This study investigates spatial and temporal patterns in zooplankton abundance and community composition in the Fram Strait between 2011 and 2023. The region is shaped by contrasting hydrographic regimes: warm, saline Atlantic Water transported northward by the West Spitsbergen Current (WSC) in the east, and cold, fresh Polar Water carried southward by the East Greenland Current (EGC) in the west. While the WSC has been extensively studied, the EGC remains less well understood. Yet, monitoring both regimes is essential to detect changes related to ongoing Atlantification. As part of the LTER observatory HAUSGARTEN, zooplankton were sampled at nine sites across Fram Strait using vertical multinet hauls (150 mu m mesh, 1500-1000-500-200-50-0 m depth). Preserved samples were scanned and analysed using the web-based platform EcoTaxa. In total 86 taxa were identified, with Copepoda, particularly Oithona and Calanus spp., dominating the communities. Abundances in the EGC were relatively stable and consistently lower than in the WSC. In the WSC, zooplankton abundances varied considerably and occasionally reached extremely high values that might be linked to mesoscale hydrographic features. We applied Generalized Additive Models to our data, that revealed significantly declining abundances over time in the WSC while they increased in the EGC. Shannon diversity and Pielou's evenness, however, had increased slightly in both regions in the last three years. Climate change is likely to affect the two regimes differently, leading to diverging ecosystem responses. Thus, our findings underscore the importance of sustained long-term monitoring to fully understand zooplankton dynamics in the entire Fram Strait.

Embedded in the interdisciplinary design of pelagic and benthic long-term observations at the Arctic long-term ecological research site HAUSGARTEN in Fram Strait (78 degrees 36,46 ' N-79 degrees 56,63 ' N 10 degrees 50,91 ' E-5 degrees 22,24 ' W; 250-5500 m), environmental DNA (eDNA)-based biodiversity studies span multiple habitats and trophic levels. These studies provide valuable insights into marine ecosystem functionality and carbon fluxes, including seasonal and interannual dynamics. Since 2009, eDNA studies of pelagic eukaryotic microbial communities have regularly been conducted. Recently, these studies have been expanded to cover the biodiversity of bacteria, archaea and metazoa such as zooplankton (including gelatinous taxa) and nekton (cephalopods and fish) alongside with microbial eukaryotes. The eDNA surveys have been coupled with measurements of physical and chemical properties, such as sea-ice coverage and water masses. By applying these approaches across spatial and trophic scales in a nested approach, we characterize the dynamics in biodiversity, its role in ecological processes, with a focus on current climate change. This is achieved by considering the complexity and interconnectivity of organisms, integrating information from different disciplines across scales and habitats, and focusing on functional processes including species composition. The resulting data improve our predictive capabilities for conservation and sustainable marine management in a rapidly changing Arctic. Here, we review the design of the first interdisciplinary Arctic eDNA long-term observatory and the results of more than ten years of method development and observations.

Seasonal vertical migration of large lipid-rich copepods is often described as a mass descent of animals when primary production ceases, with important implications for mesopelagic food webs and global carbon sequestration. This view ignores the existence of surface-resident individuals, but here we show that non-migrants can form a substantial part of the populations of polar migrant species. In the Central Arctic Ocean, the biomass-dominant Calanus hyperboreus was evenly distributed throughout the water column from November 2019 to March 2020, with ~20% of subadults and adult females remaining in the upper 200 m and ~41% migrating to 1000-2000 m. These vertical positions aligned with differences in the copepods' cholesterol content, which can enhance the tissue density at higher temperatures. Gonad development and the vertical distribution of their offspring indicate that both non-migrant and migrant females contribute to the population recruitment. We reinterpret copepod seasonal migration as a bet-hedging strategy that balances nutritional benefits near the surface with survival benefits at depth, and thereby contributes to the species' resilience under climatic change.

