Long-term effects of ocean warming on the prokaryotic community: evidence from the vibrios

The long-term effects of ocean warming on prokaryotic communities are unknown because of lack of historical data. We overcame this gap by applying a retrospective molecular analysis to the bacterial community on formalin-fixed samples from the historical Continuous Plankton Recorder archive, which is one of the longest and most geographically extensive collections of marine biological samples in the world. We showed that during the last half century, ubiquitous marine bacteria of the Vibrio genus, including Vibrio cholerae, increased in dominance within the plankton-associated bacterial community of the North Sea, where an unprecedented increase in bathing infections related to these bacteria was recently reported. Among environmental variables, increased sea surface temperature explained 45% of the variance in Vibrio data, supporting the view that ocean warming is favouring the spread of vibrios and may be the cause of the globally increasing trend in their associated diseases.

Prey landscapes help identify potential foraging habitats for leatherback turtles in the NE Atlantic

Identifying key marine megavertebrate habitats has become ever more important as concern increases regarding global fisheries bycatch and accelerated climate change. This will be aided by a greater understanding of the patterns and processes determining the spatiotemporal distribution of species of conservation concern. We identify probable foraging grounds for leatherback turtles in the NE Atlantic using monthly landscapes of gelatinous organism distribution constructed from Continuous Plankton Recorder Survey data. Using sightings data (n = 2013 records, 1954 to 2003) from 9 countries (UK, Ireland, France, Belgium, The Netherlands, Denmark, Germany, Norway and Sweden), we show sea surface temperatures of approximately 10 to 12 degree C most likely indicate the lower thermal threshold for accessible habitats during seasonal foraging migrations to high latitudes. Integrating maps of gelatinous plankton as a possible indicator of prey distribution with thermal tolerance parameters demonstrates the dynamic (spatial and temporal) nature of NE Atlantic foraging habitats. We highlight the importance of body size- related thermal constraints in structuring leatherback foraging populations and demonstrate a latitudinal gradient in body size (Bergmann's rule) where smaller animals are excluded from higher latitude foraging areas. We highlight the marine area of the European continental shelf edge as being both thermally accessible and prey rich, and therefore potentially supporting appreciable densities of foraging leatherbacks, with some suitable areas not yet extensively surveyed.

Modelling the future biogeography of North Atlantic zooplankton communities in response to climate change

Advances in habitat and climate modelling allow us to reduce uncertainties of climate change impacts on species distribution. We evaluated the impacts of future climate change on community structure, diversity, distribution and phenology of 14 copepod species in the North Atlantic. We developed and validated habitat models for key zooplankton species using continuous plankton recorder (CPR) survey data collected at mid latitudes of the North Atlantic. Generalized additive models (GAMs) were applied to relate the occurrence of species to environmental variables. Models were projected to future (2080–2099) environmental conditions using coupled hydroclimatix–biogeochemical models under the Intergovernmental Panel on Climate Change (IPCC) A1B climate scenario, and compared to present (2001–2020) conditions. Our projections indicated that the copepod community is expected to respond substantially to climate change: a mean poleward latitudinal shift of 8.7 km per decade for the overall community with an important species range variation (–15 to 18 km per decade); the species seasonal peak is expected to occur 12–13 d earlier for Calanus finmarchicus and C. hyperboreus; and important changes in community structure are also expected (high species turnover of 43–79% south of the Oceanic Polar Front). The impacts of the change expected by the end of the century under IPCC global warming scenarios on copepods highlight poleward shifts, earlier seasonal peak and changes in biodiversity spatial patterns that might lead to alterations of the future North Atlantic pelagic ecosystem. Our model and projections are supported by a temporal validation undertaken using the North Atlantic climate regime shift that occurred in the 1980s: the habitat model built in the cold period (1970–1986) has been validated in the warm period (1987–2004).

High dispersal potential has maintained long-term population stability in the North Atlantic copepod Calanus finmarchicus

The cool-water copepod Calanus finmarchicus is a key species in North Atlantic marine ecosystems since it represents an important food resource for the developmental stages of several fish of major economic value. Over the last 40 years, however, data from the Continuous Plankton Recorder survey have highlighted a 70 per cent reduction in C. finmarchicus biomass, coupled with a gradual northward shift in the species's distribution, which have both been linked with climate change. To determine the potential for C. finmarchicus to track changes in habitat availability and maintain stable effective population sizes, we have assessed levels of gene flow and dispersal in current populations, as well as using a coalescent approach together with palaeodistribution modelling to elucidate the historical population demography of the species over previous changes in Earth's climate. Our findings indicate high levels of dispersal and a constant effective population size over the period 359 000-566 000 BP and suggest that C. finmarchicus possesses the capacity to track changes in available habitat, a feature that may be of crucial importance to the species's ability to cope with the current period of global climate change.

