Arctic Diatom Research Articles

Major Diatom Microfossils from the Arctic Region: A Review

Abstract Diatoms-the unicellular, eukaryotic, photosynthetic phytoplankton belonging to Class Bacillariophyceae are an excellent tool to understand past ocean surface conditions, especially in high latitude regions. Along with sea surface temperature and sea ice, which they are primarily sensitive to, diatom assemblages could potentially be used to record runoff from melting glaciers and atmospheric teleconnections such as North Atlantic Oscillations (NAO) in Arctic fjords. However, understanding the ecology and geographical distribution of diatoms is extremely important before they can be used for palaeoceanographic reconstructions. In this study, four significant Arctic diatoms namely Bacterosira bathyomphala, Fragilariopsis cylindrus, Fragilaripsis oceanica and Thalassiosira antarctica var. borealis resting spores (R.S.) have been reviewed to understand their current status in their use for palaeoceanographic reconstructions. These four diatoms have been selected as they represent a typical Arctic environment. Numerous studies in the past thirty years had classified these diatoms into different assemblage groups, indicating specific ocean surface conditions. Here, we review the previous studies and to present the up-to-date classification of these species. We also compared the change in abundance of these four species through mid to late Holocene and modern period to their respective sea surface temperature and sea ice reconstructions, to show the robustness of their usage in palaeoceanographic reconstructions. The findings of this study will help to understand the up-to-date ecological characteristics of Arctic diatoms used in palaeoceanographic as well as palaeoecological reconstructions and the gap areas in diatom research such as a requirement for pan Arctic Sea ice diatom dataset.

Harmonizing marine Arctic diatom taxonomy for improving paleoenvironmental reconstructions

Harmonizing marine Arctic diatom taxonomy for improving paleoenvironmental reconstructions

Marine Arctic Diatoms working group

Marine Arctic Diatoms working group

Responses to herbicides of Arctic and temperate microalgae grown under different light intensities

In aquatic ecosystems, microalgae are exposed to light fluctuations at different frequencies due to daily and seasonal changes. Although concentrations of herbicides are lower in Arctic than in temperate regions, atrazine and simazine, are increasingly found in northern aquatic systems because of long-distance aerial dispersal of widespread applications in the south and antifouling biocides used on ships. The toxic effects of atrazine on temperate microalgae are well documented, but very little is known about their effects on Arctic marine microalgae in relation to their temperate counterparts after light adaptation to variable light intensities. We therefore investigated the impacts of atrazine and simazine on photosynthetic activity, PSII energy fluxes, pigment content, photoprotective ability (NPQ), and reactive oxygen species (ROS) content under three light intensities. The goal was to better understand differences in physiological responses to light fluctuations between Arctic and temperate microalgae and to determine how these different characteristics affect their responses to herbicides. The Arctic diatom Chaetoceros showed stronger light adaptation capacity than the Arctic green algae Micromonas. Atrazine and simazine inhibited the growth and photosynthetic electron transport, affected the pigment content, and disturbed the energy balance between light absorption and utilization. As a result, during high light adaptation and in the presence of herbicides, photoprotective pigments were synthesized and NPQ was highly activated. Nevertheless, these protective responses were insufficient to prevent oxidative damage caused by herbicides in both species from both regions, but at different extent depending on the species. Our study demonstrates that light is important in regulating herbicide toxicity in both Arctic and temperate microalgal strains. Moreover, eco-physiological differences in light responses are likely to support changes in the algal community, especially as the Arctic ocean becomes more polluted and bright with continued human impacts.

Unlocking environmental archives in the Arctic—insights from modern diatom-environment relationships in lakes and ponds across Greenland

Given the current rate of Arctic warming, the associated ecological changes need to be put into a longer-term context of natural variability. Palaeolimnology offers tools to explore archives stored in the sediments of Arctic lakes and ponds. The interpretation of these archives requires a sound knowledge of the ecology and distribution of the sedimentary proxy organisms used. Here we explored the relationship between diatoms, a widely used proxy group of siliceous algae, and the environmental drivers defining their assemblages and diversity in 115 lakes and ponds in Greenland, a markedly understudied arctic region covering extensive climate and environmental gradients. The main environmental drivers of diatom communities were related to climate and lake ontogeny, including both measured and unmeasured (spatially structured) environmental variables. The lakes and ponds in the northern study regions showed a distinctive dominance of small benthic fragilarioid species, while diatom communities in the South(west) of Greenland were more varied, including many epiphytes, owing to the longer growing season and higher habitat diversity of these lakes and ponds. The newly established lakes in the Ilulissat region host markedly different communities compared to all other sites. Species diversity followed an overall clear latitudinal decline towards the North. Despite the large distances between our study regions, diatom dispersal appeared not to be limited. Based on our results, diatoms are an excellent proxy for climate-mediated lake ecosystem change in the Arctic and thus a valuable tool for climate reconstructions in the region. Particular consideration should be given to often unmeasured climate-related drivers, such as in-lake habitat availability, due to their apparent importance in defining Arctic diatom communities.