Due to the high sensitivity of zooplankton to environmental fluctuations, monitoring their taxonomic composition, abundance and biomass is of high priority to identify changes in the ecosystem. Recent advances in imaging and molecular technologies promise to greatly accelerate the processing of samples to determine both the diversity and quantity of the zooplankton community. In our study, we analyzed the diversity and quantity of an epipelagic Arctic zooplankton community using multi-marker metabarcoding and imaging analysis (ZooScan). We identified a total of 11 phyla and 58 species in the northern Barents Sea and the Nansen Basin. Metabarcoding identified more taxa than image analysis, while imaging provided quantitative information on abundance and biomass. Multivariate analyses revealed overall the same significant environmental drivers (temperature and percentage of Polar Surface Water in the sampling depth layer) explaining the similarity and spatial distribution of the zooplankton community. For all approaches, similar spatial patterns of the zooplankton community were found. Abundance, biovolume and biomass decreased with increasing latitude within the analyzed regions. Based on this study, we recommend ZooScan image analysis in combination with COI metabarcoding for future monitoring of Arctic zooplankton diversity and quantification to ensure the detection of changes in both aspects of these communities.

The international and interdisciplinary sea-ice drift expedition "The Multidisciplinary drifting Observatory for the Study of Arctic Climate" (MOSAiC) was conducted from October 2019 to September 2020. The aim of MOSAiC was to study the interconnected physical, chemical, and biological characteristics and processes from the atmosphere to the deep sea of the central Arctic system. The ecosystem team addressed current knowledge gaps and explored unknown biological properties over a complete seasonal cycle focusing on three major research areas: biodiversity, biogeochemical cycles, and linkages to the environment. In addition to the measurements of core properties along a complete seasonal cycle, dedicated projects covered specific processes and habitats, or organisms on higher taxonomic or temporal resolution in specific time windows. A wide range of sampling instruments and approaches, including sea-ice coring, lead sampling with pumps, rosette-based water sampling, plankton nets, remotely operated vehicles, and acoustic buoys, was applied to address the science objectives. Further, a broad range of process-related measurements to address, for example, productivity patterns, seasonal migrations, and diversity shifts, were made both in situ and onboard RV Polarstern. . This article provides a detailed overview of the sampling approaches used to address the three main science objectives. It highlights the core sampling program and provides examples of habitat- or process-specific sampling. The initial results presented include high biological activities in wintertime and the discovery of biological hotspots in underexplored habitats. The unique interconnectivity of the coordinated sampling efforts also revealed insights into cross-disciplinary interactions like the impact of biota on Arctic cloud formation. This overview further presents both lessons learned from conducting such a demanding field campaign and an outlook on spin-off projects to be conducted over the next years.

Polar cod (Boreogadus saida) is an endemic key species of the Arctic Ocean ecosystem. The ecology of this forage fish is well studied in Arctic shelf habitats where a large part of its population lives. However, knowledge about its ecology in the central Arctic Ocean (CAO), including its use of the sea-ice habitat, is hitherto very limited. To increase this knowledge, samples were collected at the under-ice surface during several expeditions to the CAO between 2012 and 2020, including the Multidisciplinary drifting Observatory for the Study of Arctic Climate (MOSAiC) expedition. The diet of immature B. saida and the taxonomic composition of their potential prey were analysed, showing that both sympagic and pelagic species were important prey items. Stomach contents included expected prey such as copepods and amphipods. Surprisingly, more rarely observed prey such as appendicularians, chaetognaths, and euphausiids were also found to be important. Comparisons of the fish stomach contents with prey distribution data suggests opportunistic feeding. However, relative prey density and catchability are important factors that determine which type of prey is ingested. Prey that ensures limited energy expenditure on hunting and feeding is often found in the stomach contents even though it is not the dominant species present in the environment. To investigate the importance of prey quality and quantity for the growth of B. saida in this area, we measured energy content of dominant prey species and used a bioenergetic model to quantify the effect of variations in diet on growth rate potential. The modeling results suggest that diet variability was largely explained by stomach fullness and, to a lesser degree, the energetic content of the prey. Our results suggest that under climate change, immature B. saida may be at least equally sensitive to a loss in the number of efficiently hunted prey than to a reduction in the prey's energy content. Consequences for the growth and survival of B. saida will not depend on prey presence alone, but also on prey catchability, digestibility, and energy content.

