Continuous Plankton Recorder Survey Research Articles

Mapping Selected Emergent Marine Toxin-Producing Organisms Using Historical Samples with Two Methods (Biosensors and Real-Time PCR): A Comparison of Resolution

The Continuous Plankton Recorder (CPR) survey is a valuable resource for mapping changes in plankton distribution and understanding harmful algal ecology because of its breadth and longevity. Preservation methods with formalin degrade DNA, making it difficult to use as a molecular tool for archived marine samples. DNA was extracted from CPR samples immediately after collection, seven months later and after nine years of storage from a cruise track along the Iberian Peninsula. PCR reactions performed from the nine-year timepoint were hybridized to probes in an electrochemical biosensor and compared to results obtained from RT-PCR performed at two earlier time points. The successful identification of Pseudo-nitzschia spp., Prorocentrum lima, Alexandrium minutum, Alexandrium ostenfeldii, Gambierdiscus spp. and Coolia spp. was documented. The biosensor analysis outperformed RT-PCR, allowing us to document certain tropical toxic dinoflagellates, viz., Gambierdiscus and Coolia, that produce human ciguatoxins and Coolia toxins, respectively. These non-native algal toxins can accumulate, pervade the food web and negatively impact human food security. This supports the northerly movement of microalgae with climate change in offshore Iberian peninsular waters. This study highlights biosensors as a cost-effective tool for the offshore monitoring of HAB species and advances molecular technologies for long-term CPR datasets that have limited records of harmful algae. DNA from formalin-preserved CPR samples is degraded, so the use of a short, multiprobe biosensor can augment historical plankton records with contemporary methods that also capture infrequently occurring benthic taxa carried in surface waters. The integration of probe-based biosensor technologies offers a promising avenue for exploring plankton dynamics in response to environmental changes.

At what scale should we assess the health of pelagic habitats? Trade-offs between small-scale manageable pressures and the need for regional upscaling

Major planktonic lifeforms such as diatoms, dinoflagellates, meroplankton and holoplankton have recently shown significant and alarming changes in abundance - mainly downwards trends - around the northwest European shelf. This has major implications for food web connections and for ecosystem services including seafood provision and carbon storage. We have quantified these changes in abundance for 2006–2019/20 using a Plankton Index (PI) and show that the scale of spatial aggregation is critical to the ability of the PI to detect change, understand causal mechanisms, and provide advice to policymakers.We derived PI statistics in the Celtic and North Seas from data from the Continuous Plankton Recorder survey offshore and England’s Environment Agency inshore using three sets of spatial units: (i) Ecohydrodynamic (EHD) units based on hydro-biogeochemical modelling, (ii) ‘COMP4′ areas based on cluster analysis of satellite data for chlorophyll a and primary productivity, and (iii) English coastal and estuarine Water Framework Directive (WFD) waterbodies. For the largest scale areas, the EHD units (median size 87,000 km2), we find greater change in plankton communities than previously reported, suggesting that these shifts have continued and possibly intensified in recent years. The smaller-scale COMP4 areas (median size 6,700 km2) appear to encompass more spatially coherent changes in plankton community structure than EHD units; at this scale PI values indicate community shifts of greater magnitude. These COMP4 areas provide a reasonable compromise scale for linking offshore plankton communities to large-scale drivers of change such as climate warming. For inshore plankton communities, larger changes are detected at the smaller WFD waterbody scale (median size 11 km2). This scale allows direct links to coastal management measures and is more suitable for linking to land-sourced pressures.Recent integration of the UK’s OSPAR and WFD plankton monitoring data management enables the exploration of changes across spatial scales to develop a holistic understanding of ecosystem health. Regional-sea scale derivation of the PI for coastal waters provides a clear indication that changes are occurring, at least in phytoplankton communities, while localised PI statistics offer an additional layer of information which can be an important tool for linking to localised drivers of change including coastal anthropogenic pressures. Broadscale inshore zooplankton monitoring is needed to evaluate the coastal plankton community holistically; zooplankton communities offshore are also changing but these changes cannot currently be linked to coastal processes. Layering information across spatial scales provides a breadth of system-level understanding beyond what any one typology can provide.

