The abundances of zooplankton species supporting Northeast Atlantic food webs have declined over the past 60 years, and their future is uncertain given continuing climate change. Here, we develop a new high-resolution machine-learning model of key taxa, driven by past ocean conditions and trained on Continuous Plankton Recorder observations. We apply it to an ensemble of regional climate projections for the Northwest European Shelf under the high emissions scenario RCP8.5 to enhance the detectability of a clear signal of climate-driven change in zooplankton abundance. The results project large declines in abundance: ensemble mean of 58%-72% by 2050 and 84%-93% by 2100 for small copepods and large Calanus. The projections are not geographically uniform, with evidence of regional refugia. The future declines are primarily driven by increasing temperature and decreasing salinity and nutrient concentrations, which shift conditions away from the species’ environmental niches and water mass associations. Impacts on food webs would be severe unless alternative zooplankton assemblages emerged to support existing planktivorous fish and their predators. Large declines in abundance of key zooplankton species on Northwest European Shelf can be driven by rising temperatures, falling nutrient concentrations, and a slowdown in the Atlantic Meridional Overturning Circulation, according to machine learning projections of a high emissions scenario.

Declining body sizes are prevalent in marine fish and have been suggested to be a response to increasing temperatures. However, the evidence is mixed, and the underlying causes are often unknown. Here, we explore drivers of spatio-temporal patterns in size in juvenile lesser sandeel Ammodytes marinus, focusing on ongoing size declines in parts of the North Sea. We combine experimental and field data with theory to develop a biologically realistic dynamic energy budget model that explicitly models feeding, metabolism and energy allocation to produce daily predictions of sandeel length during the growth season from 1979 to 2016 in 4 North Sea sub-populations. When forced with daily temperature estimates and zooplankton data from the Continuous Plankton Recorder, model predictions largely match observed spatio-temporal patterns. Our results suggest that the most plausible driver of observed size declines in the western North Sea is declining prey densities. In contrast, the direct effect of temperature on sandeel size is small but interacts with local prey availability so that increasing temperatures may boost growth rates in areas with high food availability but reduce growth rates in areas with low food availability. Our results thus suggest that to understand the effects of climate change on fish size, we need to account for both direct physiological effects and changes in resource availability. Finally, we show that early-life phenology and turbidity (via its impact on intake rates in the visually foraging sandeel) may also impact sandeel size, highlighting the importance of broadening our view of potential drivers of size declines.

Declining body sizes are prevalent in marine fish. While these declines have been suggested to be a response to increasing temperatures, the evidence is mixed and the underlying causes of observed declines often unknown. Here, we explore drivers of spatio-temporal patterns in size in lesser sandeel (Ammodytes marinus), an important prey for seabirds and marine mammals, focusing on ongoing size declines in the North Sea. We combine experimental and field data with ecological theory to develop a biologically realistic dynamic energy budget model that explicitly models feeding, metabolism and energy allocation to produce daily predictions of size during the juvenile growth season from 1979 to 2016. When forced with daily temperature estimates and zooplankton data from the Continuous Plankton Recorder, model predictions reproduce observed spatio-temporal patterns in size well. Our results suggest that the most plausible driver of observed size declines in the western North Sea is declining prey densities. In contrast, the direct effect of temperature on sandeel size is small, but interacts with local prey availability so that the effect varies in both size and direction over space. Our results thus suggest that to understand effects of climate change on fish size we need to account for both direct physiological effects and changes in resource availability. Finally, we use the model to show that early-life phenology and turbidity (via its impact on intake rates in the visually foraging sandeel) may also impact sandeel size, highlighting the importance of broadening our view of potential drivers of size declines. ### Competing Interest Statement The authors have declared no competing interest.

