The effects of abrupt topography on plankton dynamics

Plankton population dynamics in the upper layer of the ocean depends on upwelling processes that bring nutrients from deeper waters. In turn, these depend on the structure of the vertical velocity field. In coastal areas and in oceanic regions characterized by the presence of strong submarine topographic features, the variable bottom topography induces significant effects on vertical velocities and upwelling/downwelling patterns. As a consequence, large plankton and fish abundances are frequently observed above seamounts, canyons and steep continental shelves. In this work, the dynamics of an NPZ (nutrient-phytoplankton-zooplankton) system is numerically studied by coupling the ecosystem model with a quasi two-dimensional (2D) fluid model with topography. At variance with classical 2D approaches, this formulation allows for an explicit expression of the vertical motions produced when fluid columns are squeezed and stretched as they experience changes of depth. Thus, input or output of nutrients at the surface are associated with fluid motion over the bottom topography. We examine the dynamics of a cyclonic vortex over two basic topographies: a steep escarpment and a submarine mountain. We show that plankton abundance over the escarpment is modulated by the passing of topographic Rossby waves, generated by the vortex-topography interaction. In such configuration, advection effects driven by the flow over the escarpment are of limited relevance for the dynamics of biological fields. By contrast, we find that the flow resulting from the interaction of a vortex with a seamount is sufficiently strong and persistent to allow for a remarkable increase of nutrients, and a corresponding enhancement of phytoplankton and zooplankton concentrations. Over the seamount, advection effects associated with trapped flow perturbations around the summit play an essential role.     

The effects of abrupt topography on plankton dynamics

Plankton population dynamics in the upper layer of the ocean depends on upwelling processes that bring nutrients from deeper waters. In turn, these depend on the structure of the vertical velocity field. In coastal areas and in oceanic regions characterized by the presence of strong submarine topographic features, the variable bottom topography induces significant effects on vertical velocities and upwelling/downwelling patterns. As a consequence, large plankton and fish abundances are frequently observed above seamounts, canyons and steep continental shelves. In this work, the dynamics of an NPZ (nutrient-phytoplankton-zooplankton) system is numerically studied by coupling the ecosystem model with a quasi two-dimensional (2D) fluid model with topography. At variance with classical 2D approaches, this formulation allows for an explicit expression of the vertical motions produced when fluid columns are squeezed and stretched as they experience changes of depth. Thus, input or output of nutrients at the surface are associated with fluid motion over the bottom topography. We examine the dynamics of a cyclonic vortex over two basic topographies: a steep escarpment and a submarine mountain. We show that plankton abundance over the escarpment is modulated by the passing of topographic Rossby waves, generated by the vortex-topography interaction. In such configuration, advection effects driven by the flow over the escarpment are of limited relevance for the dynamics of biological fields. By contrast, we find that the flow resulting from the interaction of a vortex with a seamount is sufficiently strong and persistent to allow for a remarkable increase of nutrients, and a corresponding enhancement of phytoplankton and zooplankton concentrations. Over the seamount, advection effects associated with trapped flow perturbations around the summit play an essential role.     

Seasonal plankton dynamics along a cross-shelf gradient

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.     

Modeling the effects of doliolids on the plankton community structure of the southeastern US continental shelf

A model of the lower trophic levels that consists of a system of coupled ordinary differential equations was developed to investigate the time-dependent behavior of doliolid populations associated with upwelling features on the outer southeastern US continental shelf. Model equations describe the interactions of doliolids with two phytoplankton size fractions, five copepod developmental states and detrital pool. Additional equations describe nitrate and ammonia. Model dynamics are based primarily upon data obtained from field and laboratory experiments for southeastern US continental shelf plankton populations. Variations on a reference simulation, which represents average upwelling conditions without doliolids, were carried out to determine the effect of inclusion of doliolids, temperature and nutrient variations, and variations in ambient food concentrations on the basic plankton community structure. These simulations provide a measure of the role of environmental versus biological interactions in structuring the planktonic food web on the southeastern US continental shelf. Simulations show that he copepod population is significantly reduce when doliolids are present. This happens primarily as a result of direct predation of the doliolids on copepod eggs and juveniles as opposed to an increase in competition for phytoplankton, the primary food source. Additional simulations show that the cooler temperatures associated with the newly upwelled water temporarily decrease the growth rates of doliolids and copepods, bestowing an event greater advantage on the rapidly reproducing doliolids.      