Reinterpreting two regime shifts in North Sea plankton communities through the lens of functional traits

AbstractAimThe so‐called regime shifts in North Sea plankton communities provide an important historical case study to understand marine regime shifts. Previous studies characterized regime shifts using a variety of community metrics (e.g., indicator species abundances, taxonomic composition and chlorophyll biomass) but left the functional traits of plankton unassessed. Here, we explicitly re‐assess the historically recognized North Sea regime shifts through the lens of plankton functional traits to gain a better understanding of these events.LocationThe North Sea (NW European shelf).Time period1958–2018, focusing on the 1980s and 1996–2003 regime shifts.Major taxa studiedMarine phyto‐ and zooplankton.MethodsWe compute trait spaces for both phyto‐ and zooplankton of the North Sea using traits from the literature and a Gower's distance‐based method. Using abundance data from the Continuous Plankton Recorder Survey, we then compute monthly time series of the centroids of the communities, an indicator of functional composition. We then use principal component analysis on the centroids to assess the main temporal changes in plankton functional composition associated with the 1980s and 1996–2003 regime shifts.ResultsLittle change in plankton functional composition was associated with the 1980s regime shift. In contrast, the functional composition of plankton communities changed markedly after the 1996–2003 regime shift, with an increase in the summer relative abundance of non‐motile autotrophs (i.e., diatoms) and the spring relative abundance of meroplankton.Main conclusionsThe North Sea regime shifts were not associated systematically with changes in functional composition, calling into question the definition of regime shifts and illustrating the importance of taking different metrics into account to interpret ecological events accurately. Taking into account functional composition, we interpret the 1980s so‐called regime shift as a latitudinal shift in communities that was insufficient to impact functional composition and the 1996–2003 so‐called regime shift as a period of change in bentho‐pelagic coupling.

Historical biomonitoring of pollution trends in the North Pacific using archived samples from the Continuous Plankton Recorder Survey

First started in 1931, the Continuous Plankton Recorder (CPR) Survey is the longest-running and most geographically extensive marine plankton sampling program in the world. This pilot study investigates the feasibility of biomonitoring the spatiotemporal trends of marine pollution using archived CPR samples from the North Pacific. We selected specimens collected from three different locations (British Columbia Shelf, Northern Gulf of Alaska, and Aleutian Shelf) in the North Pacific between 2002 and 2020. Comprehensive profiling of the plankton chemical exposome was conducted using liquid and gas chromatography coupled with tandem mass spectrometry (LC-MS/MS and GC–MS/MS). Our results show that phthalates, plasticizers, persistent organic pollutants (POPs), pesticides, pharmaceuticals, and personal care products were present in the plankton exposome, and that many of these pollutants have decreased in amount over the last two decades, which was most pronounced for tri-n-butyl phosphate. In addition, the plankton exposome differed significantly by regional human activities, with the most polluted samples coming from the nearshore area. Exposome-wide association analysis revealed that bioaccumulation of environmental pollutants was highly correlated with the biomass of different plankton taxa. Overall, this study demonstrates that exposomic analysis of archived samples from the CPR Survey is effective for long-term biomonitoring of the spatial and temporal trends of environmental pollutants in the marine environment.

The species chromatogram, a new graphical method to represent, characterize, and compare the ecological niches of different species.