In Fram Strait, we combined underway-sampling using the remote-controlled Automated Filtration System for Marine Microbes (AUTOFIM) with CTD-sampling for eDNA analyses, and with high-resolution optical measurements in an unprecedented approach to determine variability in plankton composition in response to physical forcing in a sub-mesoscale filament. We determined plankton composition and biomass near the surface with a horizontal resolution of ~ 2 km, and addressed vertical variability at five selected sites. Inside and near the filament, plankton composition was tightly linked to the hydrological dynamics related to the presence of sea ice. The comprehensive data set indicates that sea-ice melt related stratification near the surface inside the sub-mesoscale filament resulted in increased sequence abundances of sea ice-associated diatoms and zooplankton near the surface. In analogy to the physical data set, the underway eDNA data, complemented with highly sampled phytoplankton pigment data suggest a corridor of 7 km along the filament with enhanced photosynthetic biomass and sequence abundances of sea-ice associated plankton. Thus, based on our data we extrapolated an area of 350 km2 in Fram Strait with enhanced plankton abundances, possibly leading to enhanced POC export in an area that is around a magnitude larger than the visible streak of sea-ice.

In marine ecosystems, most physiological, ecological, or physical processes are size dependent. These include metabolic rates, the uptake of carbon and other nutrients, swimming and sinking velocities, and trophic interactions, which eventually determine the stocks of commercial species, as well as biogeochemical cycles and carbon sequestration. As such, broad-scale observations of plankton size distribution are important indicators of the general functioning and state of pelagic ecosystems under anthropogenic pressures. Here, we present the first global datasets of the Pelagic Size Structure database (PSSdb), generated from plankton imaging devices. This release includes the bulk particle normalized biovolume size spectrum (NBSS) and the bulk particle size distribution (PSD), along with their related parameters (slope, intercept, and R2) measured within the epipelagic layer (0–200 m) by three imaging sensors: the Imaging FlowCytobot (IFCB), the Underwater Vision Profiler (UVP), and benchtop scanners. Collectively, these instruments effectively image organisms and detrital material in the 7–10 000 µm size range. A total of 92 472 IFCB samples, 3068 UVP profiles, and 2411 scans passed our quality control and were standardized to produce consistent instrument-specific size spectra averaged to 1° × 1° latitude and longitude and by year and month. Our instrument-specific datasets span most major ocean basins, except for the IFCB datasets we have ingested, which were exclusively collected in northern latitudes, and cover decadal time periods (2013–2022 for IFCB, 2008–2021 for UVP, and 1996–2022 for scanners), allowing for a further assessment of the pelagic size spectrum in space and time. The datasets that constitute PSSdb's first release are available at https://doi.org/10.5281/zenodo.11050013 (Dugenne et al., 2024b). In addition, future updates to these data products can be accessed at https://doi.org/10.5281/zenodo.7998799.

Calanus hyperboreus is one of the dominant copepod species in the Arctic zooplankton communities. The impact of climate change varies among regions within the Arctic, implying that C. hyperboreus populations may be differently affected at different locations, but knowledge on seasonal population dynamics in relation to biogeography is scarce. To fill this gap, we counted C. hyperboreus in samples from sediment traps that were moored from 2009 to 2014 in three regions of the Arctic Ocean (eastern Fram Strait, northern Chukchi Sea and MacKenzie Trough). The C. hyperboreus flux increased between April and May in all regions, likely associated with the ascent from overwintering depth to the surface. In the descent period, high fluxes were observed between July and September in the Fram Strait, between September and November in the northern Chukchi Sea, and between August and October in the MacKenzie Trough, suggesting that the timing of descent varied among the regions characterized by differences in light regime, phytoplankton development and water temperature. The copepodite stage composition in the eastern Fram Strait and the MacKenzie Trough varied with season, suggesting successful local reproduction while it was uniform in the northern Chukchi Sea, possibly because the population is fueled by advection.

