Molecular tools prove little auks from Svalbard are extremely selective for Calanus glacialis even when exposed to Atlantification.

Two Calanus species, C. glacialis and C. finmarchicus, due to different life strategies and environmental preferences act as an ecological indicators of Arctic Atlantification. Their high lipid content makes them important food source for higher trophic levels of Arctic ecosystems including the most abundant Northern Hemisphere's seabird, the little auk (Alle alle). Recent studies indicate a critical need for the use of molecular methods to reliably identify these two sympatric Calanus species. We performed genetic and morphology-based identification of 2600 Calanus individuals collected in little auks foraging grounds and diet in summer seasons 2019-2021 in regions of Svalbard with varying levels of Atlantification. Genetic identification proved that 40% of Calanus individuals were wrongly classified as C. finmarchicus according to morphology-based identification in both types of samples. The diet of little auks consisted almost entirely of C. glacialis even in more Atlantified regions. Due to the substantial bias in morphology-based identification, we expect that the scale of the northern expansion of boreal C. finmarchicus may have been largely overestimated and that higher costs for birds exposed to Atlantification could be mostly driven by a decrease in the size of C. glacialis rather than by shift from C. glacialis to C. finmarchicus.

Dark plumes of glacial meltwater affect vertical distribution of zooplankton in the Arctic.

In polar regions, the release of glacial meltwater resulting in turbid plumes is expected to transform coastal waters with numerous consequences on the marine ecosystem. This study aimed to determine the influence of turbidity regimes on the vertical distribution of copepods together with their potential food (chlorophyll a fluorescence) and non-visual predators (gelatinous zooplankton). Hydrography, turbidity, suspended particulate matter and chlorophyll a were studied in July and August 2019 in West Spitsbergen waters (European Arctic). Fine-scale vertical distribution patterns of zooplankton were assessed by an optical counter (LOPC) and underwater camera (UVP) and verified by plankton nets. In waters with the shallow impact of dark plumes, Calanus spp. and gelatinous zooplankton were concentrated in the upper water layers, whereas in areas with a thick turbid layer, they were distributed evenly in the water column. However, chlorophyll a peaks were found to be restricted to the surface in the turbid waters and there were subsurface maxima in the shallow turbidity regime. Regardless of the region, the turbidity regime was a significant factor shaping the vertical distribution of Calanus spp. We speculate that similar trends might be observed in other rapidly emerging turbid ecosystems and urge that future plankton research should also include relatively simple turbidity measurements.

Gone with the wind - Wind speed affects prey accessibility for a High Arctic zooplanktivorous seabird, the little auk Alle alle.

Foraging ecology of chick rearing seabirds is affected mainly by the food availability on feeding grounds, but it can be also modulated by environmental conditions during the foraging trip, in that wind force. Considering predicted strengthening of surface winds over the Arctic Ocean, this factor may have a growing impact on the foraging performance of Arctic seabirds. Here, we studied how wind speed could affect prey accessibility for the High Arctic zooplanktivorous seabird, the little auk Alle alle breeding in Svalbard in 2015-2019. First, we estimated availability of its preferred prey, a cold water copepod Calanus glacialis, based on wider-scale mesozooplankton biomass model and environmental conditions. Then we estimated prey accessibility by including wind speed, the factor affecting the flapping flight performance of little auks commuting from/to the colony. Finally, we compared reproductive performance of the little auks (chick diet, growth rate and survival and duration of foraging flights of adults) between the studied years differing in wind and food availability conditions. We found that wind speed could affect significantly food accessibility for a zooplanktivorous seabird. Despite high spatial and temporal variability in prey availability and accessibility in shelf waters of SW Spitsbergen, interannual differences in duration of foraging flights and chick growth rate, little auks were able to sustain high breeding success confirming their capacity to buffer suboptimal foraging conditions. Our multidisciplinary work, combining multi-year remote sensing of oceanographic conditions, zooplankton availability and accessibility modelling, little auks diet composition and chick growth and survival emphasizes the importance of including wind conditions in the studies of foraging ecology of seabirds.

Phenology of Oithona similis demonstrates that ecological flexibility may be a winning trait in the warming Arctic.

Rapidly warming Arctic is facing significant shifts in the zooplankton size-spectra manifested as increasing numbers of the small-sized copepod Oithona similis. Here we present a unique continuous data set covering 22 months, on its copepodite structure along with environmental drivers in the Atlantic-influenced high Arctic fjord Isfjorden (Spitsbergen). Abundance maxima of O. similis were observed in September when the highest seawater temperature was recorded. A high concentration of the indicator species of Atlantification Oithona atlantica was also observed at that time. The clear dominance of O. similis in the zooplankton community during the dark, theoretically unproductive season emphasizes its substantial role in sustaining a continuous carbon flow, when most of the large herbivorous copepods fall into sleeping state. The high sex ratio observed twice in both years during periods of high primary production suggests two main reproductive events per year. O. similis reproduced even in very low temperatures (< 0 °C) previously thought to limit their fecundity, which proves its unique thermal tolerance. Our study provides a new insight on ecology of this key copepod of marine ecosystems across the globe, and thus confirm the Climatic Variability Hypothesis assuming that natural selection favour species with such flexible adaptive traits as O. similis.

The interplay between plankton and particles in the Isfjorden waters influenced by marine- and land-terminating glaciers.