Long-term changes in abundance and distribution of microzooplankton in the NE Atlantic and North Sea

Long-term changes in mesozooplankton and phytoplankton populations have been well documented in the North Atlantic region, whereas data for microzooplankton are scarce. This neglected component of the plankton is a vital link in marine food-webs, grazing on smaller flagellates and cyanobacteria and in turn providing food for the larger mesozooplankton. We use the latest tintinnid (Ciliophora, Protista) data from the Continuous Plankton Recorder (CPR) survey in the NE Atlantic and North Sea to examine the phenology, distribution and abundance of this important group of ciliates. Presence/absence data came from 167 122 CPR samples collected between 1960 and 2009 and abundance data from 49 662 samples collected between 1996 and 2009. In the North Atlantic the genus Dictyocysta spp. dominated and Parafavella gigantea showed an increase in abundance around Iceland and Greenland. In the North Sea higher densities of Tintinnopsis spp., Favella serrata and Ptychocylis spp. were found. The presence of tintinnids in CPR samples collected in the North Atlantic has increased over the last 50 years and the seasonal window of high abundance has lengthened. Conversely in the North Sea there has been an overall reduction in abundance. We discuss possible drivers for these long-term changes and point the way forward to more holistic studies that examine how ecosystems, rather than just selected taxa, are responding to climate change.

Long-term changes in the diel vertical migration behaviour of zooplankton

Samples historically collected and analysed by the Continuous Plankton Recorder survey were used to examine long-term (1958 to 1994) patterns in the normal diel vertical migration (NDVM) behaviour of 7 copepod taxa in the North Sea: Calanus finmarchicus C5-C6; Calanus spp. C1-C4; Centropages typicus; Centropages hamatus; Temora longicornis; Acartia clausii and Para-Pseudocalanus (this last group included all Paracalanus and Pseudocalanus species). The ratio of night:day abundance near the surface was used as a measure of the extent of NDVM. For all 7 taxa, the extent of NDVM between 1958 and 1994 co-varied with the abundance of herring Clupea harengus in the North Sea. Fisheries data show that during this period the herring stock was a good indicator of the overall abundance of planktivorous fish in the North Sea. These results suggest that changes in the abundance of planktivorous fish in the North Sea over recent decades have resulted in modifications in the NDVM behaviour of many zooplankton taxa.

Ontogenetic and seasonal variation in the Diel Vertical Migration of the Copepods Metridia lucens and Metridia longa

Plankton samples collected and analyzed by the Continuous Plankton Recorder survey were used to examine the length of time spent near the surface in the North Atlantic and the North Sea by three closely related groups of zooplankton (copepodite stages 1-4 Metridia spp., copepodite stages 5-6 Metridia lucens, and copepodite stages 5-6 Metridia longa). For all three groups, the mean daily length of time spent near the surface in each month of the year covaried seasonally with day length. In addition, the amount of time spent near the surface varied significantly between the three groups, being longest for the copepods of smallest body size (C 1-C4 Metridia spp.) and shortest for the copepods of largest body size (C5-C6 M. longa). These results support the suggestion that diel vertical migration serves to reduce the risk of mortality from visually orienting predators

Diel vertical migration behaviour of Calanus hyperboreus at temperate latitudes

Samples historically collected and analysed by the Continuous Plankton Recorder survey in the North Atlantic and adjacent seas were used to examine temporal patterns in the near-surface abundance of copepodite stages V and VI Calanus hyperboreus (n = a total of 32836 samples; mean latitude of samples = 54* N, SD = 6.7*). The occurrence of CV and VI C. hyperboreus was largely restricted to samples collected in April, May and June. During this period there was a diel cycle in near-surface abundance which was consistent with a normal pattern of diel vertical migration. For example, CV-VI C. hyperboreus were 1.9 times more abundant at night than during the day, a difference which was highly significant (t-test, t5072 = 3.9, p < 0.001). The ratio of night:day abundance was similar in CV-VI C. finmarchicus (night:day abundance = 1.4), but markedly higher in CV-VI Metridia lucens (night:day abundance = 12.3) and CV-VI M. longa (night:day abundance = 15.0).

Have jellyfish in the Irish Sea benefited from climate change and overfishing?