Climate change is transforming marine ecosystems, opening new fishing possibilities for large-scale trawling vessels in the Arctic. This study investigates the potential for new fishing grounds to emerge in West Greenland. We employed a maximum entropy model to predict fishing suitability based on climatological and time-invariant variables alongside public fishing vessel data. The model, validated with high accuracy, identified maximum depth, ice thickness and ice concentration as the most important predictors of fishing suitability. Results indicate a 6.2% increase in whole domain suitable fishing grounds from the 2010s to the 2040s, and a 11.4% increase from the 2010s to the 2090s. This change is driven by decreasing ice coverage, allowing extended access to the productive shelf edge. Increased fishing suitability could enhance fishing opportunities leading to increased economic benefits. However, the prolonged fishing season also raises concerns about overexploitation, ecological sustainability and sediment dispersion. Earlier access would enable trawling over carbon-rich sediments for extended periods, leading to a resuspension of sediment-bound carbon. Our findings highlight the need for adaptive management strategies to balance economic gains with the protection of marine ecosystems in the face of ongoing climate changes.

Atlantic salmon (Salmo salar) populations have suffered declines across their range in recent decades, largely attributed to decreasing marine survival rates. The first few months at sea are thought to be a time of particular vulnerability, but investigation into drivers of survival is limited by a lack of knowledge of migratory paths. Here, we model the early marine migration of Atlantic salmon from Scottish and Irish rivers over 27 years. Movement is simulated over the first 3 months at sea using a Lagrangian particle tracking model (FVCOM i-state configuration model) coupled with an active swimming model that is dependent on ocean currents, salinity, and compass direction. Our model is driven by the reanalysis of a high-resolution ocean model (Scottish Shelf Waters Reanalysis Service). Differences in the speed of migrations, the proportion of time spent in different oceanographic regions, and the proportion of migrations reaching the Norwegian Sea are seen between years. These differences are related to changes in local ocean conditions: years with lower on-shelf salinity, stronger on-shelf northwest currents, and stronger shelf-edge currents were associated with greater migration success. Within years, differences in modelled migrations between rivers were best explained by their minimum distance from the continental shelf edge.

Return rates of Atlantic salmon (Salmo salar) from the sea to European rivers have declined in recent decades. The first months at sea are critical for growth and survival; recent evidence suggests that reduced food availability may be a contributory factor to the observed declines. Here, zooplankton abundance data are used to derive a measure of prey energy available to forage fish prey of salmon during early marine migration. This zooplankton prey energy has significantly and dramatically declined over much of the northeast Atlantic, and specifically within key salmon migration domains, over the past 60 years. Marine return rates from a set of southern European populations are found to exhibit clustering not entirely predictable from geographical proximity. Variability in grouped return rates from these populations is correlated with zooplankton energy on a range of scales, demonstrating the potential use of zooplankton energy as an indicator of salmon marine survival. Comparison with environmental variables derived from ocean model reanalysis data suggests zooplankton energy is regulated by a combination of climate change impacts on ecosystem productivity and multi-decadal variability in water mass influence along the migration routes.

Arctic marine ecosystems are strongly influenced by the extreme seasonality of light in the region. Accurate determination of light is essential for building a comprehensive understanding of the dynamics of animal and aquatic algae populations. Current approaches to underwater light field parameterisations rely upon shortwave radiation (300–3000 nm) estimates from satellites or surface radiometry measurements to populate full radiative transfer software. Due to the inaccessibility of many regions in the Arctic, measured data is not widely available. This study presents a model of spectrally resolved underwater light in ice‐free conditions in the Barents Sea. Given a location and time, the model accounts for downwelling spectral irradiance in the photosynthetically active radiation (PAR, 400–700 nm) range ( E D PAR ) at the ocean surface from solar, lunar, and galactic light sources, modulated by local cloud cover. We demonstrate the ability to extend over the full year into the period of Polar Night, validated in both broadband PAR and spectral domains. Using a bio‐optical model of diffuse attenuation developed for the Barents Sea, we show accurate calculations to depth for inhomogeneous water columns over a spatial‐temporal range, validated against time series irradiance data from the ArcLight observatory in Ny‐Ålesund, Svalbard and in‐situ irradiance sensors deployed in the Barents Sea. Finally, in comparison to state‐of‐the‐art radiative transfer models, averaged over the water column we demonstrate a typical mean absolute error of <1 μmol m −2 s −1 in E D PAR for overcast conditions (<6 μmol m −2 s −1 for clear‐sky) and reduced execution time of factor 20.