Seasonal and habitat-based variations in vertical export of biogenic sea-ice proxies in Hudson Bay

Despite their wide use in past sea-ice reconstructions, the seasonal, habitat and species-based sources of sedimentary sea-ice proxies are poorly understood. Here, we conduct direct observations of the community composition of diatoms, dinoflagellate cysts and highly branched isoprenoid lipids within the sea ice, water column, sediment traps and sediment surface in the Belcher Islands Archipelago, Hudson Bay throughout spring 2019. We find that Arctic diatom and dinoflagellate cysts species commonly used as sea-ice proxies appear to be only indirectly linked to sea-ice conditions, and that the sediment assemblages of these groups overrepresent summertime pelagic blooms. Species contributing to the diverse sea-ice diatom communities are rare in the sediment. Dinoflagellate cysts form a typical Arctic assemblage in the sediment, although they are virtually absent in the sea ice and water column in spring. We also find that certain highly branched isoprenoid lipids that were previously considered indicators of open water, can be produced in sea-ice. We conclude that contextual knowledge and a multiproxy approach are necessary in reconstruction, encouraging further studies on the sources and controls of sea-ice proxy production in different geographic areas.

Toward the development of a new toxicity test with the Arctic alga Nitzschia frigida

There are limited data available related to the sensitivity of Arctic species to environmental contaminants, and this knowledge gap creates uncertainty in environmental risk assessments (ERAs). To help address this concern, we optimized culturing conditions to allow for toxicity tests with an Arctic diatom, Nitzschia frigida. We found optimal conditions for growth were Harrison's medium with natural seawater at 2 °C under a continuous photoperiod of 90 μmol photons m2 s−1. We then compared the response of N. frigida with the temperate standard diatom species Skeletonema costatum. We performed concurrent and repeated exposures of the two species to three compounds (zinc, copper, and 1-methylnaphthalene). EC50 values calculated from N. frigida exposures were consistently lower than those from S. costatum tests for metals, but not 1-methylnaphthalene. Overall, we have taken the inaugural steps toward the development of a new toxicity test method using an Arctic species to inform ERAs in northern regions.

A story of resilience: Arctic diatom Chaetoceros gelidus exhibited high physiological plasticity to changing CO2 and light levels.

Arctic phytoplankton are experiencing multifaceted stresses due to climate warming, ocean acidification, retreating sea ice, and associated changes in light availability, and that may have large ecological consequences. Multiple stressor studies on Arctic phytoplankton, particularly on the bloom-forming species, may help understand their fitness in response to future climate change, however, such studies are scarce. In the present study, a laboratory experiment was conducted on the bloom-forming Arctic diatom Chaetoceros gelidus (earlier C. socialis) under variable CO2 (240 and 900 µatm) and light (50 and 100 µmol photons m-2 s-1) levels. The growth response was documented using the pre-acclimatized culture at 2°C in a closed batch system over 12 days until the dissolved inorganic nitrogen was depleted. Particulate organic carbon and nitrogen (POC and PON), pigments, cell density, and the maximum quantum yield of photosystem II (Fv/Fm) were measured on day 4 (D4), 6 (D6), 10 (D10), and 12 (D12). The overall growth response suggested that C. gelidus maintained a steady-state carboxylation rate with subsequent conversion to macromolecules as reflected in the per-cell POC contents under variable CO2 and light levels. A substantial amount of POC buildup at the low CO2 level (comparable to the high CO2 treatment) indicated the possibility of existing carbon dioxide concentration mechanisms (CCMs) that needs further investigation. Pigment signatures revealed a high level of adaptability to variable irradiance in this species without any major CO2 effect. PON contents per cell increased initially but decreased irrespective of CO2 levels when nitrogen was limited (D6 onward) possibly to recycle intracellular nitrogen resources resulting in enhanced C: N ratios. On D12 the decreased dissolved organic nitrogen levels could be attributed to consumption under nitrogen starvation. Such physiological plasticity could make C. gelidus "ecologically resilient" in the future Arctic.

