Tove M. Gabrielsen

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].

Potential misidentifications of two climate indicator species of the marine arctic ecosystem: Calanus glacialis and C. finmarchicus

Calanoid copepods of the genus Calanus represent an important, nutrient-rich food source for a multitude of Arctic marine organisms. Although morphologically very similar, their life histories and ecological roles differ. Because the distribution of Calanus glacialis and C. finmarchicus corresponds to Arctic and Atlantic water masses, respectively, they are regularly used as climate indicators. A correct identification of these species is therefore necessary in many ecological, environmental and climatological studies. In this study, we aimed at validating the traditionally used morphological characteristics (combining prosome length and copepodite stage) for separation of species of Calanus by using molecular tools (PCR–RFLP of the 16S mtDNA). A total of 418 specimens of copepodite stages CIV, CV and CVI(af) from three Arctic fjords have been identified both morphologically and genetically. We find that the morphological identification systematically overestimates the abundance of C. finmarchicus at the expense of C. glacialis. Hence, parts of the C. glacialis populations are found to be structurally smaller and the within population size range thus larger than previously assumed. Consequently, using the traditional morphological species delimitation poses a serious problem in the use of these two species as indicators of Atlantic versus Arctic water masses and thus as climatic indicators. Furthermore, it obscures our understanding of the life history differences between the two species and of their relative importance as food for a number of ecologically and economically important species in the Arctic.

Strong Seasonality of Marine Microbial Eukaryotes in a High-Arctic Fjord (Isfjorden, in West Spitsbergen, Norway)

ABSTRACT The Adventfjorden time series station (IsA) in Isfjorden, West Spitsbergen, Norway, was sampled frequently from December 2011 to December 2012. The community composition of microbial eukaryotes (size, 0.45 to 10 μm) from a depth of 25 m was determined using 454 sequencing of the 18S V4 region amplified from both DNA and RNA. The compositional changes throughout the year were assessed in relation to in situ fjord environmental conditions. Size fractionation analyses of chlorophyll a showed that the photosynthetic biomass was dominated by small cells (<10 μm) most of the year but that larger cells dominated during the spring and summer. The winter and early-spring communities were more diverse than the spring and summer/autumn communities. Dinophyceae were predominant throughout the year. The Arctic Micromonas ecotype was abundant mostly in the early-bloom and fall periods, whereas heterotrophs, such as marine stramenopiles (MASTs), Picozoa, and the parasitoid marine alveolates (MALVs), displayed higher relative abundance in the winter than in other seasons. Our results emphasize the extreme seasonality of Arctic microbial eukaryotic communities driven by the light regime and nutrient availability but point to the necessity of a thorough knowledge of hydrography for full understanding of their succession and variability.

Evolution of the Arctic Calanus complex: an Arctic marine avocado?

Before man hunted the large baleen whales to near extinction by the end of the nineteenth century, Arctic ecosystems were strongly influenced by these large predators. Their main prey were zooplankton, among which the calanoid copepod species of the genus Calanus, long considered key elements of polar marine ecosystems, are particularly abundant. These herbivorous zooplankters display a range of adaptations to the highly seasonal environments of the polar oceans, most notably extensive energy reserves and seasonal migrations to deep waters where the non-feeding season is spent in diapause. Classical work in marine ecology has suggested that slow growth, long lifespan and large body size in zooplankton are specific adaptations to life in cold waters with short and unpredictable feeding seasons. Here, we challenge this understanding and, by using an analogy from the evolutionary and contemporary history of the avocado, argue that predation pressure by the now nearly extinct baleen whales was an important driving force in the evolution of life history diversity in the Arctic Calanus complex.

Key Arctic phototrophs are widespread in the polar night

Small photosynthetic pico- and nanoeukaryotes contribute substantially to the biomass and primary production in the Arctic, often producing large blooms during spring and summer. During the civil polar night, which in the Svalbard Archipelago lasts from the middle of November to the end of February, no light is present, thus providing photosynthetic organisms with the challenge of how to survive several months of darkness. The small green alga Micromonas pusilla and the haptophyte Phaeocystis pouchetii are two key phototrophs in the Arctic, commonly blooming during the arctic spring and summer. Their occurence in Arctic waters during the polar night period is, however, less well known. In the present study, we used a molecular approach to show that M. pusilla and P. pouchetii are widely distributed in Svalbard waters also at the height of the polar night. Both species were detected in pelagic samples from both fjords and the open ocean, ice-covered and ice-free locations, shallow and deep water and from Atlantic, Arctic and coastal water masses. PCR screening was performed on both DNA and RNA samples, the latter allowing the detection of viable cells of both species in the mesopelagos. As far as we are aware this is the first systematic study on the persistence of these important photosynthetic organisms through the polar night.

