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.

Flexibility of little auks foraging in various oceanographic features in a changing Arctic.

Using GPS-tracked individuals, we compared foraging ecology and reproductive output of a High-Arctic zooplanktivorous seabird, the little auk Alle alle, between three years differing in environmental conditions (sea surface temperature). Despite contrasting environmental conditions, average foraging fights distance and duration were generally similar in all studied years. Also, in all years foraging locations visited by the little auk parents during short trips (ST, for chick provisioning) were significantly closer to the colony compared to those visited during long trips (LTs, mainly for adults' self-maintenance). Nevertheless, we also found some differences in the little auk foraging behaviour: duration of LTs was the longest in the coldest year suggesting more time for resting for adults compared to warmer years. Besides, birds foraged closer to the colony and in significantly colder water in the coldest year. Interestingly, these differences did not affect chick diet: in all the years, the energy content of food loads was similar, with the Arctic copepod, Calanus glacialis copepodite stage V being the most preferred prey item (>73% of items by number and >67% by energy content). Also chick survival was similar in all the study years. However, when examining chicks growth rate we found that their peak body mass was lower in warmer years suggesting that overall conditions in the two warm years were less favourable. While our results, demonstrate a great foraging flexibility by little auks, they also point out their vulnerability to changing environmental conditions.

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.

Identification, Discrimination, and Discovery of Species of Marine Planktonic Ostracods Using DNA Barcodes.

The Ostracoda (Crustacea; Class Ostracoda) is a diverse, frequently abundant, and ecologically important component of the marine zooplankton assemblage. There are more than 200 described species of marine planktonic ostracods, many of which (especially conspecific species) can be identified only by microscopic examination and dissection of fragile morphological characters. Given the complexity of species identification and increasing lack of expert taxonomists, DNA barcodes (short DNA sequences for species discrimination and identification) are particularly useful and necessary. Results are reported from analysis of 210 specimens of 78 species of marine planktonic ostracods, including two novel species, and 51 species for which barcodes have not been previously published. Specimens were collected during 2006 to 2008 from the Atlantic, Indian, and Southern Oceans, Greenland Sea and Gulf of Alaska. Samples were collected from surface to 5,000 m using various collection devices. DNA sequence variation was analyzed for a 598 base-pair region of the mitochondrial cytochrome oxidase subunit I (COI) gene. Kimura-2-Parameter (K2P) genetic distances within described species (mean = 0.010 ± 0.017 SD) were significantly smaller than between species (0.260 + 0.080), excluding eight taxa hypothesized to comprise cryptic species due to morphological variation (especially different size forms) and/or collection from different geographic regions. These taxa showed similar K2P distance values within (0.014 + 0.026) and between (0.221 ± 0.068) species. All K2P distances > 0.1 resulted from comparisons between identified or cryptic species, with no overlap between intra- and interspecific genetic distances. A Neighbor Joining tree resolved nearly all described species analyzed, with multiple sequences forming monophyletic clusters with high bootstrap values (typically 99%). Based on taxonomically and geographically extensive sampling and analysis (albeit with small sample sizes), the COI barcode region was shown to be a valuable character for discrimination, recognition, identification, and discovery of species of marine planktonic ostracods.

A re-description of Discoconchoecia elegans (Sars, 1865) (Ostracoda: Halocyprididae) from high latitudes in the North Atlantic.

Discoconchoecia elegans (Sars, 1865) is one of the most frequently recorded species of halocyprid ostracods and specimens are collected in abundance from various latitudes throughout the world oceans. This species is often dominant or subdominant member of the mesopelagic assemblages. However, its body size varies substantially with latitude, posing the question as to whether D. elegans is either a single, highly variable species, or a complex of cryptic and sibling species. Evaluation of the hypothesis that D. elegans is a complex of species requires comparison between the type material and specimens collected from different latitudes. The inadequacy of the original description from the type locality, off the Lofoten Islands (NW Norway), combined with a lack of the type material is preventing critical rating. In this paper Discoconchoecia elegans is redescribed from specimens collected from an area close to Svalbard, using detailed drawings, morphometric measurements of all limbs, and SEM photographs, and it is compared with specimens collected from an area close to the species type locality. The individuals from those two localities show no significant differences, probably because the hydrographic conditions are similar between the two sites.

Supplementary diet components of little auk chicks in two contrasting regions on the West Spitsbergen coast.

The complete diet composition structure of the most numerous planktivorous sea bird, little auk (Alle alle), in the European Arctic, is still not fully recognized. Although regular constituents of little auk chick diets, the copepods, Calanus glacialis and C. finmarchicus have been previously relatively well described, more taxa were frequent ingredients of the bird's meals. Therefore, the role of the little auks supplementary diet components (SDCs) at two colonies in the Svalbard Archipelago, Hornsund and Magdalenefjorden, in 2007-2009, is a main subject of this comparative study. Because the SDCs often consisted of scarce but large zooplankters, this investigation was focused on biomass as a proxy of the SDCs' energy input. Although the total biomass of the food delivered to chicks in both colonies was similar, in Magdalenefjorden, the proportion of SDCs was twice that found in Hornsund. The main SDCs in Hornsund were Decapoda larvae (with predominating Pagurus pubescens) and Thysanoessa inermis, whereas the main SDCs in Magdalenefjorden were C. hyperboreus and Apherusa glacialis. Previous investigations, which indicated lipid richness of SDCs, together with our ecological results from the colonies, suggest that this category might play a compensatory role in little auk chick diets. The ability to forage on diverse taxa may help the birds to adapt to ongoing Arctic ecosystem changes.