Simulated annual plankton production in the northeastern Pacific Coastal Upwelling Domain

A microcomputer simulation model is presented that describesthe generalized plankton production dynamics, in the surfacemixed layer, of the Juan de Fuca Eddy located on the southwesternBritish Columbia continental shelf. The Juan de Fuca Eddy simulationmodel evaluates how the annual biomass production of diatoms,copepods and euphausiids is forced by plankton feeding interactions,seasonal variability in upwelling, water temperature and solarradiation, and generalized fish predation. The model estimatesannual primary production of 345 g C m^–2 year^–1and secondary production of 19.4 g C m^–2 year^–1for copepods and 6 g C m^–2 year^–1 for euphausiids,during 1985–89; -90% of the annual plankton productionwas generated during the April-October upwelling season. Perturbationsof 22 abiotic and biotic parameters, one at a time by ±10%of nominal values, indicated that oceanic variability (e.g.upwelling rate) most strongly affected primary production. Conversely,zooplankton production was most sensitive to variability inbiological parameters describing zooplankton grazing potentialand growth (e.g. gross growth efficiency). Simulated seasonalbiomass patterns of diatoms, copepods and euphausiids were foundto closely match empirical data. However, euphausiid biomassproduction in the Juan de Fuca Eddy alone was unable to meetthe demands of estimated pelagic fish consumption. Local Eddyeuphausiid populations had to be supplemented, from regionaleuphausiids. by a mechanism that is proposed to be linked tothe seasonal pattern and intensity of positive Ekman transport(upwelling).     

The Effects of Hydrology on Plankton Biomass in Shallow Lakes of the Pampa Plain

Climatic and hydrological variability is usually high in the Pampa Plain (Argentina). However it has not studied yet how this variability may affect the phytoplankton and zooplankton biomass and community structure in aquatic systems of this region. The main purpose of this study was to assess flushing effects on nutrient and plankton dynamics in two interconnected very shallow lakes of the Pampa Plain. In order to study the impact of hydrology on the plankton biomass and community structure, we compared the summer plankton community among three consecutive years with contrasting hydrological characteristics. Water residence time varied an order of magnitude among years and this variability was correlated to strong changes in physicochemical and biological lake characteristics. Depending on the water discharge level, the hydrological regime within the lakes ranged from lentic to more lotic conditions. Nutrient and phytoplankton biomass were positively related to water discharges. During high flushing periods, nutrients import from intensive agriculture lands leads to a dramatic increase in trophic conditions. On the other hand, macrozooplankton biomass was positively related to water residence time and showed a dramatic decrease during high flushing years. Rotifers biomass was not affected by interannual water discharge variability during the study period. Our results support that in case of lakes with high flushing rates, zooplankton development is dependent on water residence time and that hydrology may have stronger effects on macrozooplankton biomass than top-down control by planktivores.     

Predicting population consequences of ocean climate change for an ecosystem sentinel,the seabird Cassin's auklet

Forecasting the ecological effects of climate change on marine species is critical for informing greenhouse gas mitigation targets and developing marine conservation strategies that remain effective and increase species' resilience under changing climate conditions. Highly productive coastal upwelling systems are predicted to experience substantial effects from climate change, making them priorities for ecological forecasting. We used a population modeling approach to examine the consequences of ocean climate change in the California Current upwelling ecosystem on the population growth rate of the planktivorous seabird Cassin's auklet (Ptychoramphus aleuticus), a demographically sensitive indicator of marine climate change. We use future climate projections for sea surface temperature and upwelling intensity from a regional climate model to forecast changes in the population growth rate of the auklet population at the important Farallon Island colony in central California. Our study projected that the auklet population growth rate will experience an absolute decline of 11–45% by the end of the century, placing this population on a trajectory toward extinction. In addition, future changes in upwelling intensity and timing of peak upwelling are likely to vary across auklet foraging regions in the California Current Ecosystem (CCE), producing a mosaic of climate conditions and ecological impacts across the auklet range. Overall, the Farallon Island Cassin's auklet population has been declining during recent decades, and ocean climate change in this century under a mid‐level emissions scenario is projected to accelerate this decline, leading toward population extinction. Because our study species has proven to be a sensitive indicator of oceanographic conditions in the CCE and a powerful predictor of the abundance of other important predators (i.e. salmon), the significant impacts we predicted for the Cassin's auklet provide insights into the consequences that ocean climate change may have for other plankton predators in this system.     