As Arctic sea ice deteriorates, more light enters the ocean, causing largely unknown effects on the ecosystem. Using an autonomous biophysical observatory, we recorded zooplankton vertical distribution under Arctic sea ice from dusk to dawn of the polar night. Here we show that zooplankton ascend into the under-ice habitat during autumn twilight, following an isolume of 2.4 × 10 −4 W m −2 . We applied this trigger isolume to CMIP6 model outputs accounting for incoming radiation after sunset and before sunrise of the polar night. The models project that, in about three decades, the total time spent by zooplankton in the under-ice habitat could be reduced by up to one month, depending on geographic region. This will impact zooplankton winter survival, the Arctic foodweb, and carbon and nutrient fluxes. These findings highlight the importance of biological processes during the twilight periods for predicting change in high-latitude ecosystems.

Abstract As Arctic sea ice deteriorates, more light enters the Arctic Ocean, causing largely unknown effects on the ecosystem. A novel autonomous bio-physical observatory provided the first record of zooplankton vertical distribution under sea ice drifting across the Arctic Ocean from dusk to dawn of the polar night. Its measurements revealed that zooplankton ascend into the under-ice layer during autumn twilight, following an isolume of 5.5 10-4 W m-2. We applied this trigger isolume to IPCC models enabled to incorporate incoming radiation after sunset and before sunrise of the polar night. The models project that, in about three decades, the total time spent by zooplankton in the under-ice layer will be reduced by up to one month, depending on geographic region. This will impact zooplankton winter survival, the Arctic foodweb, carbon- and nutrient fluxes. These findings highlight the importance of processes in the twilight periods for predicting change in high-latitude ecosystems.

Zooplankton play a crucial role in the ocean’s ecology, as they form a foundational component in the food chain by consuming phytoplankton or other zooplankton, supporting various marine species and influencing nutrient cycling. The vertical distribution of zooplankton in the ocean is patchy, and its relation to hydrographical conditions cannot be fully deciphered using traditional net casts due to the large depth intervals sampled. The Lightframe On-sight Keyspecies Investigation (LOKI) concentrates zooplankton with a net that leads to a flow-through chamber with a camera taking images. These high-resolution images allow for the determination of zooplankton taxa, often even to genus or species level, and, in the case of copepods, developmental stages. Each cruise produces a substantial volume of images, ideally requiring onboard analysis, which presently consumes a significant amount of time and necessitates internet connectivity to access the EcoTaxa Web service. To enhance the analyses, we developed an AI-based software framework named DeepLOKI, utilizing Deep Transfer Learning with a Convolution Neural Network Backbone. Our DeepLOKI can be applied directly on board. We trained and validated the model on pre-labeled images from four cruises, while images from a fifth cruise were used for testing. The best-performing model, utilizing the self-supervised pre-trained ResNet18 Backbone, achieved a notable average classification accuracy of 83.9%, surpassing the regularly and frequently used method EcoTaxa (default) in this field by a factor of two. In summary, we developed a tool for pre-sorting high-resolution black and white zooplankton images with high accuracy, which will simplify and quicken the final annotation process. In addition, we provide a user-friendly graphical interface for the DeepLOKI framework for efficient and concise processes leading up to the classification stage. Moreover, performing latent space analysis on the self-supervised pre-trained ResNet18 Backbone could prove advantageous in identifying anomalies such as deviations in image parameter settings. This, in turn, enhances the quality control of the data. Our methodology remains agnostic to the specific imaging end system used, such as Loki, UVP, or ZooScan, as long as there is a sufficient amount of appropriately labeled data available to enable effective task performance by our algorithms.