The ecological niche sensu Hutchinson is defined as the set of environmental conditions allowing a species to grow, maintain, and reproduce. This conception of the niche, which is assimilated to a p‐dimensional hypervolume, with p representing all environmental variables, has been widely applied in ecology. However, displaying the niche hypervolume has proved challenging when more than three environmental dimensions are considered simultaneously. We propose a simple method (implemented in the specieschrom R package) that displays the full multidimensionality of the ecological niche of a species into a two‐dimensional space by means of a graphic we call species chromatogram. This method gives a graphical summary of the niche by representing together abundance gradients with respect to all environmental variables. A chromatogram enables niche optimums and breaths to be rapidly quantified, and when several chromatograms are examined (one per species), rapid comparisons can be made. From our chromatograms, we proposed a procedure that quantifies niche optimum and breadth as well as niche overlapping (index D) and the identification of the most discriminant combination of environmental variables. We apply these analyses on eight planktonic species collected by the Continuous Plankton Recorder (CPR) survey in the North Atlantic Ocean using 10 environmental variables. We display their full multidimensional niches and quantify their niche optimums and breadths along each dimension. We also compare our index D with other indices by means of hypervolume and dynRB R packages. By catching the full complexity of the niche, species chromatograms allow many different niche properties to be rapidly assessed and compared among species from niche optimums and breadths to the identification of the most relevant environmental parameters and the degree of niche overlapping among species. Species chromatograms may be seen as species’ fingerprint and may also allow a better identification of the mechanisms involved in species assembly.

Occurrence and chemical characteristics of microplastic paint flakes in the North Atlantic Ocean

Non-fibrous microplastics sampled by the Continuous Plankton Recorder (CPR) Survey throughout the North Atlantic Ocean during 2018 have been recorded and a selection (n = 17, or 16.7%) physically and chemically characterised. The average abundance of non-fibrous particles captured by the plankton silks and detectable by microscopy was estimated to be around 0.01 m-3, with the highest concentrations evident in shelf seas of northwest Europe. Amongst the samples analysed, median size was 180 μm and, based on visible properties (e.g., brittleness, layering) and infra-red spectra, all but one were identified as flakes of paint. Semi-quantitative analysis by energy-dispersive X-ray fluorescence spectrometry with a collimated beam revealed that six flakes from European shelf seas were Cu-based antifouling formulations (without evidence of organo-Sn compounds), and five with a broader geographical distribution were Pb-based formulations of likely marine origin. Other elements regularly detected included Cr, Fe, Ti and Zn that were present in pigments or as contaminants from the underlying substrate. After fibres, paint flakes appear to be the most abundant type of microplastic in the oceans that, because of the abundance and mobility of metallic additives, deserve closer scientific attention.

Temperature and Patterns of Occurrence and Abundance of Key Copepod Taxa in the Northeast Pacific

The Northeast Pacific is a highly heterogeneous and productive ecosystem, yet it is vulnerable to climate change and extreme events such as marine heat waves. Recent heat wave induced die-offs of fish, marine mammals, and seabirds in the Gulf of Alaska were associated with the loss of large, lipid-rich copepods, which are a vital food resource for forage fishes. The critical and temperature sensitive role of copepods in this ecosystem motivates our investigation into the impacts of temperature on copepod occurrence, abundance, and phenology. Here, we pair long term in situ copepod data from Continuous Plankton Recorder surveys with satellite temperature data to determine the influence of water temperature on three key copepod taxa: Neocalanus plumchrus, Calanus pacificus, and Oithona spp. Through the use of linear models and thermal threshold methods, we demonstrate that N. plumchrus is most vulnerable to warming and future marine heat waves in this region. Linear models demonstrate that N. plumchrus abundance is negatively related to temperature, and thermal threshold methods reveal that N. plumchrus has an upper thermal threshold of 11.5°C for occurrence, and 10.5°C for abundance. Additionally, examining N. plumchrus abundance before and during the 2014–2016 marine heat wave demonstrates reduced species abundance during past warming events. Oithona spp. and C. pacificus appear to be less vulnerable to warm temperatures. However, their presence will not be sufficient to supplement the loss of the larger-bodied and lipid-rich N. plumchrus. Our findings demonstrate the power of using long-term in situ data to determine thermal tolerances, and suggest the need to further examine the potential resilience of N. plumchrus to climate change.

