Carbon export is facilitated by sea urchins transforming kelp detritus.

With the increasing imperative for societies to act to curb climate change by increasing carbon stores and sinks, it has become critical to understand how organic carbon is produced, released, transformed, transported, and sequestered within and across ecosystems. In freshwater and open-ocean systems, shredders play a significant and well-known role in transforming and mobilizing carbon, but their role in the carbon cycle of coastal ecosystems is largely unknown. Marine plants such as kelps produce vast amounts of detritus, which can be captured and consumed by shedders as it traverses the seafloor. We measured capture and consumption rates of kelp detritus by sea urchins across four sampling periods and over a range of kelp detritus production rates and sea urchin densities, in northern Norway. When sea urchin densities exceeded 4 m-2, the sea urchins captured and consumed a high percentage (ca. 80%) of kelp detritus on shallow reefs. We calculated that between 1.3 and 10.8 kg of kelp m-2 are shredded annually from these reefs. We used a hydrodynamic dispersal model to show that transformation of kelp blades to sea urchin feces increased its export distance fourfold. Our findings show that sea urchins can accelerate and extend the export of carbon to neighboring areas. This collector-shredder pathway could represent a significant flow of small particulate carbon from kelp forests to deep-sea areas, where it can subsidize benthic communities or contribute to the global carbon sink.

Ocean warming expands habitat of a rich natural resource and benefits a national economy.

Geographic redistribution of living natural resources changes access and thereby harvesting opportunities between countries. Internationally shared fish resources can be sensitive to shifts in the marine environment and this may have great impact on the economies of countries and regions that rely most heavily on fisheries to provide employment and food supply. Here we present a climate change-related biotic expansion of a rich natural resource with substantial economic consequences, namely the appearance of northeast Atlantic mackerel (Scomber scombrus) in Greenlandic waters. In recent years, the summer temperature has reached record highs in the Irminger Current, and this development has expanded the available and realized mackerel habitat in time and space. Observations in the Irminger Current in east Greenland in 2011 of this temperature-sensitive epipelagic fish were the first records so far northwest in the Atlantic. This change in migration pattern was followed by a rapid development of a large-scale fishery of substantial importance for the national economy of Greenland (23% of Greenland's export value of all goods in 2014). A pelagic trawl survey was conducted in mid-summer 2014 and the results showed that the bulk of ~1 million Mg (=t) of mackerel in the Irminger Current in southeast Greenland were located in the relatively warm (>8.5°C) surface layer. Mackerel was also observed in southwest Greenland. Finally, 15 CMIP5 Earth System Model projections of future marine climate were used to evaluate the epipelagic environment in Greenland. These projections for moderate and high CO2 emission scenarios (representative concentration pathways [RCP] 4.5 and 8.5) suggest how the available mackerel habitat may expand further in space and time. Overall, our results indicate that, if the stock remains large, productive, and continues its current migration pattern, then climate change has provided Greenland with a new unique opportunity for commercial exploitation. However, positive cases like this should not be cherry-picked and misused as arguments against timely and effective mitigation of climate change.

Mechanistic insights into the effects of climate change on larval cod.

Understanding the biophysical mechanisms that shape variability in fisheries recruitment is critical for estimating the effects of climate change on fisheries. In this study, we used an Earth System Model (ESM) and a mechanistic individual-based model (IBM) for larval fish to analyze how climate change may impact the growth and survival of larval cod in the North Atlantic. We focused our analysis on five regions that span the current geographical range of cod and are known to contain important spawning populations. Under the SRES A2 (high emissions) scenario, the ESM-projected surface ocean temperatures are expected to increase by >1 °C for 3 of the 5 regions, and stratification is expected to increase at all sites between 1950-1999 and 2050-2099. This enhanced stratification is projected to decrease large (>5 µm ESD) phytoplankton productivity and mesozooplankton biomass at all 5 sites. Higher temperatures are projected to increase larval metabolic costs, which combined with decreased food resources will reduce larval weight, increase the probability of larvae dying from starvation and increase larval exposure to visual and invertebrate predators at most sites. If current concentrations of piscivore and invertebrate predators are maintained, larval survival is projected to decrease at all five sites by 2050-2099. In contrast to past observed responses to climate variability in which warm anomalies led to better recruitment in cold-water stocks, our simulations indicated that reduced prey availability under climate change may cause a reduction in larval survival despite higher temperatures in these regions. In the lower prey environment projected under climate change, higher metabolic costs due to higher temperatures outweigh the advantages of higher growth potential, leading to negative effects on northern cod stocks. Our results provide an important first large-scale assessment of the impacts of climate change on larval cod in the North Atlantic.