Interactive effects of temperature and nitrogen source on the elemental stoichiometry of a polar diatom

AbstractA recent study hypothesized that the near‐zero temperatures that generally prevail in Arctic waters negate the influence that different nitrogen (N) sources can otherwise have on the growth and elemental stoichiometry of marine micro‐algae. Here we test this hypothesis experimentally by evaluating how temperature (0–9°C) affects the growth and elemental stoichiometry of an ecologically relevant Arctic diatom Chaetoceros gelidus growing on different N sources (ammonium, nitrate, urea) at saturating irradiance. Following an initial acclimation period in which steady growth rates were achieved under each experimental treatment, changes in cellular concentrations of chlorophyll a and particulate carbon (C), nitrogen (N), phosphorus (P), and biogenic silica (Si) were monitored. While N source effects on growth rate became manifest as temperature rose above 0°C, the estimated optimal growth temperature was similar in all cases (Topt = 8.3°C). A positive effect of temperature on the N : P ratio occurred only at 6°C. Above this temperature, the N : P ratio decreased to values close to those observed at 0°C and 3°C. By contrast, the C : N ratio remained nearly invariant between 0°C and 6°C but increased substantially at 9°C. Overall, the results suggest that the presently widespread and successful diatom C. gelidus possesses the ability to remain competitive despite ongoing environmental changes and that its responses to warming and the availability of different N sources may impact elemental fluxes in the future.

Photoacclimation of the polar diatom Chaetoceros neogracilis at low temperature.

Polar microalgae face two major challenges: 1- growing at temperatures (-1.7 to 5°C) that limit enzyme kinetics; and 2- surviving and exploiting a wide range of irradiance. The objective of this study is to understand the adaptation of an Arctic diatom to its environment by studying its ability to acclimate to changes in light and temperature. We acclimated the polar diatom Chaetoceros neogracilis to various light levels at two different temperatures and studied its growth and photosynthetic properties using semi-continuous cultures. Rubisco content was high, to compensate for low catalytic rates, but did not change detectably with growth temperature. Contrary to what is observed in temperate species, in C. neogracilis, carbon fixation rate (20 min 14C incorporation) equaled net growth rate (μ) suggesting very low or very rapid (<20 min) re-oxidation of the newly fixed carbon. The comparison of saturation irradiances for electron transport, oxygen net production and carbon fixation revealed alternative electron pathways that could provide energy and reducing power to the cell without consuming organic carbon which is a very limiting product at low temperatures. High protein contents, low re-oxidation of newly fixed carbon and the use of electron pathways alternative to carbon fixation may be important characteristics allowing efficient growth under those extreme environmental conditions.

Diversity and biogeography of planktonic diatoms in Svalbard fjords: The role of dispersal and Arctic endemism in phytoplankton community structuring

Understanding the processes that shape the community structure of Arctic phytoplankton is crucial for predicting responses of Arctic ecosystems to the ongoing ocean warming. In particular, little is known about the importance of phytoplankton dispersal by the North Atlantic Current and the prevalence and maintenance of Arctic endemism. We investigated the diversity and biogeography of diatoms from five Svalbard fjords and the Hausgarten observatory (Fram Strait) by combining diatom cultivation and 18S rRNA gene metabarcoding. In total, 50 diatom strains were isolated from the area during the HE492 cruise in August 2017. The strains were identified taxonomically using molecular and morphological approaches, and their biogeographic distribution was mapped using the local metabarcoding dataset and a global compilation of published metabarcoding datasets. Biogeographic analysis was also conducted for the locally most abundant diatom metabarcoding amplicon sequence variants. The biogeographic analyses demonstrated that Arctic diatoms exhibit three general biogeographic distribution types: Arctic, Arctic-temperate, and cosmopolitan. At Hausgarten and in outer Isfjorden on the west coast of Svalbard, the communities were dominated by genotypes with Arctic-temperate and cosmopolitan distribution. Diatom communities in nearby Van Mijenfjorden, inner Isfjorden and Kongsfjorden were dominated by genotypes with Arctic-temperate distribution, and cosmopolitan species were less abundant. The genotypes endemic to the Arctic had lower abundance on the west coast of Svalbard. The two northernmost fjords (Woodfjorden and Wijdefjorden) had a higher abundance of genotypes endemic to the Arctic. Our results demonstrate that the diatom communities in the Svalbard area consist of genotypes endemic to the Arctic, and genotypes with broader biogeographic distribution, all of which are further structured by local environmental gradients. Finer biogeographic patterns observed within Arctic-temperate and cosmopolitan genotypes suggest that certain genotypes can be used as indicators of increasing influence of Atlantic waters on the phytoplankton community structure in the Svalbard area.