In the Pacific Northwest, blooms of the diatom Pseudo-nitzschia (PN) sometimes produce domoic acid, a neurotoxin that causes amnesic shellfish poisoning, leading to a Harmful Algal Bloom (HAB) event. The Pacific Northwest (PNW) HAB Bulletin project, a partnership between academic, government, and tribal stakeholders, uses a combination of beach and offshore monitoring data and ocean forecast modeling to better understand the formation, evolution, and transport of HABs in this region. This project produces periodic Bulletins to inform local stakeholders of current and forecasted conditions. The goal of this study was to help improve how the forecast model is used in the Bulletin's preparation through a retrospective particle-tracking experiment. Using past observations of beach PN cell counts, events were identified that likely originated in the Juan de Fuca eddy, a known PN hotspot, and then particle tracks were used in the model to simulate these events. A variety of "beaching definitions" were tested, based on both water depth and distance offshore, to define when a particle in the model was close enough to the coast that it was likely to correspond to cells appearing in the intertidal zone and in shellfish diets, as well as a variety of observed PN cell thresholds to determine what cell count should be used to describe an event that would warrant further action. The skill of these criteria was assessed by determining the fraction of true positives, true negatives, false positives, and false negatives within the model in comparison with observations, as well as a variety of derived model performance metrics. This analysis suggested that for our stakeholders' purposes, the most useful beaching definition is the 30 m isobath and the most useful PN cell threshold for coincident field-based sample PN density estimates is 10,000 PN cells/L. Lastly, the performance of a medium-resolution (1.5 km horizontal resolution) version of the model was compared with that of a high-resolution (0.5 km horizontal resolution) version, the latter currently used in forecasting for the PNW HAB Bulletin project. This analysis includes a direct comparison of the two model resolutions for one overlapping year (2017). These results suggested that a narrower, more realistic beaching definition is most useful in a high-resolution model, while a wider beaching definition is more appropriate in a lower resolution model like the medium-resolution version used in this analysis. Overall, this analysis demonstrated the importance of incorporating stakeholder needs into the statistical approach in order to generate the most effective decision-support information from oceanographic modeling.

The phenology, distribution, and size composition of plankton communities are changing rapidly in response to warming. This may lead to shifts in the prey fields of planktivorous fish, which play a key role in transferring energy up marine food chains. Here, we use 60 + years of Continuous Plankton Recorder data to explore temporal trends in key taxa and community traits in the prey field of planktivorous lesser sandeels (Ammodytes marinus) in the North Sea, the Faroes and southern Iceland. We found marked spatial variation in the prey field, with Calanus copepods generally being much more common in the northern part of the study area. In the western North Sea, the estimated amount of available energy in the prey field has decreased by more than 50% since the 1960s. This decrease was accompanied by declining abundances of small copepods, and shifts in the timing of peak annual prey abundances. Further, the estimated average prey community body size has increased in several of the locations considered. Overall, our results point to the importance of regional studies of prey fields, and caution against inferring ecological consequences based only on large-scale trends in key taxa or mean community traits.

The Arctic marine ecosystem is shaped by the seasonality of the solar cycle, spanning from 24-h light at the sea surface in summer to 24-h darkness in winter. The amount of light available for under-ice ecosystems is the result of different physical and biological processes that affect its path through atmosphere, snow, sea ice and water. In this article, we review the present state of knowledge of the abiotic (clouds, sea ice, snow, suspended matter) and biotic (sea ice algae and phytoplankton) controls on the underwater light field. We focus on how the available light affects the seasonal cycle of primary production (sympagic and pelagic) and discuss the sensitivity of ecosystems to changes in the light field based on model simulations. Lastly, we discuss predicted future changes in under-ice light as a consequence of climate change and their potential ecological implications, with the aim of providing a guide for future research.