Microplanktonic regeneration of ammonium and dissolved organic nitrogen in the upwelling area of the NW of Spain: relationships with dissolved organic carbon production and phytoplankton size-structure

Ammonium regeneration and dissolved organic nitrogen (DON) releasewere studied experimentally in the euphotic zone of shelf andoceanic waters of NW Spain in relation to coastal upwellingdynamics and the size-structure of phytoplankton communities.Incubations of plankton labelled with [¹⁵N]ammonium were madeduring four cruises, two of which also included size-fractionateddeterminations of chlorophyll a and primary production, andexperimental determinations of production rates of dissolvedorganic carbon (DOC) using ¹⁴C. Inorganic nitrogen concentrationswere mainly related to nitrate enrichment by upwelling pulses,while ammonium concentrations were generally low in all situations.Ammonium did not accumulate in the study area, suggesting adaily time scale coupling between regeneration and uptake. Incontrast, DON largely exceeded inorganic nitrogen in all situationsand generally increased from spring to autumn. Ammonium regenerationwas positively correlated with DON release and both rates showedthe largest variation in summer, with minimum values duringactive upwelling and maximum values when upwelling relaxed.Comparison of DON stocks and rates in different shelf areassuggests that DON release near the coast during summer was morepersistent in the water than DON release in off-shelf and oceanicareas. The carbon:nitrogen ratio of DOC and DON release rateswas highly variable, revealing a large excess of DOC comparedwith DON. This excess can be attributed to either an underestimateof total DON release (as only release from ammonium was measured)or to an enhanced production of carbon-rich organic substancesby diatoms in coastal areas. By considering a broad range oftrophic situations, this study reveals a fundamental differencebetween short term release of DOC and DON by plankton. Physiologicalprocesses (such as carbohydrate exudation by diatoms) seem tobe the cause of large DOC excess, whereas trophic processes(such as grazing) are more likely the cause of enhanced DONrelease.     

Seasonal Synechococcus and Thaumarchaeal population dynamics examined with high resolution with remote in situ instrumentation

Monterey Bay, CA is an Eastern boundary upwelling system that is nitrogen limited much of the year. In order to resolve population dynamics of microorganisms important for nutrient cycling in this region, we deployed the Environmental Sample Processor with quantitative PCR assays targeting both ribosomal RNA genes and functional genes for subclades of cyanobacteria (Synechococcus) and ammonia-oxidizing Archaea (Thaumarchaeota) populations. Results showed a strong correlation between Thaumarchaea abundances and nitrate during the spring upwelling but not the fall sampling period. In relatively stratified fall waters, the Thaumarchaeota community reached higher numbers than in the spring, and an unexpected positive correlation with chlorophyll concentration was observed. Further, we detected drops in Synechococcus abundance that occurred on short (that is, daily) time scales. Upwelling intensity and blooms of eukaryotic phytoplankton strongly influenced Synechococcus distributions in the spring and fall, revealing what appear to be the environmental limitations of Synechococcus populations in this region. Each of these findings has implications for Monterey Bay biogeochemistry. High-resolution sampling provides a better-resolved framework within which to observe changes in the plankton community. We conclude that controls on these ecosystems change on smaller scales than are routinely assessed, and that more predictable trends will be uncovered if they are evaluated within seasonal (monthly), rather than on annual or interannual scales.     