Zooplankton communities along a Southern Ocean monitoring transect at 110° E from three CPR surveys (Dec 2014, Jan 2015, Mar 2015)

Many studies using surface plankton data from the Continuous Plankton Recorder (CPR) have suggested that micro- and meso-sized zooplankton species/taxa can serve as indicators of environmental change. But while regular CPR sampling would allow for the detection of changes in zooplankton communities, few intra-annual CPR surveys have been undertaken along single transects. We report the temporal variability of zooplankton community structure using CPR data collected along the Japanese monitoring transect (110° E) in December 2014, January 2015, and March 2015. We found zooplankton abundances in both the Polar Frontal Zone (PFZ) and Antarctic Zone (AZ) to increase from December to January, and mean AZ abundance to peak abruptly in January (381.9 ind. m−3) before decreasing to about 25% of that value in March. We attribute changes in total zooplankton abundance to fluctuations in the numbers of the two small dominant copepods, Oithona similis and Ctenocalanus citer. High AZ phytoplankton concentrations in December likely increased surface food availability, influencing population growth, and/or the timing of copepod seasonal ascension. We recommend future CPR surveys include sampling with finer mesh for more accurate determination of abundances of small-sized zooplankton, to better understand the ramifications of climate change on Southern Ocean trophic structure.

Overwintering distribution, inflow patterns and sustainability of Calanus finmarchicus in the North Sea

Calanoid copepods are key taxa in the North Sea as they are the main food source for many fish stocks, such as herring, mackerel and cod. In this study we use an individual-based model for Calanus finmarchicus embedded in the NORWegian ECOlogical Model system (NORWECOM) to investigate important population parameters such as biomass and abundance, distribution and interannual variability of the overwintering population, as well as the inflow of C. finmarchicus into the North Sea from adjacent areas for the 2000–2016 period. The modelled spatial–temporal patterns of C. finmarchicus abundance is comparable with the Continuous Plankton Recorder (CPR) Survey data in the northern North Sea. The simulated annual mean biomass of C. finmarchicus amounts to 0.94 million-tonnes of carbon. High overwintering biomass appears in the Norwegian Trench as well as in the north-west shelf region of the North Sea. A decreasing trend in the overwintering biomass has been detected on the path of the East Shetland Atlantic Inflow (ESAI) over the simulated period. The inflow of C. finmarchicus biomass into the North Sea from the north constitutes on average 41% of the annual mean biomass in the North Sea during the simulated 17 years, and thus determines the interannual variability of the biomass. We conclude that the C. finmarchicus population in the North Sea is not self-sustained and is highly dependent on the inflow of C. finmarchicus from the Faroe-Shetland Channel and south of the Norwegian Sea. C. finmarchicus enter the North Sea via three branches of the North Atlantic current with variable depths depending on seasons and topography. Beside the western flank of the Norwegian Trench (carrying 57% of the inflow biomass), we suggest that the ESAI is also an important agent carrying 37% of the total C. finmarchicus inflow biomass through the shelf area into the north-west of the North Sea. The annual mean outflow biomass is larger than the inflow biomass (0.52 versus 0.39 million-tonnes carbon per year), which indicates that the North Sea serves as a feeding ground and growth region for C. finmarchicus. This study is a first step towards a better understanding and quantification of the exchange of C. finmarchicus between the open seas, coastal waters and the fjords.

Plankton biogeography in the North Atlantic Ocean and its adjacent seas: Species assemblages and environmental signatures.

Plankton biodiversity is a key component of marine pelagic ecosystems. They are at the base of the food web, control the productivity of marine ecosystems, and provide many provisioning and regulating ecological services. It is therefore important to understand how plankton are organized in both space and time. Here, we use data of varying taxonomic resolution, collected by the Continuous Plankton Recorder (CPR) survey, to map phytoplankton and zooplankton biodiversity in the North Atlantic and its adjacent seas. We then decompose biodiversity into 24 species assemblages and investigate their spatial distribution using ecological units and ecoregions recently proposed. Finally, we propose a descriptive method, which we call the environmental chromatogram, to characterize the environmental signature of each plankton assemblage. The method is based on a graphic that identifies where species of an assemblage aggregate along an environmental gradient composed of multiple ecological dimensions. The decomposition of the biodiversity into species assemblages allows us to show (a) that most marine regions of the North Atlantic are composed of coenoclines (i.e., gradients of biocoenoses or communities) and (b) that the overlapping spatial distribution of assemblages is the result of their environmental signatures. It follows that neither the ecoregions nor the ecological units identified in the North Atlantic are characterized by a unique assemblage but instead by a mosaic of assemblages that overlap in many places.