Pigmentation is often overlooked in zooplankton, since these organisms are mostly colorless to fit the translucid water medium. However, one of the dominant zooplankton taxa in aquatic ecosystems-copepods-often show a bright red-orange or blue coloration owing to the accumulation of carotenoid pigments in some parts of their bodies. Even though there are many functional traits describing copepod's performance (e.g., size, feeding, and reproductive modes), it is surprising that the role of such a simple and visible trait as coloration has not been studied in a coherent manner yet. Here, by reviewing 95 studies, we demonstrate that carotenoid-based pigmentation (mainly caused by astaxanthin molecules) is a widespread functional trait in freshwater and marine copepods. We propose a way to disentangle the complex and thus intriguing patterns of pigment expression along latitudinal and altitudinal gradients, addressing its relationship to diet quality and quantity, temperature, ultraviolet radiation stress, predation pressure, lipid metabolism, and reproduction. We show that large-scale variations in pigmentation are difficult to tackle because of the fundamental plasticity of this trait at short time scales (i.e., hours, days), and the most recent information about carotenoid bioconversion are addressed (genes and enzyme identification, and influence of microbiota). From this literature review, we hypothesize that pigments play a "Swiss-army knife" role for copepod's fitness, useful in various ecosystem conditions owing to the strong antioxidant power and the finely-tuned metabolism of astaxanthin. With larger antioxidant capacities (survival), higher metabolisms (growth), and more offspring in better condition (reproduction), red morphs appear more successful than their uncolored siblings. Also, the potential camouflage strategies allowed by red and blue pigmentation are discussed. We finally formulate new directions and future research fields from molecular to ecosystem scales. Routine quantifications of copepod's pigmentation through trait-based approaches could be useful (1) to obtain an accurate copepod fitness indicator and (2) to better estimate the transfer of antioxidant to higher trophic levels in ecosystems, including humans.

Plankton imaging systems supported by automated classification and analysis have improved ecologists' ability to observe aquatic ecosystems. Today, we are on the cusp of reliably tracking plankton populations with a suite of lab-based and in situ tools, collecting imaging data at unprecedentedly fine spatial and temporal scales. But these data have potential well beyond examining the abundances of different taxa; the individual images themselves contain a wealth of information on functional traits. Here, we outline traits that could be measured from image data, suggest machine learning and computer vision approaches to extract functional trait information from the images, and discuss promising avenues for novel studies. The approaches we discuss are data agnostic and are broadly applicable to imagery of other aquatic or terrestrial organisms.

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.

Comau Fjord is a stratified Chilean Patagonian Fjord characterized by a shallow brackish surface layer and a >400 m layer of aragonite-depleted subsurface waters. Despite the energetic burden of low aragonite saturation levels to calcification, Comau Fjord harbours dense populations of cold-water corals (CWC). While this paradox has been attributed to a rich supply of zooplankton, supporting abundance and biomass data are so far lacking. In this study, we investigated the seasonal and diel changes of the zooplankton community over the entire water column. We used a Nansen net (100 µm mesh) to take stratified vertical hauls between the surface and the bottom (0-50-100-200-300-400-450 m). Samples were scanned with a ZooScan, and abundance, biovolume and biomass were determined for 41 taxa identified on the web-based platform EcoTaxa 2.0. Zooplankton biomass was the highest in summer (209 g dry mass m−2) and the lowest in winter (61 g dry mass m−2). Abundance, however, peaked in spring, suggesting a close correspondence between reproduction and phytoplankton spring blooms (Chl a max. 50.86 mg m−3, 3 m depth). Overall, copepods were the most important group of the total zooplankton community, both in abundance (64–81%) and biovolume (20–70%) followed by mysids and chaetognaths (in terms of biovolume and biomass), and nauplii and Appendicularia (in terms of abundance). Throughout the year, diel changes in the vertical distribution of biomass were found with a daytime maximum in the 100–200 m depth layer and a nighttime maximum in surface waters (0–50 m), associated with the diel vertical migration of the calanoid copepod family Metridinidae. Diel differences in integrated zooplankton abundance, biovolume and biomass were probably due to a high zooplankton patchiness driven by biological processes (e.g., diel vertical migration or predation avoidance), and oceanographic processes (estuarine circulation, tidal mixing or water column stratification). Those factors are considered to be the main drivers of the zooplankton vertical distribution in Comau Fjord.