Drivers of reef fish assemblages in an upwelling region from the Eastern Tropical Pacific Ocean

Reef fish assemblages are exposed to a wide range of anthropogenic threats as well as chronic natural disturbances. In upwelling regions, for example, there is a seasonal influx of cool nutrient-rich waters that may shape the structure and composition of reef fish assemblages. Given that climate change may disrupt the natural oceanographic processes by altering the frequency and strength of natural disturbances, understanding how fish assemblages respond to upwelling events is essential to effectively manage reef ecosystems under changing ocean conditions. This study used the baited remote underwater video stations (BRUVS) and the traditional underwater visual census (UVC) to investigate the spatiotemporal patterns of reef fish assemblages in an upwelling region in the North Pacific of Costa Rica. A total of 183 reef fish species from 60 families were recorded, of which 166 species were detected using BRUVS and 122 using UVC. Only 66% of all species were detected using both methods. This study showed that the upwelling had an important role in shaping reef fish assemblages in this region, but there was also a significant interaction between upwelling and location. In addition, other drivers such as habitat complexity and habitat composition had an effect on reef fish abundances and species. To authors’ knowledge, this is the first study in the Eastern Tropical Pacific that combines BRUVS and UVC to monitor reef fish assemblages in an upwelling region, which provides more detailed information to assess the state of reef ecosystems in response to multiple threats and changing ocean conditions.     

Invasion Waves in Populations with Excitable Dynamics

Whilst the most obvious mechanism for a biological invasion is the occupation of a new territory as a result of direct ingress by individuals of the invading population, a more subtle “invasion” may occur without significant motion of invading individuals if the population dynamics in a predator prey scenario has an “excitable” character. Here, “excitable” means that a local equilibrium state, either of coexistence of predator and prey, or of prey only, may, when disturbed by a small perturbation, switch to a new, essentially invaded state. In an invasion of this type little spatial movement of individuals occurs, but a wave of rapid change of population level nevertheless travels through the invaded territory. In this article we summarise and review recent modelling research which shows that the macroscopic features of these invasion waves depend strongly on the detailed spatial dynamics of the predator–prey relationship; the models assume simple (linear) diffusion and pursuit-evasion, represented by (non-linear) cross-diffusion, as examples. In the context of plankton population dynamics, such waves may be produced by sudden injections of nutrient and consequent rapid increase in plankton populations, brought about, for example, by the upwelling caused by a passing atmospheric low pressure system.     

The making of a productivity hotspot in the coastal ocean

Background

Highly productive hotspots in the ocean often occur where complex physical forcing mechanisms lead to aggregation of primary and secondary producers. Understanding how hotspots persist, however, requires combining knowledge of the spatio-temporal linkages between geomorphology, physical forcing, and biological responses with the physiological requirements and movement of top predators.

Methodology/Principal Findings

Here we integrate remotely sensed oceanography, ship surveys, and satellite telemetry to show how local geomorphology interacts with physical forcing to create a region with locally enhanced upwelling and an adjacent upwelling shadow that promotes retentive circulation, enhanced year-round primary production, and prey aggregation. These conditions provide an area within the upwelling shadow where physiologically optimal water temperatures can be found adjacent to a region of enhanced prey availability, resulting in a foraging hotspot for loggerhead sea turtles (Caretta caretta) off the Baja California peninsula, Mexico.

Significance/Conclusions

We have identified the set of conditions that lead to a persistent top predator hotspot, which increases our understanding of how highly migratory species exploit productive regions of the ocean. These results will aid in the development of spatially and environmentally explicit management strategies for marine species of conservation concern.     