Inter‐ and intraspecific variation in body‐ and genome size in calanoid copepods from temperate and arctic waters

Summary The tendency of ectotherms to get larger in the cold (Bergmann clines) has potentially great implications for individual performance and food web dynamics. The mechanistic drivers of this trend are not well understood, however. One fundamental question is to which extent variation in body size is attributed to variation in cell size, which again is related to genome size. In this study, we analyzed body and genome size in four species of marine calanoid copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Paraeuchaeta norvegica, with populations from both south Norwegian fjords and the High Arctic. The Calanus species showed typical interspecific Bergmann clines, and we assessed whether they also displayed similar intraspecific variations—and if correlation between genome size and body size differed between species. There were considerable inter‐ as well as intraspecific variations in body size and genome size, with the northernmost populations having the largest values of both variables within each species. Positive intraspecific relationships suggest a functional link between body and genome size, although its adaptiveness has not been settled. Impact of additional drivers like phylogeny or specific adaptations, however, was suggested by striking divergences in body size – genome size ratios among species. Thus, C. glacialis and C. hyperboreus, had fairly similar genome size despite very different body size, while P. norvegica, of similar body size as C. hyperboreus, had the largest genome sizes ever recorded from copepods. The inter‐ and intraspecific latitudinal body size clines suggest that climate change may have major impact on body size composition of keystone species in marine planktonic food webs.

Microbial eukaryotes in an arctic under-ice spring bloom north of Svalbard

Microbial eukaryotes can play prominent roles in the Arctic marine ecosystem, but their diversity and variability is not well known in the ice-covered ecosystems. We determined the community composition of microbial eukaryotes in an Arctic under-ice spring bloom north of Svalbard using metabarcoding of DNA and RNA from the hypervariable V4 region of 18S nrDNA. At the two stations studied, the photosynthetic biomass was dominated by protists >3 μm and was concentrated in the upper 70–80 m, above the thermocline and halocline. Hierarchical cluster analyses as well as ordination analyses showed a distinct clustering of the microbial eukaryote communities according to a combination of water mass and local environmental characteristics. While samples collected in the surface mixed layer differed distinctly between the two sites, the deeper communities collected in Atlantic Water were fairly similar despite being geographically distant. The differentiation of the microbial eukaryote communities of the upper mixed water was probably driven by local development and advection, while the lack of such differentiation in the communities of Atlantic Water reflects the homogenizing effect of water currents on microbial communities.

Proton-pumping rhodopsins are abundantly expressed by microbial eukaryotes in a high-Arctic fjord: Rhodopsin expression in Arctic marine eukaryotes

Proton-pumping rhodopsins provide an alternative pathway to photosynthesis by which solar energy can enter the marine food web. Rhodopsin genes are widely found in marine bacteria, also in the Arctic, and were recently reported from several eukaryotic lineages. So far, little is known about rhodopsin expression in Arctic eukaryotes. In this study, we used metatranscriptomics and 18S rDNA tag sequencing to examine the mid-summer function and composition of marine protists (size 0.45-10 µm) in the high-Arctic Billefjorden (Spitsbergen), especially focussing on the expression of microbial proton-pumping rhodopsins. Rhodopsin transcripts were highly abundant, at a level similar to that of genes involved in photosynthesis. Phylogenetic analyses placed the environmental rhodopsins within disparate eukaryotic lineages, including dinoflagellates, stramenopiles, haptophytes and cryptophytes. Sequence comparison indicated the presence of several functional types, including xanthorhodopsins and a eukaryotic clade of proteorhodopsin. Transcripts belonging to the proteorhodopsin clade were also abundant in published metatranscriptomes from other oceanic regions, suggesting a global distribution. The diversity and abundance of rhodopsins show that these light-driven proton pumps play an important role in Arctic microbial eukaryotes. Understanding this role is imperative to predicting the future of the Arctic marine ecosystem faced by a changing light climate due to diminishing sea-ice.

Seasonality of bivalve larvae within a high Arctic fjord

The temporal and spatial distribution of larval plankton of high latitudes is poorly understood. The objective of this work is to identify the occurrence and abundance of pelagic bivalve larvae within a high Arctic fjord (Adventfjorden, Svalbard) and to reveal their seasonal dynamics in relation to environmental variables—temperature, salinity and chlorophyll a—between December 2011 and January 2013. We applied a combination of DNA barcoding of mitochondrial 16S ribosomal RNA and morphological analysis to identify the bivalve larvae found within the plankton and demonstrate a strong seasonality in the occurrence of bivalve larvae, largely coinciding with periods of primary productivity. Seasonal occurrences of bivalve larval species differ from those known for other populations across species’ biogeographic distribution ranges. Serripes groenlandicus, which is of circum-Arctic distribution, demonstrated a later occurrence than Mya truncata or Hiatella arctica, which are of predominantly boreal or cosmopolitan distribution, respectively. S. groenlandicus larvae demonstrate the most pronounced response to seasonality, with the shortest presence in the water column. Establishing latitudinal differences in the occurrence of bivalve larvae enhances our understanding of how reproductive traits of marine invertebrates may respond to climate-driven seasonal shifts in the occurrence of primary productivity.

Feeding by Calanus glacialis in a high arctic fjord: potential seasonal importance of alternative prey

Original Article Feeding by Calanus glacialis in a high arctic fjord: potential seasonal importance of alternative prey Alison C. Cleary*, Janne E. Søreide, Daniela Freese, Barbara Niehoff, and Tove M. Gabrielsen University Centre in Svalbard, Longyearbyen N-9171, Norway Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research, 27570 Bremerhaven, Germany *Corresponding author: tel: þ47 96 88 53 94; fax: þ47 77 75 05 01; e-mail: alison_cleary@my.uri.edu Present address: Norwegian Polar Institute, Fram Centre, Tromsø NO-9296, Norway.