The pelagic ecosystem of the Arctic Ocean is threatened by severe changes such as the reduction in sea‐ice coverage and increased inflow of warmer Atlantic water. The latter is already altering the zooplankton community, highlighting the need for monitoring studies. It is therefore essential to accelerate the taxonomic identification to speed up sample analysis, and to expand the analysis to biomass and size assessments, providing data for modeling efforts. Our case study in Fram Strait illustrates that image‐based analyses with the ZooScan provide abundance data and taxonomic resolutions that are comparable to microscopic analyses and are suitable for zooplankton monitoring purposes in the Arctic. We also show that image analysis allows to differentiate developmental stages of the key species Calanus spp. and Metridia longa and, thus, to study their population dynamics. Our results emphasize that older preserved samples can be successfully reanalyzed with ZooScan. To explore the applicability of image parameters for calculating total mesozooplankton and Calanus spp. biomasses, we used (1) conversion factors (CFs) translating wet mass to dry mass (DM), and (2) length–mass (LM) relationships. For Calanus spp., the calculated biomass values yielded similar results as direct DM measurements. Total mesozooplankton biomass ranged between 1.6 and 15 (LM) or 2.4 and 21 (CF) g DM m −2 , respectively, which corresponds to previous studies in Fram Strait. Ultimately, a normalized biomass size spectra analysis provides 1 st insights into the mesozooplankton size structure at different depths, revealing steep slopes in the linear fit in communities influenced by Atlantic water inflow.

The retreating ice cover of the Central Arctic Ocean (CAO) fuels speculations on future fisheries. However, very little is known about the existence of harvestable fish stocks in this 3.3 million-square kilometer ecosystem around the North Pole. Crossing the Eurasian Basin, we documented an uninterrupted 3170-kilometer-long deep scattering layer (DSL) with zooplankton and small fish in the Atlantic water layer at 100- to 500-meter depth. Diel vertical migration of this central Arctic DSL was lacking most of the year when daily light variation was absent. Unexpectedly, the DSL also contained low abundances of Atlantic cod, along with lanternfish, armhook squid, and Arctic endemic ice cod. The Atlantic cod originated from Norwegian spawning grounds and had lived in Arctic water temperature for up to 6 years. The potential fish abundance was far below commercially sustainable levels and is expected to remain so because of the low productivity of the CAO.

Consumer regulation of lipid composition during assimilation of dietary items is related to their ecology, habitat, and life cycle, and may lead to extra energetic costs associated with the conversion of dietary material into the fatty acids (FAs) necessary to meet metabolic requirements. For example, lipid-rich copepods from temperate and polar latitudes must convert assimilated dietary FAs into wax esters, an efficient type of energy storage which enables them to cope with seasonal food shortages and buoyancy requirements. Lipid-poor copepods, however, tend to not be as constrained by food availability as their lipid-rich counterparts and, thus, should have no need for modifying dietary FAs. Our objective was to test the assumption that Temora longicornis, a proxy species for lipid-poor copepods, does not regulate its lipid composition. Isotopically-enriched (13C) diatoms were fed to copepods during a 5-day laboratory experiment. Compound-specific stable isotope analysis of algae and copepod samples was performed in order to calculate dietary FA assimilation, turnover, and assimilation efficiency into copepod FAs. Approximately 65% of the total dietary lipid carbon (C) assimilated (913 ± 68 ng C ind-1 at the end of the experiment) was recorded as polyunsaturated FAs, with 20 and 15% recorded as saturated and monounsaturated FAs, respectively. As expected, T. longicornis assimilated dietary FAs in an unregulated, non-homeostatic manner, as evidenced by the changes in its FA profile, which became more similar to that of their diet. Copepods assimilated 11% of the total dietary C (or 40% of the dietary lipid C) ingested in the first two days of the experiment. In addition, 34% of their somatic growth (in C) after two days was due to the assimilation of dietary C in FAs. Global warming may lead to increased proportions of smaller copepods in the oceans, and to a lower availability of algae-produced essential FAs. In order for changes in the energy transfer in marine food webs to be better understood, it is important that future investigations assess a broader range of diets as well as lipid-poor zooplankton from oceanographic areas throughout the world’s oceans.