Pelagic and ice‐associated microalgae under elevated light and pCO2: Contrasting physiological strategies in two Arctic diatoms

AbstractSea ice retreat, changing stratification, and ocean acidification are fundamentally changing the light availability and physico‐chemical conditions for primary producers in the Arctic Ocean. However, detailed studies on ecophysiological strategies and performance of key species in the pelagic and ice‐associated habitat remain scarce. Therefore, we investigated the acclimated responses of the diatoms Thalassiosira hyalina and Melosira arctica toward elevated irradiance and CO2 partial pressures (pCO2). Next to growth, elemental composition, and biomass production, we assessed detailed photophysiological responses through fluorometry and gas‐flux measurements, including respiration and carbon acquisition. In the pelagic T. hyalina, growth rates remained high in all treatments and biomass production increased strongly with light. Even under low irradiances cells maintained a high‐light acclimated state, allowing them to opportunistically utilize high irradiances by means of a highly plastic photosynthetic machinery and carbon uptake. The ice‐associated M. arctica proved to be less plastic and more specialized on low‐light. Its acclimation to high irradiances was characterized by minimizing photon harvest and photosynthetic efficiency, which led to lowered growth. Comparably low growth rates and strong silification advocate a strategy of persistence rather than of fast proliferation, which is also in line with the observed formation of resting stages under low‐light conditions. In both species, responses to elevated pCO2 were comparably minor. Although both diatom species persisted under the applied conditions, their competitive abilities and strategies differ strongly. With the anticipated extension of Arctic pelagic habitats, flexible high‐light specialists like T. hyalina seem to face a brighter future.

Shifts in growth light optima among diatom species support their succession during the spring bloom in the Arctic

Abstract Diatoms of the Arctic Ocean annually experience extreme changes of light environment linked to photoperiodic cycles and seasonal variations of the snow and sea‐ice cover extent and thickness which attenuate light penetration in the water column. Arctic diatom communities exploit this complex seasonal dynamic through a well‐documented species succession during spring, beginning in sea‐ice and culminating in massive phytoplankton blooms underneath sea‐ice and in the marginal ice zone. The pattern of diatom taxa sequentially dominating this succession is relatively well conserved interannually, and taxonomic shifts seem to align with habitat transitions. To understand whether differential photoadaptation strategies among diatom taxa explain these recurring succession sequences, we coupled laboratory experiments with field work in Baffin Bay at 67.5°N. Based on field data, we selected five diatom species typical of different ecological niches and measured their growth rates under light intensity ranges representative of their natural habitats. To characterize their photoacclimative responses, we sampled pigments and total particulate carbon, and conducted 14C‐uptake photosynthesis response curves and variable fluorescence measurements. We documented a gradient in species respective light intensity for maximal growth suggesting divergent light response plasticity, which for the most part align with species sequential dominance. Other photophysiological parameters supported this ecophysiological framing, although contrasts were always clear only between succession endmembers, Nitzschia frigida and Chaetoceros neogracilis. To validate that these photoacclimative responses are representative of in situ dynamics, we compared them to the chlorophyll a‐specific light‐limited slope (α*) and saturated rate of photosynthesis (), monitored in Baffin Bay on sea‐ice and planktonic communities. This complementary approach confirmed that unusual responses in α* and as a function of light history intensity are similar between sentinel sympagic species N. frigida and natural ice‐core communities. While no light‐history‐dependent trends were observed in planktonic communities, their α* and values were in the range of measurements from our monospecific cultures. Synthesis. Our results suggest that Arctic diatoms species photoadaptation strategy is tuned to the light environment of the habitats in which they dominate and indeed drives the seasonal taxonomic succession.

Revealing environmentally driven population dynamics of an Arctic diatom using a novel microsatellite PoolSeq barcoding approach.