Contribution of heterotrophic plankton to nitrogen regeneration in the upwelling ecosystem of A Coruna (NW Spain)

The contribution of heterotrophic plankton to nitrogen (N) regenerationin the water column, and its significance for the requirementsof phytoplankton, were studied at the seasonal scale in thecoastal upwelling ecosystem of A Coruña (Galicia, NWSpain). During 1995–1997, monthly measurements were takenof hydrographic conditions, dissolved nutrients, and abundanceand biomass of microplanktonic heterotrophs (bacteria, flagellatesand ciliates), phytoplankton and mesozooplankton (>200 µm).Additionally, series of experiments were conducted to quantifyN fluxes, including primary production (¹⁴C method), phytoplanktonuptake of nitrate, ammonium and urea (¹⁵N-labelling techniques),microheterotrophic regeneration of ammonium, mesozooplanktongrazing (chlorophyll gut-content method) and excretion of ammoniumby mesozooplankton. Two N budgets were built for the averagesituations of high (>100 mg C m^-2 h^-1) and low (<100 mgC m^-2 h^-1) primary production. The results revealed that phytoplanktonrelied strongly on regenerated ammonium all year round (33 and43% of total N uptake in high and low production situations,respectively). This demand for ammonium was closely matchedby regeneration rates of microplankton (0.14–0.25 mmolN m^-2 h^-1), whereas zooplankton contributed on average <10%to N regeneration. Likewise, zooplankton grazing had littledirect control on phytoplanktonic biomass. The results obtainedindicate that in the A Coruña upwelling system, N biomassof heterotrophic plankton is generally higher than phytoplanktonN biomass. The high rates of N regeneration measured also suggestthat a large proportion of the organic matter produced afteran upwelling pulse is recycled in the water column through themicrobial food web.     

Optimal wind patterns for biological production in shelf ecosystems driven by coastal upwelling

Major upwelling systems around the world provide marine productivity and fishery yield out of proportion to their area. Upwelling winds have the counteracting effects that stronger winds upwell more nutrients to the surface for higher production, but they also transport that production off continental shelves where it may not be consumed by shelf-dwelling species. Because the patterns of wind fluctuations vary in these systems, we determined the conditions for maximal biological production using a simple conveyor belt model. The conditions are that: (1) the average cross-shelf velocity produced by the winds be the value that provides the maximum production with constant winds and (2) that the wind pattern be periodic with period equal to the cross-shelf transport time that results from maximizing production with constant winds. Examination of an example using winds in central California indicated wind patterns optimal for phytoplankton occurred more frequently than those for zooplankton.     

Climatic control of upwelling variability along the western North-American coast

The high biological production of the California Current System (CCS) results from the seasonal development of equatorward alongshore winds that drive coastal upwelling. While several climatic fluctuation patterns influence the dynamics and biological productivity of the CCS, including the El Niño-Southern Oscillation (ENSO), the Pacific Decadal Oscillation index (PDO) and the North Pacific Gyre Oscillation (NPGO), the mechanisms of interaction between climatic oscillations and the CCS upwelling dynamics have remained obscure. Here, we use Singular Spectral Analysis (SSA) to reveal, for the first time, low-frequency concordance between the time series of climatic indices and upwelling intensity along the coast of western North America. Based on energy distributions in annual, semiannual and low-frequency signals, we can divide the coast into three distinct regions. While the annual upwelling signal dominates the energy spectrum elsewhere, low-frequency variability is maximal in the regions south of 33°N. Non-structured variability associated with storms and turbulent mixing is enhanced at northerly locations. We found that the low-frequency signal is significantly correlated with different climatic indices such as PDO, NPGO and ENSO with the correlation patterns being latitude-dependent. We also analyzed the correlations between this upwelling variability and sea surface temperature (SST) and sea level pressure (SLP) throughout the North Pacific to visualize and interpret the large-scale teleconnection dynamics in the atmosphere that drive the low-frequency coastal winds. These results provide new insights into the underlying mechanisms connecting climatic patterns with upwelling dynamics, which could enhance our prediction and forecast capabilities of the effects of future oceanographic and climatic variability in the CCS.     

Upwelling and downwelling at Cabo Frio (Brazil): comparison of biomass and primary production responses

Based on a comparative study of the different phases of upwellingand downwelling, a comprehensive model of average biologicalfeatures at Cabo Frio region was established. For identicalupwelling conditions, biological responses of equal range occur,allowing the establishment of some regional patterns. Thesepatterns are represented by chlorophyll values that range from0.5 to 6.0 mg m^–3, and primary production ranging from2 to 14 mg C m^–3 h^–1. During winter, in the absenceof upwelling, organic detritus constitutes the main energy sourcemaintaining the higher levels of the food web.     

