which occurs during the last copepodite stage (C5). The current knowledge of lipid degradation pathways during diapause termination is limited. We characterized changes in lipid fullness and generated transcriptional profiles in C5s during termination of diapause and progression towards adulthood. Lipid fullness of C5s declined linearly during developmental progression, but more β-oxidation genes were upregulated in early C5s compared to late C5s and adults. We identified four possible master regulators of energy metabolism, which all were generally upregulated in early C5s, compared to late C5s and adults. We discovered that one of two enzymes in the carnitine shuttle is absent from the calanoid copepod lineage. Based on the geographical location of the sampling site, the field-samples were initially presumed to consist of C. finmarchicus. However, the identification of C. glacialis in some samples underlines the need for performing molecular analyses to reliably identify Calanus species. Our findings contributes to a better understanding of molecular events occurring during diapause and diapause termination in calanoid copepods.publishedVersio

Towards population genomics in non-model species with large genomes : a case study of the marine zooplankton Calanus finmarchicus

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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.publishedVersionPaid Open Acces

Body burdens of PAHs and alkylated phenols in copepods in the Ekofisk region - Water column monitoring 2021

This report summarizes the findings from the Water Column Monitoring in April 2021 where zooplankton (Calanus finmarchicus) were collected and sampled around the Ekofisk region in the North Sea. Extraction and analyses of polycyclic aromatic hydrocarbons and alkylated phenols in copepod were determined to provide data for comparison to DREAM-MER simulations.Body burdens of PAHs and alkylated phenols in copepods in the Ekofisk region - Water column monitoring 2021publishedVersio