Ecological stability under environmental change is determined by both interspecific and intraspecific processes. Particularly for planktonic microorganisms, it is challenging to follow intraspecific dynamics over space and time. We propose a new method, microsatellite PoolSeq barcoding (MPB), for tracing allele frequency changes in protist populations. We successfully applied this method to experimental community incubations and field samples of the diatom Thalassiosira hyalina from the Arctic, a rapidly changing ecosystem. Validation of the method found compelling accuracy in comparison with established genotyping approaches within different diversity contexts. In experimental and environmental samples, we show that MPB can detect meaningful patterns of population dynamics, resolving allelic stability and shifts within a key diatom species in response to experimental treatments as well as different bloom phases and years. Through our novel MPB approach, we produced a large dataset of populations at different time-points and locations with comparably little effort. Results like this can add insights into the roles of selection and plasticity in natural protist populations under stable experimental but also variable field conditions. Especially for organisms where genotype sampling remains challenging, MPB holds great potential to efficiently resolve eco-evolutionary dynamics and to assess the mechanisms and limits of resilience to environmental stressors.

The ice-nucleating activity of Arctic sea surface microlayer samples and marine algal cultures

Abstract. In recent years, sea spray as well as the biological material it contains has received increased attention as a source of ice-nucleating particles (INPs). Such INPs may play a role in remote marine regions, where other sources of INPs are scarce or absent. In the Arctic, these INPs can influence water–ice partitioning in low-level clouds and thereby the cloud lifetime, with consequences for the surface energy budget, sea ice formation and melt, and climate. Marine aerosol is of a diverse nature, so identifying sources of INPs is challenging. One fraction of marine bioaerosol (phytoplankton and their exudates) has been a particular focus of marine INP research. In our study we attempt to address three main questions. Firstly, we compare the ice-nucleating ability of two common phytoplankton species with Arctic seawater microlayer samples using the same instrumentation to see if these phytoplankton species produce ice-nucleating material with sufficient activity to account for the ice nucleation observed in Arctic microlayer samples. We present the first measurements of the ice-nucleating ability of two predominant phytoplankton species: Melosira arctica, a common Arctic diatom species, and Skeletonema marinoi, a ubiquitous diatom species across oceans worldwide. To determine the potential effect of nutrient conditions and characteristics of the algal culture, such as the amount of organic carbon associated with algal cells, on the ice nucleation activity, Skeletonema marinoi was grown under different nutrient regimes. From comparison of the ice nucleation data of the algal cultures to those obtained from a range of sea surface microlayer (SML) samples obtained during three different field expeditions to the Arctic (ACCACIA, NETCARE, and ASCOS), we found that they were not as ice active as the investigated microlayer samples, although these diatoms do produce ice-nucleating material. Secondly, to improve our understanding of local Arctic marine sources as atmospheric INPs we applied two aerosolization techniques to analyse the ice-nucleating ability of aerosolized microlayer and algal samples. The aerosols were generated either by direct nebulization of the undiluted bulk solutions or by the addition of the samples to a sea spray simulation chamber filled with artificial seawater. The latter method generates aerosol particles using a plunging jet to mimic the process of oceanic wave breaking. We observed that the aerosols produced using this approach can be ice active, indicating that the ice-nucleating material in seawater can indeed transfer to the aerosol phase. Thirdly, we attempted to measure ice nucleation activity across the entire temperature range relevant for mixed-phase clouds using a suite of ice nucleation measurement techniques – an expansion cloud chamber, a continuous-flow diffusion chamber, and a cold stage. In order to compare the measurements made using the different instruments, we have normalized the data in relation to the mass of salt present in the nascent sea spray aerosol. At temperatures above 248 K some of the SML samples were very effective at nucleating ice, but there was substantial variability between the different samples. In contrast, there was much less variability between samples below 248 K. We discuss our results in the context of aerosol–cloud interactions in the Arctic with a focus on furthering our understanding of which INP types may be important in the Arctic atmosphere.