Statistically downscaled CMIP6 ocean variables for European waters.

Climate change impact studies need climate projections for different scenarios and at scales relevant to planning and management, preferably for a variety of models and realizations to capture the uncertainty in these models. To address current gaps, we statistically downscaled (SD) 3-7 CMIP6 models for five key indicators of marine habitat conditions: temperature, salinity, pH, oxygen, and chlorophyll across European waters for three climate scenarios SSP1-2.6, SSP2-4.5, and SSP5-8.5. Results provide ensemble averages and uncertainty estimates that can serve as input data for projecting the potential success of a range of Nature-based Solutions, including the restoration of habitat-forming species such as seagrass in the Mediterranean and kelp in coastal areas of Portugal and Norway. Evaluation of the ensemble with observations from four European regions (North Sea, Baltic Sea, Bay of Biscay, and Mediterranean Sea) indicates that the SD projections realistically capture the climatological conditions of the historical period 1993-2020. Model skill (Liu-mean efficiency, Pearson correlation) clearly improves for both surface temperature and oxygen across all regions with respect to the original ESMs demonstrating a higher skill for temperature compared to oxygen. Warming is evident across all areas and large differences among scenarios fully emerge from the background uncertainties related to internal variability and model differences in the second half of the century. Scenario-specific differences in acidification significantly emerge from model uncertainty and internal variability leading to distinct trajectories in surface pH starting before mid-century (in some cases starting from present day). Deoxygenation is also present across all domains, but the climate signal was significantly weaker compared to the other two indicators when compared to model uncertainty and internal variability, and the impact of different greenhouse gas trajectories is less distinct. The substantial regional and local heterogeneity in these three abiotic indicators underscores the need for highly spatially resolved physical and biogeochemical projections to understand how climate change may impact marine ecosystems.

Impacts of climate change on water quality, benthic mussels, and suspended mussel culture in a shallow, eutrophic estuary.

Climate change is a global problem that causes severe local changes to marine biota, ecosystem functioning, and ecosystem services. The Limfjorden is a shallow, eutrophic estuary influenced by episodic summer hypoxia with an important mussel fishery and suspended mussel culture industry. Three future climate change scenarios ranging from low greenhouse gas emissions (SSP1-2.6), to intermediate (SSP2-4.5) and very high emissions (SSP5-8.5) were combined with nutrient load reductions according to the National Water Plans to investigate potential impacts on natural benthic mussel populations and suspended mussel culture for the two periods 2051-2060 and 2090-2099, relative to a reference period from 2009 to 2018. The FlexSem model combined 3D hydrodynamics with a pelagic biogeochemical model, a sediment-benthos model, and a dynamic energy budget - farm scale model for mussel culture. Model results showed that the Limfjorden was sensitive to climate change impacts with the strongest responses of physics and water quality in the worst case SSP5-8.5 scenario with no nutrient reductions. In the two low emissions scenarios, expected improvements of bottom oxygen and Chlorophyll a concentrations due to reduced nutrient loads were counteracted by climate change impacts on water physics (warming, freshening, stronger stratification). Hence, higher nutrient reductions in the Water Plans would be needed to reach a good ecological status under the influence of climate change. Suspended mussel culture was intensified in all scenarios showing a high potential harvest, whereas the benthic mussels suffered from reduced food supply and hypoxia. Provided the environmental changes and trends in social demands, in the future, it is likely that suspended mussel cultivation will become the primary source of mussels for the industry. Model scenarios can be used to inform managers, mussel farmers, fishermen, and the local population on potential future changes in bivalve harvesting and ecosystem health, and to find solutions to mitigate climate change impacts.