Bayesian Network Analysis reveals resilience of the jellyfish Aurelia aurita to an Irish Sea regime shift

Robust time-series of direct observations of jellyfish abundance are not available for many ecosystems, leaving it difficult to determine changes in jellyfish abundance, the possible causes (e.g. climate change) or the consequences (e.g. trophic cascades). We sought an indirect ecological route to reconstruct jellyfish abundance in the Irish Sea: since zooplankton are jellyfish prey, historic variability in zooplankton communities may provide proxies for jellyfish abundance. We determined the Bayesian ecological network of jellyfish–zooplankton dependencies using jellyfish- and zooplankton-abundance data obtained using nets during a 2-week cruise to the Irish Sea in 2008. This network revealed that Aurelia aurita abundance was dependent on zooplankton groups Warm Temperate and Temperate Oceanic as defined by previous zooplankton ecology work. We then determined historic zooplankton networks across the Irish Sea from abundance data from Continuous Plankton Recorder surveys conducted between 1970 and 2000. Transposing the 2008 spatial dependencies onto the historic networks revealed that Aurelia abundance was more strongly dependent over time on sea surface temperature than on the zooplankton community. The generalist predatory abilities of Aurelia may have insulated this jellyfish over the 1985 regime shift when zooplankton composition in the Irish Sea changed abruptly, and also help explain its globally widespread distribution.

Assuming independence in spatial latent variable models: Consequences and implications of misspecification

Multivariate spatial data, where multiple responses are simultaneously recorded across spatially indexed observational units, are routinely collected in a wide variety of disciplines. For example, the Southern Ocean Continuous Plankton Recorder survey collects records of zooplankton communities in the Indian sector of the Southern Ocean, with the aim of identifying and quantifying spatial patterns in biodiversity in response to environmental change. One increasingly popular method for modeling such data is spatial generalized linear latent variable models (GLLVMs), where the correlation across sites is captured by a spatial covariance function in the latent variables. However, little is known about the impact of misspecifying the latent variable correlation structure on inference of various parameters in such models. To address this gap in the literature, we investigate how misspecifying and assuming independence for the latent variables' correlation structure impacts estimation and inference in spatial GLLVMs. Through both theory and numerical studies, we show that performance of maximum likelihood estimation and inference on regression coefficients under misspecification depends on a combination of the response type, the magnitude of true regression coefficient, and the corresponding loadings, and, most importantly, whether the corresponding covariate is (also) spatially correlated. On the other hand, estimation and inference of truly nonzero loadings and prediction of latent variables is consistently not robust to misspecification of the latent variable correlationstructure.

Annual phytoplankton succession results from niche-environment interaction

AbstractAnnual plankton succession has been investigated for many decades with hypotheses ranging from abiotic to biotic mechanisms being proposed to explain these recurrent patterns. Here, using data collected by the Continuous Plankton Recorder (CPR) survey and models originating from the MacroEcological Theory on the Arrangement of Life, we investigate Annual Phytoplankton Succession (APS) in the North Sea at a species level. Our results show that this phenomenon can be predicted well by models combining photosynthetically active radiation, temperature and macro-nutrients. Our findings suggest that APS originates from the interaction between species’ ecological niches and the annual environmental fluctuations at a community level. We discuss our results in the context of traditional hypotheses formulated to explain this recurrent pattern in the marine field.

Capturing open ocean biodiversity: Comparing environmental DNA metabarcoding to the continuous plankton recorder.

