Decadal changes in climate and ecosystems in the North Atlantic Ocean and adjacent seas

Climate change is unambiguous and its effects are clearly detected in all functional units of the Earth system. This study presents new analyses of sea-surface temperature changes and show that climate change is affecting ecosystems of the North Atlantic. Changes are seen from phytoplankton to zooplankton to fish and are modifying the dominance of species and the structure, the diversity and the functioning of marine ecosystems. Changes also range from phenological to biogeographical shifts and have involved in some regions of the Atlantic abrupt ecosystem shifts. These alterations reflect a response of pelagic ecosystems to a warmer temperature regime. Mechanisms are complex because they are nonlinear exhibiting tipping points and varying in space and time. Sensitivity of organisms to temperature changes is high, implicating that a small temperature modification can have sustained ecosystem effects. Implications of these changes for biogeochemical cycles are discussed. Two observed changes detected in the North Sea that could have opposite effects on carbon cycle are discussed. Increase in phytoplankton, as inferred from the phytoplankton colour index derived from the Continuous Plankton Recorder (CPR) survey, has been detected in the North Sea. This pattern has been accompanied by a reduction in the abundance of the herbivorous species Calanus finmarchicus. This might have reduced the grazing pressure and increase diatomaceous ‘fluff’, therefore carbon export in the North Sea. Therefore, it could be argued that the biological carbon pump might increase in this region with sea warming. In the meantime, however, the mean size of organisms (calanoid copepods) has dropped. Such changes have implications for the turnover time of biogenic carbon in plankton organisms and the mean residence time of particulate carbon they produce. The system characterising the warmer period is more based on recycling and less on export. The increase in the minimum turnover time indicates an increase in the ecosystem metabolism, which can be considered as a response of the pelagic ecosystems to climate warming. This phenomenon could reduce carbon export. These two opposite patterns of change are examples of the diversity of mechanisms and pathways the ecosystems may exhibit with climate change. Oversimplification of current biogeochemical models, often due to lack of data and biological understanding, could lead to wrong projection on the direction ecosystems and therefore some biogeochemical cycles might take in a warmer world.

Mesoscale fronts as foraging habitats: composite front mapping reveals oceanographic drivers of habitat use for a pelagic seabird

The oceanographic drivers of marine vertebrate habitat use are poorly understood yet fundamental to our knowledge of marine ecosystem functioning. Here, we use composite front mapping and high-resolution GPS tracking to determine the significance of mesoscale oceanographic fronts as physical drivers of foraging habitat selection in northern gannets Morus bassanus. We tracked 66 breeding gannets from a Celtic Sea colony over 2 years and used residence time to identify area-restricted search (ARS) behaviour. Composite front maps identified thermal and chlorophyll-a mesoscale fronts at two different temporal scales—(i) contemporaneous fronts and (ii) seasonally persistent frontal zones. Using generalized additive models (GAMs), with generalized estimating equations (GEE-GAMs) to account for serial autocorrelation in tracking data, we found that gannets do not adjust their behaviour in response to contemporaneous fronts. However, ARS was more likely to occur within spatially predictable, seasonally persistent frontal zones (GAMs). Our results provide proof of concept that composite front mapping is a useful tool for studying the influence of oceanographic features on animal movements. Moreover, we highlight that frontal persistence is a crucial element of the formation of pelagic foraging hotspots for mobile marine vertebrates.

Distribution and abundance of sardine (Sardina pilchardus) eggs in the English Channel from Continuous Plankton Recorder sampling

Continuous Plankton Recorder (CPR) samples from the English Channel and adjacent Celtic shelf, taken over the period 1958-1980, were analysed for sardine (Sardina pilchardus) eggs. Results showed the progression of sardine spawning along the English Channel from west to east from March to August and a return from east to west from September to November. This corresponds with the two seasonal peaks of sardine egg abundance in the western Channel: the main summer peak being in May/June, with a smaller autumn peak in October/November. Long-term changes in sardine egg abundance in CPR samples showed a decline in summer spawning from the late 1960s, but no clear trend in autumn-spawned egg abundance. Similar patterns were observed in the numbers of sardine eggs sampled by conventional plankton net tows at the time-series Station L5 off Plymouth. This supports the use of the longer time-series of sardine egg data at L5 as being representative of a wider area and emphasizes the importance in continuation of the L5 time-series.