Abstract. The spring phytoplankton bloom is a key event in temperate and polar seas, yet the mechanisms that trigger it remain under debate. Some hypotheses claim that the spring bloom onset occurs when light is no longer limiting, allowing phytoplankton division rates to surpass a critical threshold. In contrast, the Disturbance Recovery Hypothesis (DRH) proposes that the onset responds to an imbalance between phytoplankton growth and loss processes, allowing phytoplankton biomass to start accumulating, and this can occur even when light is still limiting. Although many studies have shown that the DRH explains the spring bloom onset in oceanic waters, it is less certain whether and how it also applies to coastal areas. To address this question at a coastal location in the Scottish North Sea, we combined 21 years (1997–2017) of weekly in situ data with meteorological information. The onset of phytoplankton biomass accumulation occurred around the same date each year, 16 ± 11 days (mean ± SD) after the winter solstice, when light limitation for growth was strongest. Also, negative and positive biomass accumulation rates (r) occurred respectively before and after the winter solstice at similar light levels. The seasonal change from negative to positive r was mainly driven by the rate of change in light availability rather than light itself. Our results support the validity of the DRH for the studied coastal region and suggest its applicability to other coastal areas.

The changing Arctic environment is affecting zooplankton that support its abundant wildlife. We examined how these changes are influencing a key zooplankton species, Calanus finmarchicus, principally found in the North Atlantic but expatriated to the Arctic. Close to the ice-edge in the Fram Strait, we identified areas that, since the 1980s, are increasingly favourable to C. finmarchicus. Field-sampling revealed part of the population there to be capable of amassing enough reserves to overwinter. Early developmental stages were also present in early summer, suggesting successful local recruitment. This extension to suitable C. finmarchicus habitat is most likely facilitated by the long-term retreat of the ice-edge, allowing phytoplankton to bloom earlier and for longer and through higher temperatures increasing copepod developmental rates. The increased capacity for this species to complete its life-cycle and prosper in the Fram Strait can change community structure, with large consequences to regional food-webs.

Bowhead whales ( Balaena mysticetus ) visit Disko Bay, West Greenland in winter and early spring to feed on Calanus spp., at a time of year when the copepods are still mostly in diapause and concentrated in near-bottom patches. Combining past observations of copepod abundance and distribution with detailed observations of bowhead whale foraging behaviour from telemetry suggests that if the whales target the highest-density patches, they likely consume 26–75% of the Calanus standing stock annually. A parallel bioenergetic calculation further suggests that the whales' patch selection must be close to optimally efficient at finding hotspots of high density copepods near the sea floor in order for foraging in Disko Bay to be a net energetic gain. Annual Calanus consumption by bowhead whales is similar to median estimates of consumption by each of three zooplankton taxa (jellies, chaetognaths, and predatory copepods), and much greater than the median estimate of consumption by fish larvae, as derived from seasonal abundance and specific ingestion rates from the literature. The copepods' self-concentration during diapause, far from providing a refuge from predation, is the behaviour that makes this strong trophic link possible. Because the grazing impact of the whales comes 6–10 months later than the annual peak in primary production, and because Disko Bay sits at the end of rapid advective pathways (here delineated by a simple numerical particle-tracking experiment), it is likely that these Calanus populations act in part as a long-distance energetic bridge between the whales and primary production hundreds or thousands of km away.

The spring phytoplankton bloom is a key event in temperate and polar seas, yet the mechanisms that trigger it remain under debate. Some hypotheses claim that the spring bloom onset occurs when light is no longer limiting, allowing phytoplankton division rates to surpass a critical threshold. In contrast, the Disturbance Recovery Hypothesis (DRH) proposes that the onset responds to an imbalance between phytoplankton growth and loss processes, allowing phytoplankton biomass to start accumulating, and this can occur even when light is still limiting. Although several studies have shown that the DRH can explain the spring bloom onset in oceanic waters, it is less certain whether and how it also applies to coastal areas. To address this question at a coastal location in the Scottish North Sea, we combined 21 years (1997–2017) of weekly in situ chlorophyll and environmental data with meteorological information. Additionally, we also analyzed phytoplankton cell counts estimated using microscopy (2000–2017) and flow cytometry (2015–2017). The onset of phytoplankton biomass accumulation occurred around the same date each year, 16 ± 11 d (mean ± SD) after the winter solstice, when light limitation for growth was strongest. Also, negative and positive biomass accumulation rates (r) occurred respectively before and after the winter solstice at similar light levels. The seasonal change from negative to positive r was mainly driven by the rate of change in light availability rather than light itself. Our results support the validity of the DRH for the studied coastal region and suggest its applicability to other coastal areas.