Environmental DNA (eDNA) metabarcoding is emerging as a novel, objective tool for monitoring marine metazoan biodiversity. Zooplankton biodiversity in the vast open ocean is currently monitored through continuous plankton recorder (CPR) surveys, using ship-based bulk plankton sampling and morphological identification. We assessed whether eDNA metabarcoding (2L filtered seawater) could capture similar Southern Ocean zooplankton biodiversity as conventional CPR bulk sampling (~1,500L filtered seawater per CPR sample). We directly compared eDNA metabarcoding with (a) conventional morphological CPR sampling and (b) bulk DNA metabarcoding of CPR collected plankton (two transects for each comparison, 40 and 44 paired samples, respectively). A metazoan-targeted cytochrome c oxidase I (COI) marker was used to characterize species-level diversity. In the 2L seawater eDNA samples, this marker amplified large amounts of non-metazoan picoplanktonic algae, but eDNA metabarcoding still detected up to 1.6 times more zooplankton species than morphologically analysed bulk CPR samples. COI metabarcoding of bulk DNA samples mostly avoided nonmetazoan amplifications and recovered more zooplankton species than eDNA metabarcoding. However, eDNA metabarcoding detected roughly two thirds of metazoan species and identified similar taxa contributing to community differentiation across the subtropical front separating transects. We observed a diurnal pattern in eDNA data for copepods which perform diel vertical migrations, indicating a surprisingly short temporal eDNA signal. Compared to COI, a eukaryote-targeted 18S ribosomal RNA marker detected a higher proportion, but lower diversity, of metazoans in eDNA. With refinement and standardization of methodology, eDNA metabarcoding could become an efficient tool for monitoring open ocean biodiversity.

The fifth shade of green: A novel approach to phytoplankton color index assessment in an oligotrophic system

AbstractMeasurements of phytoplankton color index (PCI), an indicator of phytoplankton biomass, represent one of the largest datasets acquired through the continuous plankton recorder (CPR) survey. PCI has been used to describe changes in the spatial and temporal distribution of phytoplankton, especially in the North Atlantic, where the methodology was developed and primarily applied. The method functions by grouping samples, based on their color shade, into four categories. Each category is assigned a numerical value to aid spatial and temporal averaging. Recently a CPR has been operated in the Eastern Mediterranean, one of the most oligotrophic areas of the global oceans. To capture the specific characteristics of the area, an additional greenness grading is introduced. In this study, we use satellite‐derived chlorophyll a (Chl a) data to examine whether PCI categories can be distinguished within the low and narrow range of Chl a values found in the study area and to calculate the range of Chl a concentrations expressed by each category. Furthermore, a newly calculated numerical system for the general use of PCI in oligotrophic areas and previously calculated PCI numerical value systems utilizing a variety of in situ, laboratory and remote sensing experiments, were tested for their applicability in the area. Present results indicate the feasibility of employing PCI measurements to provide an estimate of Chl a concentration in the Eastern Mediterranean. This provides confidence in the implementation of a new category system for PCI, as well as the use of the new PCI numerical system for the assessment of phytoplankton in oligotrophic waters.

Spatial distributions and seasonality of four Calanus species in the Northeast Atlantic