Climate change and overfishing may lead to ecosystem instability and may benefit nonexploited organisms such as jellyfish. In the Irish Sea, an increase in jellyfish abundance was evident (r2=0.29, P=0.03) in a 16-year time-series (1994–2009) collected during juvenile fish surveys. Jellyfish abundance correlated positively with sea surface temperature (SST) over the preceding 18 months (r=0.65, pACF<0.001) and copepod biomass in the previous year (r=0.56, pACF=0.03) and negatively with spring (February–May) precipitation (r=−0.57, pACF=0.02). Principal components regression indicated that climatic indices explained 68% of the interannual variability in jellyfish abundance (P=0.003), where the components were based on the North Atlantic Oscillation Index, SST and precipitation. The frequency of cnidarian material present in Continuous Plankton Recorder (CPR) samples has also increased since 1970, with a period of frequent outbreaks between 1982 and 1991. Before this period, the herring stock in the northern Irish Sea declined rapidly to a low level, potentially stimulating structural change in the ecosystem. In 1985, there was a step decrease in CPR copepod biomass and in 1989, a step increase in the phytoplankton colour index, suggesting a cascading regime shift during the 1980s. Subsequent overexploitation of gadids, coupled with warm temperatures and the poor recruitment of cod, led to the rapid decline in cod biomass from 1990. While the biomass of sprat has decreased in the last decade, the herring stock has recovered partially. Reductions in demersal fishing pressure since 2000, intended to stimulate cod recovery, appear to have facilitated further rises in haddock biomass. Since the 1980s regime shift, sea temperatures have increased, the fish community has altered and jellyfish abundance has risen such that jellyfish and haddock may now play an increasingly important role in the ecosystem.

Changes in marine dinoflagellate and diatom abundance under climate change

Marine diatoms and dinoflagellates play a variety of key ecosystem roles as important primary producers (diatoms and some dinoflagellates) and grazers (some dinoflagellates). Additionally some are harmful algal bloom (HAB) species and there is widespread concern that HAB species may be increasing accompanied by major negative socio-economic impacts, including threats to human health and marine harvesting1, 2. Using 92,263 samples from the Continuous Plankton Recorder survey, we generated a 50-year (1960–2009) time series of diatom and dinoflagellate occurrence in the northeast Atlantic and North Sea. Dinoflagellates, including both HAB taxa (for example, Prorocentrum spp.) and non-HAB taxa (for example, Ceratium furca), have declined in abundance, particularly since 2006. In contrast, diatom abundance has not shown this decline with some common diatoms, including both HAB (for example, Pseudo-nitzschia spp.) and non-HAB (for example, Thalassiosira spp.) taxa, increasing in abundance. Overall these changes have led to a marked increase in the relative abundance of diatoms versus dinoflagellates. Our analyses, including Granger tests to identify criteria of causality, indicate that this switch is driven by an interaction effect of both increasing sea surface temperatures combined with increasingly windy conditions in summer.

Data collection of depth and time integrated Calanus finmarchicus abundances, averaged temperatures, and integrated Chlorophyll concentration in the North Atlantic Ocean

Here we present a new, pan-North-Atlantic compilation of data on key mesozooplankton species, including the most important copepod, Calanus finmarchicus. Distributional data of eight representative zooplankton taxa, from recent (2000-2009) Continuous Plankton Recorder data, are presented, along with basin-scale data of the phytoplankton colour index. Then we present a compilation of data on C. finmarchicus, including observations of abundance, demography, egg production and female size, with accompanying data on temperature and chlorophyll. . This is a contribution by Canadian, European and US scientists and their institutions.

Biogeography of jellyfish in the North Atlantic

In recent years a global increase in jellyfish (i.e. Cnidarians and Ctenophores) abundance and a rise in the recurrence of jellyfish outbreak events have been largely debated, but a general consensus on this matter has not been achieved yet. Within this debate, it has been generally recognised that there is a lack of reliable data that could be analysed and compared to clarify whether indeed jellyfish are increasing throughout the world ocean as a consequence of anthropogenic impact and hydroclimatic variability. Here we describe different jellyfish data sets produced within the EU program EUROBASIN, which have been assembled with the aim of presenting an up to date overview on the diversity and standing stocks of North Atlantic jellyfish. Abundance and species composition were determined in samples collected in the epipelagic layer (0- 200m), using a net well adapted to quantitatively catching gelatinous zooplankton. The samples were collected in spring-summer (April-August) 2010-2013, in inshore and offshore North Atlantic waters, between 59-68LatN and 62W-5ELong. Jellyfish were also identified and counted in samples opportunistically collected by other sampling gears in the same region and in two coastal stations in the Bay of Biscay and in the Gulf of Cadiz. Continuous Plankton Recorder (CPR) samples collected in 2009-2012 were re-analysed with the aim of identifying the time and location of jellyfish blooms across the North Atlantic basin.