A realistic numerical model is used to study the circulation and mixing of the Salish Sea, a large, complex estuarine system on the United States and Canadian west coast. The Salish Sea is biologically productive and supports many important fisheries but is threatened by recurrent hypoxia and ocean acidification, so a clear understanding of its circulation patterns and residence times is of value. The estuarine exchange flow is quantified at 39 sections over 3 years (2017-2019) using the Total Exchange Flow method. Vertical mixing in the 37 segments between sections is quantified as opposing vertical transports: the efflux and reflux. Efflux refers to the rate at which deep, landward-flowing water is mixed up to become part of the shallow, seaward-flowing layer. Similarly, reflux refers to the rate at which upper layer water is mixed down to form part of the landward inflow. These horizontal and vertical transports are used to create a box model to explore residence times in a number of different sub-volumes, seasons, and years. Residence times from the box model are generally found to be longer than those based on simpler calculations of flushing time. The longer residence times are partly due to reflux, and partly due to incomplete tracer homogenization in sub-volumes. The methods presented here are broadly applicable to other estuaries.

The predation risk of many aquatic taxa is dominated by visually searching predators, commonly a function of ambient light. Several studies propose that changes in visual predation will become a major climate-change impact on polar marine ecosystems. The High Arctic experiences extreme seasonality in the light environment, from 24 h light to 24 h darkness, and therefore provides a natural laboratory for studying light and predation risk over diel to seasonal timescales. Here, we show that zooplankton (observed using acoustics) in an Arctic fjord position themselves vertically in relation to light. A single isolume (depth-varying line of constant light intensity, the value of which is set at the lower limit of photobehaviour reponses of Calanus spp. and krill) forms a ceiling on zooplankton distribution. The vertical distribution is structured by light across timescales, from the deepening of zooplankton populations at midday as the sun rises in spring, to the depth to which zooplankton ascend to feed during diel vertical migration. These results suggest that zooplankton might already follow a foraging strategy that will keep visual predation risk roughly constant under changing light conditions, such as those caused by the reduction of sea ice, but likely with energetic costs such as lost feeding opportunities as a result of altered habitat use.

In recent decades, the central Arctic Ocean has been experiencing dramatic decline in sea ice coverage, thickness and extent, which is expected to have a tremendous impact on all levels of Arctic marine life. Here, we analyze the regional and temporal changes in pan-Arctic distribution and population structure of the key zooplankton species Calanus glacialis and C. hyperboreus in relation to recent changes in ice conditions, based on historical (1993-1998) and recent (2007-2016) zooplankton collections and satellite-based sea ice observations. We found strong correlations between Calanus abundance/population structure and a number of sea ice parameters. These relationships were particularly strong for C. glacialis, with higher numbers being observed at locations with a lower ice concentration, a shorter distance to the ice edge, and more days of open water. Interestingly, early stages of C. hyperboreus followed the same trends, suggesting that these two species substantially overlap in their core distribution area in the Arctic Ocean. Calanus glacialis and C. hyperboreus have been historically classified as shelf versus basin species, yet we conclude that both species can inhabit a wide range of bottom depths and their distribution in the Arctic Ocean is largely shaped by sea ice dynamics. Our data suggest that the core distribution patterns of these key zooplankton are shifting northwards with retreating sea ice and changing climate conditions.

International Symposium Human Health & the Ocean in a Changing World, 2-3 December 2020, Monte-Carlo, Monaco