This paper analyses spatial and seasonal patterns for near-surface abundances of four Calanus species in the Northeast Atlantic based on monthly Continuous Plankton Recorder (CPR) Survey data collected during 2008–2016. C. finmarchicus, C. helgolandicus, C. hyperboreus and C. glacialis represent different ecosystems, and water masses with differing oceanographic properties and origins. Hence, these are considered as indicator species in climate change studies. A southern transect between Norway and the United Kingdom represented the northern North Sea, a central transect crossed the Norwegian and Iceland Seas, while a northern transect between Norway and Svalbard crossed the entrance to the Barents Sea. C. finmarchicus was prevalent everywhere, while C. helgolandicus was mainly confined to the northern North Sea though also documented downstream in the Norwegian Sea as far north as the entrance to the Barents Sea. The ratio of C. helgolandicus to C. finmarchicus abundances is expected to increase in the northern North Sea and southern Norwegian Sea given continued increases in seawater temperature. C. hyperboreus and C. glacialis were mainly associated with regions influenced by Arctic waters in the Iceland and Norwegian Seas, and south of Svalbard, respectively. Within Atlantic water masses in the central Norwegian Sea, a distinct second generation of C. finmarchicus with surprisingly high late-autumn concentrations of older stages were found. In Coastal water, two or more generations of C. finmarchicus occurred, while only one generation was evident in Arctic waters. Young C. finmarchicus of the first annual generation showed an earlier timing in Coastal than Atlantic and Arctic waters. The upper range of the temperature niche for both C. hyperboreus and C. glacialis was found to be around 7 °C. However, the peak occurrence within the study area was in 1–2 °C in Arctic water in the Iceland Sea in April for C. hyperboreus, while around 5 °C in Barents Sea Arctic water in July for C. glacialis. C. finmarchicus was present in all water masses with a peak occurrence in June at around 6 °C. C. finmarchicus generally dominated the Calanus-biomass, but C. hyperboreus in Arctic water in the Iceland and Norwegian Seas in spring, and C. helgolandicus in the northern North Sea, could match or exceed the biomass of C. finmarchicus.

Zooplankton in the Southern Ocean from the continuous plankton recorder: Distributions and long-term change

We provide an analysis of zooplankton distributions in the circumpolar Southern Ocean based on samples collected by the international SCAR Southern Ocean Continuous Plankton Recorder Survey (SO-CPR). We analysed SO-CPR measurements in relation to satellite and oceanographic model hind-cast data over the period 1997–2018. These environmental data were chlorophyll-a concentration, net primary productivity (VGPM model), sea-surface temperature (SST), mixed layer depth, sea ice and the spatial gradient of SST (as an indicator of ocean fronts). Boosted Regression Tree (BRT) models were used to extrapolate in space and time measurements of the abundances of six key taxonomic groups of zooplankton: Copepoda (Calanoida), Euphausiidae (numerically dominated in SO-CPR data by Thysanoessa macrura), Foraminifera, Fritillaria spp., Oithona similis, and pteropods. Based on the BRT models, we present predictions of the spatial and seasonal (October–March) environmental suitability for these groups in the Southern Ocean. Trend analysis suggests that between 1997 and 2018 the environmental suitability for copepods (both cyclopoid and calanoid), Foraminifera, and Fritillaria spp. has increased by 0.72% per year average, and at higher rates in frontal regions especially in the Indian sector of the Southern Ocean. In contrast, for pteropods in some areas (particularly over the Ross Sea shelf) the environmental suitability has significantly worsened over the last 20 years.

Lifeform indicators reveal large-scale shifts in plankton across the North-West European shelf.

Increasing direct human pressures on the marine environment, coupled with climate-driven changes, is a concern to marine ecosystems globally. This requires the development and monitoring of ecosystem indicators for effective management and adaptation planning. Plankton lifeforms (broad functional groups) are sensitive indicators of marine environmental change and can provide a simplified view of plankton biodiversity, building an understanding of change in lower trophic levels. Here, we visualize regional-scale multi-decadal trends in six key plankton lifeforms as well as their correlative relationships with sea surface temperature (SST). For the first time, we collate trends across multiple disparate surveys, comparing the spatially and temporally extensive Continuous Plankton Recorder (CPR) survey (offshore) with multiple long-term fixed station-based time-series (inshore) from around the UK coastline. These analyses of plankton lifeforms showed profound long-term changes, which were coherent across large spatial scales. For example, 'diatom' and 'meroplankton' lifeforms showed strong alignment between surveys and coherent regional-scale trends, with the 1998-2017 decadal average abundance of meroplankton being 2.3 times that of 1958-1967 for CPR samples in the North Sea. This major, shelf-wide increase in meroplankton correlated with increasing SSTs, and contrasted with a general decrease in holoplankton (dominated by small copepods), indicating a changing balance of benthic and pelagic fauna. Likewise, inshore-offshore gradients in dinoflagellate trends, with contemporary increases inshore contrasting with multi-decadal decreases offshore (approx. 75% lower decadal mean abundance), urgently require the identification of causal mechanisms. Our lifeform approach allows the collation of many different data types and time-series across the NW European shelf, providing a crucial evidence base for informing ecosystem-based management, and the development of regional adaptation plans.