Does fish larval dispersal differ between high and low latitudes?

Several factors lead to expectations that the scale of larval dispersal and population connectivity of marine animals differs with latitude. We examine this expectation for demersal shorefishes, including relevant mechanisms, assumptions and evidence. We explore latitudinal differences in (i) biological (e.g. species composition, spawning mode, pelagic larval duration, PLD), (ii) physical (e.g. water movement, habitat fragmentation), and (iii) biophysical factors (primarily temperature, which could strongly affect development, swimming ability or feeding). Latitudinal differences exist in taxonomic composition, habitat fragmentation, temperature and larval swimming, and each difference could influence larval dispersal. Nevertheless, clear evidence for latitudinal differences in larval dispersal at the level of broad faunas is lacking. For example, PLD is strongly influenced by taxon, habitat and geographical region, but no independent latitudinal trend is present in published PLD values. Any trends in larval dispersal may be obscured by a lack of appropriate information, or use of 'off the shelf' information that is biased with regard to the species assemblages in areas of concern. Biases may also be introduced from latitudinal differences in taxa or spawning modes as well as limited latitudinal sampling. We suggest research to make progress on the question of latitudinal trends in larval dispersal.

Genomic differentiation in Pacific cod using Pool-Seq.

Patterns of genetic differentiation across the genome can provide insight into selective forces driving adaptation. We used pooled whole genome sequencing, gene annotation, and environmental covariates to evaluate patterns of genomic differentiation and to investigate mechanisms responsible for divergence among proximate Pacific cod (Gadus macrocephalus) populations from the Bering Sea and Aleutian Islands and more distant Washington Coast cod. Samples were taken from eight spawning locations, three of which were replicated to estimate consistency in allele frequency estimation. A kernel smoothing moving weighted average of relative divergence (F ST) identified 11 genomic islands of differentiation between the Aleutian Islands and Bering Sea samples. In some islands of differentiation, there was also elevated absolute divergence (d XY) and evidence for selection, despite proximity and potential for gene flow. Similar levels of absolute divergence (d XY) but roughly double the relative divergence (F ST) were observed between the distant Bering Sea and Washington Coast samples. Islands of differentiation were much smaller than the four large inversions among Atlantic cod ecotypes. Islands of differentiation between the Bering Sea and Aleutian Island were associated with SNPs from five vision system genes, which can be associated with feeding, predator avoidance, orientation, and socialization. We hypothesize that islands of differentiation between Pacific cod from the Bering Sea and Aleutian Islands provide evidence for adaptive differentiation despite gene flow in this commercially important marine species.

Modeled buoyancy of eggs and larvae of the deep-sea shrimp Aristeus antennatus (Crustacea: Decapoda) in the northwestern Mediterranean Sea.

Information on the buoyancy of eggs and larvae from deep-sea species is rare but necessary for explaining the position of non-swimming larvae in the water column. Due to embryonic morphology and ecology diversities, egg buoyancy has important variations within one species and among other ones. Nevertheless, it has hardly been explored if this buoyancy variability can be a strategy for deep-sea larvae to optimize their transport beyond their spawning areas. In the northwestern Mediterranean Sea, protozoea and mysis larvae of the commercial deep-sea shrimp Aristeus antennatus were recently found in upper layers, but to present, earlier stages like eggs and nauplii have not been collected. Using a Lagrangian transport model and larval characteristics, we evaluate the buoyancy and hydrodynamic effects on the transport of A. antennatus' larvae in the northwestern Mediterranean Sea. The transport models suggested that 75% of buoyant eggs released between 500 and 800 m depth (i.e., known spawning area), reached the upper water layers (0-75 m depth). Then, according to the modeled larval drifts, three spawning regions were defined in the studied area: 1) the northern part, along a continental margin crossed by large submarine canyons; 2) the central part, with two circular circulation structures (i.e., eddies); and 3) the southern part, with currents flowing through a channel. The number of larvae in the most upper layer (0-5 m depth) was higher if the larval transport model accounted for the ascent of eggs and nauplii (81%) instead of eggs reaching the surface before hatching (50%). The larvae reaching the most water upper layer (0-5 m depth) had higher rates of dispersal than the ones transported below the surface layer (deeper than 5 m depth). The results of larval dispersal simulations have implications for the understanding of A. antennatus larval ecology and for management decisions related to the shrimp fisheries in the northwestern Mediterranean Sea.