Insights into the Microbial and Viral Dynamics of a Coastal Downwelling-Upwelling Transition

Although previous studies have described opposing states in upwelling regions, i.e., the rise of cold nutrient-rich waters and prevalence of surface warm nutrient-poor waters, few have addressed the transition from one state to the other. This study aimed to describe the microbial and viral structure during this transition and was able to obtain the taxonomic and metabolic compositions as well as physical-chemical data. This integrated approach allowed for a better understanding of the dynamics of the downwelling upwelling transition, suggesting that a wealth of metabolic processes and ecological interactions are occurring in the minute fractions of the plankton (femto, pico, nano). These processes and interactions included evidence of microbial predominance during downwelling (with nitrogen recycling and aerobic anoxygenic photosynthesis), different viral predation pressures over primary production in different states (cyanobacteria vs eukaryotes), and a predominance of diatoms and selected bacterial and archaeal groups during upwelling (with the occurrence of a wealth of nitrogen metabolism involving ammonia). Thus, the results provided insights into which microbes, viruses and microbial-mediated processes are probably important in the functioning of upwelling systems.     

Counterintuitive effects of global warming‐induced wind patterns on primary production in the Northern Humboldt Current System

It has been hypothesized that global warming will strengthen upwelling‐favorable winds in the Northern Humboldt Current System (NHCS) as a consequence of the increase of the land–sea thermal gradient along the Peruvian coast. The effect of strengthened winds in this region is assessed with the use of a coupled physical–biogeochemical model forced with projected and climatological winds. Strengthened winds induce an increase in primary production of 2% per latitudinal degree from 9.5°S to 5°S. In some important coastal upwelling sites primary production is reduced. This is due to a complex balance between nutrient availability, nutrient use efficiency, as well as eddy‐ and wind‐driven factors. Mesoscale activity induces a net offshore transport of inorganic nutrients, thus reducing primary production in the coastal upwelling region. Wind mixing, in general disadvantageous for primary producers, leads to shorter residence times in the southern and central coastal zones. Overall, instead of a proportional enhancement in primary production due to increased winds, the NHCS becomes only 5% more productive (+5 mol C m^?2 year^?1), 10% less limited by nutrients and 15% less efficient due to eddy‐driven effects. It is found that regions with a initial strong nutrient limitation are more efficient in terms of nutrient assimilation which makes them more resilient in face of the acceleration of the upwelling circulation.     

Contribution of Crenarchaeota and Euryarchaeota to the prokaryotic plankton in the coastal northwestern Black Sea

The NW Black Sea is an area with peculiar hydrological, hydrochemicaland biological features (river plume dynamics, coastal upwelling,intense eutrophication, blooms algae). We determined the abundanceand distribution of the two major groups of Archaea (Crenarchaeotaand Euryarchaeota) in three contrasting sites of the NW BlackSea from May to August 2004 to estimate the contribution ofnon-extremophilic Archaea to the coastal prokaryotic plankton.Using catalyzed reporter deposition-fluorescence in situ hybridization,Euryarchaeota were found to be particularly prominent (11–22%of total prokaryotic plankton) in the low-salinity waters duringlate-spring and summer, whereas Crenarchaeota contributed only5% to the prokaryotic community in these waters. The abundanceof Crenarchaeota was highest in nutrient- and particle-richwaters and lowest in offshore Black Sea surface waters. Theseresults confirm recent studies on the distribution of planktonicArchaea with Euryarchaeota dominating the archaeal communityin surface waters. Statistical analyses indicated significantcorrelations between crenarchaeotal abundance, ammonia and nitrateconcentrations at each sites. Our data show that the archaealplankton constitute an important and dynamic component of theprokaryotic plankton in the coastal oxygenated waters of theNW Black Sea and might play a significant role in the nitrogencycle.     

Synergistic Effects of Climate and Fishing in a Marine Ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere. Author Contributions  RRK conceived the project and GB analysed the data. RRK, GB and JAL co-wrote the paper.