Phenological shuffling of major marine phytoplankton groups over the last six decades

AbstractAimPhytoplankton form the basis of the marine food web and are responsible for approximately 50% of the world's photosynthesis. Changes to their ecology are, therefore, important: here, we examined seasonal patterns in ocean phytoplankton abundance for 45 taxa over 59 years collected from circa 410,000 km of line‐transect sampling at temperate latitudes.LocationThe North Sea.MethodsFor our analysis we used plankton abundance data from the Continuous Plankton Recorder (CPR) survey, sea surface temperature measurements from the Hadley Centre, UK Meteorological Office and wind speed data from the International Comprehensive Ocean–Atmosphere Data Set, NOAA.ResultsWe found large differences in changes in the timing of peak abundance between the major phytoplankton groups. Late‐summer blooming dinoflagellates (n = 10 taxa) tended to show a large seasonal advancement, the timing of peak abundance for dinoflagellates as group advancing 39 days over these six decades. By contrast diatoms (n = 35) did not show any change as a group in their timing of peak abundance over the time series. Granger causality testing suggested a major driver of these phenological changes has been ocean warming in general but more specifically the rate of spring temperature rise as the most important factor. We also found differences in the timing of peak abundance of harmful algal bloom taxa, with some showing peak abundance earlier while others have moved later.Main conclusionsThere has been a fundamental transformation of the classic seasonal progression from blooms of diatoms to dinoflagellates, which lies at the heart of temperate marine food chains, as the classic bimodal diatom and dinoflagellate seasonal peaks are eroded to a more continuous, single, longer‐lasting phytoplankton peak. This phenological shuffling within and between major taxonomic groups is likely to have profound implications for the transfer of energy to higher trophic levels.

Defining isoscapes in the Northeast Pacific as an index of ocean productivity

AbstractAimWe modelled isoscapes in the Northeast (NE) Pacific using satellite‐based data, with the main objective of testing whether isoscapes defined by a few key parameters can be used as a proxy for secondary productivity.LocationNortheast Pacific; 46–60° N and 125–165° W.Time periodFrom 1998 to 2017 (ongoing).Major taxa studiedZooplankton, with a focus on large herbivores.MethodsApproximately 280 summer zooplankton samples were analysed for carbon (δ13C) and nitrogen (δ15N) stable isotope (SI) ratios. Environmental conditions experienced by zooplankton organisms were extracted from satellite, in situ sensor and model databases. A generalized additive model approach was used to explain the spatial variability of δ13C and δ15N values and to predict isoscapes.ResultsSea surface temperature (SST), sea level anomaly (SLA) and chlorophyll‐a concentration emerged as the significant SI predictors. Modelled isoscapes reproduced patterns observed in δ13C and δ15N value distribution, such as a decrease from the coast to offshore. The contribution of eddies in enhancing local production in the open ocean was also well captured by the models. In the central part of the NE Pacific, higher SI values were correlated with higher large copepod biomass measured by the North Pacific Continuous Plankton Recorder (CPR) survey. However, in the area off the coast of British Columbia, high δ15N variability appeared to be associated with episodic intrusions of coastal waters, demonstrating that caution is needed when interpreting sharp changes in SI ratios.Main conclusionsAlthough the mechanisms driving SI ratio variability are complex, we demonstrate that a few parameters used as a proxy for some of these major mechanisms are able to produce successful isoscape models. This approach was proved useful to provide a qualitative estimate of secondary production, which can be particularly valuable in a region where few data are available.