The Principles of Buoyancy in Marine Fish Eggs and Their Vertical Distributions across the World Oceans.

Buoyancy acting on plankton, i.e. the difference in specific gravity between plankton and the ambient water, is a function of salinity and temperature. From specific gravity measurements of marine fish eggs salinity appears to be the only determinant of the buoyancy indicating that the thermal expansions of the fish egg and the ambient seawater are equal. We analyze the mechanisms behind thermal expansion in fish eggs in order to determine to what extent it can be justified to neglect the effects of temperature on buoyancy. Our results confirm the earlier assumptions that salinity is the basic determinant on buoyancy in marine fish eggs that, in turn, influence the vertical distributions and, consequently, the dispersal of fish eggs from the spawning areas. Fish populations have adapted accordingly by producing egg specific gravities that tune the egg buoyancy to create specific vertical distributions for each local population. A wide variety of buoyancy adaptations are found among fish populations. The ambient physical conditions at the spawning sites form a basic constraint for adaptation. In coastal regions where salinity increases with depth, and where the major fraction of the fish stocks spawns, pelagic and mesopelagic egg distributions dominate. However, in the larger part of worlds' oceans salinity decreases with depth resulting in different egg distributions. Here, the principles of vertical distributions of fish eggs in the world oceans are presented in an overarching framework presenting the basic differences between regions, mainly coastal, where salinity increases with depth and the major part of the world oceans where salinity decreases with depth. We show that under these latter conditions, steady-state vertical distribution of mesopelagic fish eggs cannot exist as it does in most coastal regions. In fact, a critical spawning depth must exist where spawning below this depth threshold results in eggs sinking out of the water column and become lost for recruitment to the population. An example of adaptation to such conditions is Cape hake spawning above the critical layer in the Northern Benguela upwelling ecosystem. The eggs rise slowly in the onshore subsurface current below the Ekman layer, hence being advected inshore where the hatched larvae concentrate with optimal feeding conditions.

Spatial match-mismatch between juvenile fish and prey provides a mechanism for recruitment variability across contrasting climate conditions in the eastern Bering Sea.

Understanding mechanisms behind variability in early life survival of marine fishes through modeling efforts can improve predictive capabilities for recruitment success under changing climate conditions. Walleye pollock (Theragra chalcogramma) support the largest single-species commercial fishery in the United States and represent an ecologically important component of the Bering Sea ecosystem. Variability in walleye pollock growth and survival is structured in part by climate-driven bottom-up control of zooplankton composition. We used two modeling approaches, informed by observations, to understand the roles of prey quality, prey composition, and water temperature on juvenile walleye pollock growth: (1) a bioenergetics model that included local predator and prey energy densities, and (2) an individual-based model that included a mechanistic feeding component dependent on larval development and behavior, local prey densities and size, and physical oceanographic conditions. Prey composition in late-summer shifted from predominantly smaller copepod species in the warmer 2005 season to larger species in the cooler 2010 season, reflecting differences in zooplankton composition between years. In 2010, the main prey of juvenile walleye pollock were more abundant, had greater biomass, and higher mean energy density, resulting in better growth conditions. Moreover, spatial patterns in prey composition and water temperature lead to areas of enhanced growth, or growth 'hot spots', for juvenile walleye pollock and survival may be enhanced when fish overlap with these areas. This study provides evidence that a spatial mismatch between juvenile walleye pollock and growth 'hot spots' in 2005 contributed to poor recruitment while a higher degree of overlap in 2010 resulted in improved recruitment. Our results indicate that climate-driven changes in prey quality and composition can impact growth of juvenile walleye pollock, potentially severely affecting recruitment variability.