Climate-induced glacial retreat in the Arctic results in an increased supply of meltwater with suspended terrigenous material into the marine environment. Despite increasing research efforts, effects of glacial retreat on functioning of plankton are not well documented and understood. Thus, we studied a hydro-optical seawater regime along with particle/plankton concentrations and composition structure in a high Arctic fjord (Isfjorden, West Spitsbergen) during mid-summer in 2019. This comprehensive study of the upper 50 m water layer presented a sharp distinction between 'muddy' waters influenced by glacial and river runoff and 'clear' open fjordic waters in the form of a notable difference in chlorophyll a concentrations, extent of euphotic zone depth, turbidity, inorganic/organic particle concentrations, and water colour. In this study, we present that the effects of glacial retreat on Arctic pelagial depend not only on different types of glaciers (marine- and land-terminating), but presumably, also on fjord topography and exposure to oceanic water inflow. The contrasting glacial, hydrological, and topographical conditions had different effects on the share of zooplankton and marine snow. Despite adaptation of the planktonic communities in the Arctic to high sediment loads and resultant light limitations, our study shows that continuing retreat of tidewater glaciers will have negative effect on planktonic communities especially in enclosed shallow fjord branches. Moreover, seawater darkening due to high turbidity could negatively affect tactile predators, such as gelatinous zooplankton. Additional division of plankton into functional groups typically used in the biogeochemical models demonstrated that diatoms, flagellates and mesozooplankton are influenced by suspended matter, whereas microzooplankton are highly adaptive to increased sediment loads. Since we investigated the largest Svalbard fjord system and incorporated multiple components of the pelagic realm, the current study delivers important recommendations for including marine snow and gelatinous zooplankton in ecosystem models applied in polar regions.

Marine snow morphology illuminates the evolution of phytoplankton blooms and determines their subsequent vertical export.

The organic carbon produced in the ocean's surface by phytoplankton is either passed through the food web or exported to the ocean interior as marine snow. The rate and efficiency of such vertical export strongly depend on the size, structure and shape of individual particles, but apart from size, other morphological properties are still not quantitatively monitored. With the growing number of in situ imaging technologies, there is now a great possibility to analyze the morphology of individual marine snow. Thus, automated methods for their classification are urgently needed. Consequently, here we present a simple, objective categorization method of marine snow into a few ecologically meaningful functional morphotypes using field data from successive phases of the Arctic phytoplankton bloom. The proposed approach is a promising tool for future studies aiming to integrate the diversity, composition and morphology of marine snow into our understanding of the biological carbon pump.

In a comfort zone and beyond-Ecological plasticity of key marine mediators.

Copepods of the genus Calanus are the key components of zooplankton. Understanding their response to a changing climate is crucial to predict the functioning of future warmer high-latitude ecosystems. Although specific Calanus species are morphologically very similar, they have different life strategies and roles in ecosystems. In this study, C. finmarchicus and C. glacialis were thoroughly studied with regard to their plasticity in morphology and ecology both in their preferred original water mass (Atlantic vs. Arctic side of the Polar Front) and in suboptimal conditions (due to, e.g., temperature, turbidity, and competition in Hornsund fjord). Our observations show that "at the same place and time," both species can reach different sizes, take on different pigmentation, be in different states of population development, utilize different reproductive versus lipid accumulation strategies, and thrive on different foods. Size was proven to be a very mutable morphological trait, especially with regard to reduced length of C. glacialis. Both species exhibited pronounced red pigmentation when inhabiting their preferred water mass. In other domains, C. finmarchicus individuals tended to be paler than C. glacialis individuals. Gonad maturation and population development indicated mixed reproductive strategies, although a surprisingly similar population age structure of the two co-occurring species in the fjord was observed. Lipid accumulation was high and not species-specific, and its variability was due to diet differences of the populations. According to the stable isotope composition, both species had a more herbivorous diatom-based diet in their original water masses. While the diet of C. glacialis was rather consistent among the domains studied, C. finmarchicus exhibited much higher variability in its feeding history (based on lipid composition). Our results show that the plasticity of both Calanus species is indeed impressive and may be regulated differently, depending on whether they live in their "comfort zone" or beyond it.

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.

Variations in the structural and functional diversity of zooplankton over vertical and horizontal environmental gradients en route to the Arctic Ocean through the Fram Strait.

A multi-scale approach was used to evaluate which spatial gradient of environmental variability is the most important in structuring zooplankton diversity in the West Spitsbergen Current (WSC). The WSC is the main conveyor of warm and biologically rich Atlantic water to the Arctic Ocean through the Fram Strait. The data set included 85 stratified vertical zooplankton samples (obtained from depths up to 1000 metres) covering two latitudinal sections (76°30'N and 79°N) located across the multi-path WSC system. The results indicate that the most important environmental variables shaping the zooplankton structural and functional diversity and standing stock variability are those associated with depth, whereas variables acting in the horizontal dimension are of lesser importance. Multivariate analysis of the zooplankton assemblages, together with different univariate descriptors of zooplankton diversity, clearly illustrated the segregation of zooplankton taxa in the vertical plane. The epipelagic zone (upper 200 m) hosted plentiful, Oithona similis-dominated assemblages with a high proportion of filter-feeding zooplankton. Although total zooplankton abundance declined in the mesopelagic zone (200-1000 m), zooplankton assemblages in that zone were more diverse and more evenly distributed, with high contributions from both herbivorous and carnivorous taxa. The vertical distribution of integrated biomass (mg DW m-2) indicated that the total zooplankton biomass in the epipelagic and mesopelagic zones was comparable. Environmental gradients acting in the horizontal plane, such as the ones associated with different ice cover and timing of the spring bloom, were reflected in the latitudinal variability in protist community structure and probably caused differences in succession in the zooplankton community. High abundances of Calanus finmarchicus in the WSC core branch suggest the existence of mechanisms advantageous for higher productivity or/and responsible for physical concentration of zooplankton. Our results indicate that regional hydrography plays a primary role in shaping zooplankton variability in the WSC on the way to the Arctic Ocean, with additional effects caused by biological factors related to seasonality in pelagic ecosystem development, resulting in regional differences in food availability or biological production between the continental slope and the deep ocean regions.