Contrasting nonphotochemical quenching patterns under high light and darkness aligns with light niche occupancy in Arctic diatoms

AbstractOver the seasons, Arctic diatom species occupy shifting habitats defined by contrasting light climates, constrained by snow and ice cover dynamics interacting with extreme photoperiod and solar angle variations. How Arctic diatom photoadaptation strategies differ across their heterogeneous light niches remains a poorly documented but crucial missing link to anticipate Arctic Ocean responses to shrinking sea‐ice and increasing light. To address this question, we selected five Arctic diatom species with diverse life traits, representative of distinct light niches across the seasonal light environment continuum: from snow‐covered dimly lit bottom ice to summer stratified waters. We studied their photoacclimation plasticity to two growth light levels and the subsequent responses of their nonphotochemical quenching (NPQ) and xanthophyll cycle to both dark incubations and light shifts. We deciphered NPQ and xanthophyll cycle tuning in darkness and their light‐dependent induction kinetics, which aligned with species' light niche occupancy. In ice‐related species, NPQ was sustained in darkness and its induction was more reactive to moderate light shifts. Open‐water species triggered strong NPQ induction in darkness and reached higher maximal NPQ under high light. Marginal ice zone species showed strong adaptation to light fluctuations with a dark response fine‐tuned depending upon light history. We argue these traits are anchored in diverging photoadaption strategies fostering Arctic diatom success in their respective light niches.

Use of organic exudates from two polar diatoms by bacterial isolates from the Arctic Ocean.

Global warming affects primary producers in the Arctic, with potential consequences for the bacterial community composition through the consumption of microalgae-derived dissolved organic matter (DOM). To determine the degree of specificity in the use of an exudate by bacterial taxa, we used simple microalgae-bacteria model systems. We isolated 92 bacterial strains from the sea ice bottom and the water column in spring-summer in the Baffin Bay (Arctic Ocean). The isolates were grouped into 42 species belonging to Proteobacteria, Bacteroidetes, Actinobacteria and Firmicutes. Forty strains were tested for their capacity to grow on the exudate from two Arctic diatoms. Most of the strains tested (78%) were able to grow on the exudate from the pelagic diatom Chaetoceros neogracilis, and 33% were able to use the exudate from the sea ice diatom Fragilariopsis cylindrus. 17.5% of the strains were not able to grow with any exudate, while 27.5% of the strains were able to use both types of exudates. All strains belonging to Flavobacteriia (n = 10) were able to use the DOM provided by C.neogracilis, and this exudate sustained a growth capacity of up to 100 times higher than diluted Marine Broth medium, of two Pseudomonas sp. strains and one Sulfitobacter strain. The variable bioavailability of exudates to bacterial strains highlights the potential role of microalgae in shaping the bacterial community composition. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Biodiversity patterns of Arctic diatom assemblages in lakes and streams: Current reference conditions and historical context for biomonitoring

Abstract Comprehensive assessments of contemporary diatom distributions across the Arctic remain scarce. Furthermore, studies tracking species compositional differences across space and time, as well as diatom responses to climate warming, are mainly limited to paleolimnological studies due to a lack of routine monitoring in lakes and streams across vast areas of the Arctic. The study aims to provide a spatial assessment of contemporary species distributions across the circum‐Arctic, establish contemporary biodiversity patterns of diatom assemblages to use as reference conditions for future biomonitoring assessments, and determine pre‐industrial baseline conditions to provide historical context for modern diatom distributions. Diatom assemblages were assessed using information from ongoing regulatory monitoring programmes, individual research projects, and from surface sediment layers obtained from lake cores. Pre‐industrial baseline conditions as well as the nature, direction and magnitude of changes in diatom assemblages over the past c.200 years were determined by comparing surface sediment samples (i.e. containing modern assemblages) with a sediment interval deposited prior to the onset of significant anthropogenic activities (i.e. containing pre‐1850 assemblages), together with an examination of diatoms preserved in contiguous samples from dated sediment cores. We identified several biotypes with distinct diatom assemblages using contemporary diatom data from both lakes and streams, including a biotype typical for High Arctic regions. Differences in diatom assemblage composition across circum‐Arctic regions were gradual rather than abrupt. Species richness was lowest in High Arctic regions compared to Low Arctic and sub‐Arctic regions, and higher in lakes than in streams. Dominant diatom taxa were not endemic to the Arctic. Species richness in both lakes and streams reached maximum values between 60°N and 75°N but was highly variable, probably reflecting differences in local and regional environmental factors and possibly sampling effort. We found clear taxon‐specific differences between contemporary and pre‐industrial samples that were often specific to both ecozone and lake depth. Regional patterns of species turnover (β‐diversity) in the past c.200 years revealed that regions of the Canadian High Arctic and the Hudson Bay Lowlands to the south showed most compositional change, whereas the easternmost regions of the Canadian Arctic changed least. As shown in previous Arctic diatom studies, global warming has already affected these remote high latitude ecosystems. Our results provide reference conditions for future environmental monitoring programmes in the Arctic. Furthermore, diatom taxa identification and harmonisation require improvement, starting with circum‐Arctic intercalibrations. Despite the challenges posed by the remoteness of the Arctic, our study shows the need for routine monitoring programmes that have a wide geographical coverage for both streams and lakes.