Recruitment variability in North Atlantic cod and match-mismatch dynamics.

BACKGROUND: Fisheries exploitation, habitat destruction, and climate are important drivers of variability in recruitment success. Understanding variability in recruitment can reveal mechanisms behind widespread decline in the abundance of key species in marine and terrestrial ecosystems. For fish populations, the match-mismatch theory hypothesizes that successful recruitment is a function of the timing and duration of larval fish abundance and prey availability. However, the underlying mechanisms of match-mismatch dynamics and the factors driving spatial differences between high and low recruitment remain poorly understood. METHODOLOGY/PRINCIPAL FINDINGS: We used empirical observations of larval fish abundance, a mechanistic individual-based model, and a reanalysis of ocean temperature data from 1960 to 2002 to estimate the survival of larval cod (Gadus morhua). From the model, we quantified how survival rates changed during the warmest and coldest years at four important cod spawning sites in the North Atlantic. The modeled difference in survival probability was not large for any given month between cold or warm years. However, the cumulative effect of higher growth rates and survival through the entire spawning season in warm years was substantial with 308%, 385%, 154%, and 175% increases in survival for Georges Bank, Iceland, North Sea, and Lofoten cod stocks, respectively. We also found that the importance of match-mismatch dynamics generally increased with latitude. CONCLUSIONS/SIGNIFICANCE: Our analyses indicate that a key factor for enhancing survival is the duration of the overlap between larval and prey abundance and not the actual timing of the peak abundance. During warm years, the duration of the overlap between larval fish and their prey is prolonged due to an early onset of the spring bloom. This prolonged season enhances cumulative growth and survival, leading to a greater number of large individuals with enhanced potential for survival to recruitment.

Real-time ichthyoplankton drift in Northeast Arctic cod and Norwegian spring-spawning herring.

BACKGROUND: Individual-based biophysical larval models, initialized and parameterized by observations, enable numerical investigations of various factors regulating survival of young fish until they recruit into the adult population. Exponentially decreasing numbers in Northeast Arctic cod and Norwegian Spring Spawning herring early changes emphasizes the importance of early life history, when ichthyoplankton exhibit pelagic free drift. However, while most studies are concerned with past recruitment variability it is also important to establish real-time predictions of ichthyoplankton distributions due to the increasing human activity in fish habitats and the need for distribution predictions that could potentially improve field coverage of ichthyoplankton. METHODOLOGY/PRINCIPAL FINDINGS: A system has been developed for operational simulation of ichthyoplankton distributions. We have coupled a two-day ocean forecasts from the Norwegian Meteorological Institute with an individual-based ichthyoplankton model for Northeast Arctic cod and Norwegian Spring Spawning herring producing daily updated maps of ichthyoplankton distributions. Recent years observed spawning distribution and intensity have been used as input to the model system. The system has been running in an operational mode since 2008. Surveys are expensive and distributions of early stages are therefore only covered once or twice a year. Comparison between model and observations are therefore limited in time. However, the observed and simulated distributions of juvenile fish tend to agree well during early fall. Area-overlap between modeled and observed juveniles September 1(st) range from 61 to 73%, and 61 to 71% when weighted by concentrations. CONCLUSIONS/SIGNIFICANCE: The model system may be used to evaluate the design of ongoing surveys, to quantify the overlap with harmful substances in the ocean after accidental spills, as well as management planning of particular risky operations at sea. The modeled distributions are already utilized during research surveys to estimate coverage success of sampled biota and immediately after spills from ships at sea.