Continuous Plankton Records - Persistence In Time-Series And The Population-Dynamics Of Pseudocalanus-Elongatus And Acartia-Clausi

Rates of population increase in early spring and the sizes of overwintering stocks were calculated for the planktonic copepods Pseudocalanus elongatus and Acartia clausi for a set of areas covering the open waters of the north-east Atlantic Ocean and the North Sea for the period 1948 to 1979. For both species, the rates of population increase were higher in the open ocean than in the North Sea and appear to be related to temperature. The overwintering stocks in the North Sea were larger than those in the open ocean and are probably related to phytoplanton concentration. P. elongatus shows higher overwintering stocks and lower rates of population increase than A. clausi, resulting in different levels of persistence in the stocks of the two species. It is suggested that this difference in persistence is responsible for differences between the two species with respect to geographical distribution in summer and different patterns of year-to-year fluctuations in abundance.

The global distribution of phytoplankton size spectrum and size classes from their light-absorption spectra derived from satellite data

The absorption spectra of phytoplankton in the visible domain hold implicit information on the phytoplankton community structure. Here we use this information to retrieve quantitative information on phytoplankton size structure by developing a novel method to compute the exponent of an assumed power-law for their particle-size spectrum. This quantity, in combination with total chlorophyll-a concentration, can be used to estimate the fractional concentration of chlorophyll in any arbitrarily-defined size class of phytoplankton. We further define and derive expressions for two distinct measures of cell size of mixed. populations, namely, the average spherical diameter of a bio-optically equivalent homogeneous population of cells of equal size, and the average equivalent spherical diameter of a population of cells that follow a power-law particle-size distribution. The method relies on measurements of two quantities of a phytoplankton sample: the concentration of chlorophyll-a, which is an operational index of phytoplankton biomass, and the total absorption coefficient of phytoplankton in the red peak of visible spectrum at 676 nm. A sensitivity analysis confirms that the relative errors in the estimates of the exponent of particle size spectra are reasonably low. The exponents of phytoplankton size spectra, estimated for a large set of in situ data from a variety of oceanic environments (similar to 2400 samples), are within a reasonable range; and the estimated fractions of chlorophyll in pico-, nano- and micro-phytoplankton are generally consistent with those obtained by an independent, indirect method based on diagnostic pigments determined using high-performance liquid chromatography. The estimates of cell size for in situ samples dominated by different phytoplankton types (diatoms, prymnesiophytes, Prochlorococcus, other cyanobacteria and green algae) yield nominal sizes consistent with the taxonomic classification. To estimate the same quantities from satellite-derived ocean-colour data, we combine our method with algorithms for obtaining inherent optical properties from remote sensing. The spatial distribution of the size-spectrum exponent and the chlorophyll fractions of pico-, nano- and micro-phytoplankton estimated from satellite remote sensing are in agreement with the current understanding of the biogeography of phytoplankton functional types in the global oceans. This study contributes to our understanding of the distribution and time evolution of phytoplankton size structure in the global oceans.

Distribution and multi-annual abundance trends of the copepod Temora longicornis in the US Northeast shelf ecosystem

The average spatial distribution and annual abundance cycle are described for the copepod Temora longicornis from samples collected on broadscale surveys (1977-2006) and along continuous plankton recorder transects (1961-2006) of the US Northeast continental shelf ecosystem. After its annual low in winter, T. longicornis abundance begins to increase in coastal waters with the northern progression of spring conditions. Annual maximum shelf concentrations were found in the more southern inshore waters of the region during the summer months. Abundance throughout most of the ecosystem increased sharply in the early 1990s and remained high through 2001. During this period, the copepod became more numerous and widespread in offshore shelf waters. Abundance declined to approximately average levels in 2002 for the remainder of the time series, but its extended offshore range remained intact. Correlation analysis found that the copepods interannual abundance variability had a significant negative relationship with surface salinity anomalies throughout the ecosystem, with higher correlations found in the northernmost subareas. Temora longicornis abundance in the ecosystem's southernmost subarea (Middle Atlantic Bight) did not increase in the 1990s and was found to be negatively correlated to surface temperature, indicating that continued global warming could adversely impact the copepods annual abundance cycle in this region.

Comparisons of zooplankton time series

Evidence for climate-correlated low frequency variability of various components of marine ecosystems has accumulated rapidly over the past 2 decades. There has also been a growing recognition that society needs to learn how the fluctuations of these various components are linked, and to predict the likely amplitude and steepness of future changes. Demographic characteristics of marine zooplankton make them especially suitable for examining variability of marine ecosystems at interannual to decadal time scales. Their life cycle duration is short enough that there is little carryover of population membership from year to year, but long enough that variability can be tracked with monthly-to-seasonal sampling. Because zooplankton are rarely fished, comparative analysis of changes in their abundance can greatly enhance our ability to evaluate the importance of and interaction between physical environment, food web, and fishery harvest as causal mechanisms driving ecosystem level changes. A number of valuable within-region analyses of zooplankton time series have been published in the past decade, covering a variety of modes of variability including changes in total biomass, changes in size structure and species composition, changes in spatial distribution, and changes in seasonal timing. But because most zooplankton time series are relatively short compared to the time scales of interest, the statistical power of local analyses is often low, and between-region and between-variable comparisons are also needed. In this paper, we review the results of recent within- and between-region analyses, and suggest some priorities for future work.

Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. “Gut flux” for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.

Phenotypic variation in shell form in the intertidal acorn barnacle Chthamalus montagui: distribution, response to predators and life history trade-offs.

The acorn barnacle Chthamalus montagui can present strong variation in shell morphology, ranging from flat conic to a highly bent form, caused by a substantial overgrowth of the rostrum plate. Shell shape distribution was investigated between January and May 2004 from geographical to microhabitat spatial scales along the western coast of Britain. Populations studied in the north (Scotland and Isle of Man) showed a higher degree of shell variation compared to those in the south (Wales and south-west England). In the north, C. montagui living at lower tidal levels and in proximity to the predatory dogwhelk, Nucella lapillus, were more bent in profile. Laboratory experiments were conducted to examine behavioural responses, and vulnerability of bent and conic barnacles to predation by N. lapillus. Dogwhelks did not attack one morphotype more than the other, but only 15 % of attacks on bent forms were successful compared to 75 % in conic forms. Dogwhelk effluent reduced the time spent feeding by C. montagui (11 %), but there was no significant difference between conic and bent forms. Examination of barnacle morphology indicated a trade-off in investment in shell structure and feeding appendages associated with being bent, but none with egg or somatic tissue mass. These results are consistent with C. montagui showing an induced defence comparable to that found in its congeners Chthamalus anisopoma and Chthamalus fissus on the Pacific coast of North America, but further work to demonstrate inducibility is required.

How does Calanus helgolandicus maintain its population in a variable environment? Analysis of a 25-year time series from the English Channel

Calanus helgolandicus is a key copepod of the NE Atlantic and fringing shelves, with a distribution that is expanding northwards with oceanic warming. The Plymouth L4 site has warmed over the past 25-years, and experiences large variations in the timing and availability of food for C. helgolandicus. Here we examine the degree to which these changes translate into variation in reproductive output and subsequently C. helgolandicus population size. Egg production rates (eggs female−1 day−1) were maximal in the spring to early-summer period of diatom blooms and high ciliate abundance, rather than during the equally large autumn blooms of autotrophic dinoflagellates. Egg hatch success was lower in spring however, with a greater proportion of naupliar deformities then also. Both the timing and the mean summer abundance of C. helgolandicus (CI–CVI) reflected those of spring total reproductive output. However this relationship was driven by inter-annual variability in female abundance and not that of egg production per female, which ranged only two-fold. Winter abundance of C. helgolandicus at L4 was much more variable than abundance in other seasons, and reflected conditions from the previous growing season. However, these low winter abundances had no clear carry-over signal to the following season’s population size. Overall, the C. helgolandicus population appears to be surprisingly resilient at this dynamic, inshore site, showing no long-term phenology shift and only a four-fold variation in mean abundance between years. This dampening effect may reflect a series of mortality sources, associated with the timing of stratification in the early part of the season, likely affecting egg sinking and loss, plus intense, density-dependent mortality of early stages in mid-summer likely through predation.

The role of advection in the distribution of plankton populations at a moored 1-D coastal observatory

The degree to which advection modulates the distribution of plankton populations at a 1-D coastal observatory was assessed at station L4 in the western English Channel (50°15′N 4°13′W, depth 50 m), part of the Western Channel Observatory (WCO). Five tidal-cycle surveys were conducted, three in spring and two in summer 2010. Observations of the physical characteristics of L4 were obtained by using a moored acoustic doppler current profiler (ADCP) and a free-falling microstructure sensor (MSS). The moored ADCP highlighted the presence of vertical shear, with typical values of U during spring tides of ∼0.5 m s−1 at the surface and ∼0.2 m s−1 at the bed. The distribution of phyto- and zooplankton populations above a size threshold of 200 μm were examined using an in-line holographic imaging system, the Holocam. Variability in time as well as depth is a common feature throughout each of the surveys, with examples of recorded numbers of phytoplankton that ranged between 1300 L−1 and 2300 L−1 at the same depth but at different points within the tidal cycle. Further, at the same points in the tidal cycle the number of recorded zooplankton was also seen to vary, specifically with the identification of gelatinous planula in spring that increased the observed number to maximums of between 140 L−1 and 220 L−1 in the upper layer, considerably higher that the corresponding WP-2 net counts for a similar period. Specific aspects of the movement and transfer of plankton relating to advection and interaction with the pycnocline are identified, both across tidal cycles and seasons.