Phylogenetic diversity and environment form assembly rules for Arctic diatom genera—A study on recent and ancient sedimentary DNA

AbstractAimThis study investigates taxonomic and phylogenetic diversity in diatom genera to evaluate assembly rules for eukaryotic microbes across the Siberian tree line. We first analysed how phylogenetic distance relates to taxonomic richness and turnover. Second, we used relatedness indices to evaluate if environmental filtering or competition influences the assemblies in space and through time. Third, we used distance‐based ordination to test which environmental variables shape diatom turnover.LocationYakutia and Taymyria, Russia: we sampled 78 surface sediments and a sediment core, extending to 7,000 years before present, to capture the forest–tundra transition in space and time respectively.TaxonArctic freshwater diatoms.MethodsWe applied metabarcoding to retrieve diatom diversity from surface and core sedimentary DNA. The taxonomic assignment binned sequence types (lineages) into genera and created taxonomic (abundance of lineages within different genera) and phylogenetic datasets (phylogenetic distances of lineages within different genera).ResultsContrary to our expectations, we find a unimodal relationship between phylogenetic distance and richness in diatom genera. We discern a positive relationship between phylogenetic distance and taxonomic turnover in spatially and temporally distributed diatom genera. Furthermore, we reveal positive relatedness indices in diatom genera across the spatial environmental gradient and predominantly in time slices at a single location, with very few exceptions assuming effects of competition. Distance‐based ordination of taxonomic and phylogenetic turnover indicates that lake environment variables, like HCO3− and water depth, largely explain diatom turnover.Main conclusionPhylogenetic and abiotic assembly rules are important in understanding the regional assembly of diatom genera across lakes in the Siberian tree line ecotone. Using a space—time approach we are able to exclude the influence of geography and elucidate that lake environmental variables primarily shape the assemblies. We conclude that some diatom genera have greater capabilities to adapt to environmental changes, whereas others will be putatively replaced or lost due to the displacement of the Arctic tundra biome under recent global warming.

Arctic (Svalbard islands) active and exported diatom stocks and cell health status

Abstract. Diatoms tend to dominate the Arctic spring phytoplankton bloom, a key event in the ecosystem including a rapid decline in surface-water pCO2. While a mass sedimentation event of diatoms at the bloom terminus is commonly observed, there are few reports on the status of diatoms' health during Arctic blooms and its possible role on sedimentary fluxes. Thus, we examine the idea that the major diatom-sinking event which occurs at the end of the regional bloom is driven by physiologically deteriorated cells. Here we quantify, using the Bottle-Net, Arctic diatom stocks below and above the photic zone and assess their cell health status. The communities were sampled around the Svalbard islands and encompassed pre- to post-bloom conditions. A mean of 24.2±6.7 % SE (standard error) of the total water column (max. 415 m) diatom standing stock was found below the photic zone, indicating significant diatom sedimentation. The fraction of living diatom cells in the photic zone averaged 59.4±6.3 % but showed the highest mean percentages (72.0 %) in stations supporting active blooms. In contrast, populations below the photic layer were dominated by dead cells (20.8±4.9 % living cells). The percentage of diatoms' standing stock found below the photic layer was negatively related to the percentage of living diatoms in the surface, indicating that healthy populations remained in the surface layer. Shipboard manipulation experiments demonstrated that (1) dead diatom cells sank faster than living cells, and (2) diatom cell mortality increased in darkness, showing an average half-life among diatom groups of 1.025±0.075 d. The results conform to a conceptual model where diatoms grow during the bloom until resources are depleted and supports a link between diatom cell health status (affected by multiple factors) and sedimentation fluxes in the Arctic. Healthy Arctic phytoplankton communities remained at the photic layer, whereas the physiologically compromised (e.g., dying) communities exported a large fraction of the biomass to the aphotic zone, fueling carbon sequestration to the mesopelagic and material to benthic ecosystems.