A regime shift in the North Sea circa 1988 linked to changes in the North Sea horse mackerel fishery

After 1987, Phytoplankton Colour (a visual estimate of chlorophyll) measured on samples taken by the continuous plankton recorder (CPR) in the North Sea increased substantially, both in level and seasonal extent, compared to earlier years since 1946. Many species of phytoplankton and zooplankton showed marked changes in abundance at about the same time. These events coincided with a large increase in catches of the western stock of the horse mackerel (Trachurus trachurus L.) in the northern North Sea reflecting a northerly expansion of the stock along the shelf edge from the Bay of Biscay to the North Sea after 1987. Using a 3D hydrodynamic model, with input from measured wind parameters, monthly transport of oceanic water into the North Sea has been calculated for the period 1976–1994, integrated for a section from Orkney to Shetland to Norway. A substantial increase in oceanic inflow occurred in the winter months, December to March, from 1988. Higher sea surface temperatures were also measured after 1987 especially in spring and summer months. These biological and physical events may be a response to observed changes in pressure distribution over the North Atlantic. From 1988 onwards, the North Atlantic Oscillation (NAO) index, the pressure difference between Iceland and the Azores, increased to the highest positive level observed in this century. Positive NAO anomalies are associated with stronger and more southerly tracks of the westerly winds and higher temperatures in western Europe. These changing wind distributions may have led to an increase in the northerly advection of water along the western edge of the European shelf and may have assisted the migration of the horse mackerel. This study is possibly a unique demonstration of a correlation between three different trophic levels of a marine ecosystem and hydrographic and atmospheric events at decadal and regional scales. The results emphasise the importance of maintaining into the future long term programmes such as the CPR.

Oceanographic responses to climate in the northwest Atlantic

Situated in an oceanographic transition zone, the Gulf of Maine/Western Scotian Shelf (GOM/WSS) region of the Northwest Atlantic is especially susceptible to changes in the climate system. Recent studies have shown that a coupled slope water system (CSWS) operates in the Northwest Atlantic and responds in a similar manner to climatic forcing over a broad range of time scales. These studies further suggest that it may be possible to associate different modes of the CSWS with the different phases of the North Atlantic Oscillation (NAO). Results from recent GLOBEC field studies in the Northwest Atlantic provide strong evidence linking physical responses of the CSWS to basin-scale forcing associated with the NAO. By placing these results in the context of time-series data collected from the GOM/WSS over the past half century, we show that we show that: (i) the region’s shelf ecosystems respond both physically and biologically to modal shifts in the CSWS; (ii) the CSWS mediates the effects on these ecosystems of basin-scale climatic forcing associated with the NAO and (iii) certain planktonic species can be good indicators of the CSWS’s modal state on inter-annual to interdecadal time scales.

Scale-dependent response diversity of seabirds to prey in the North Sea

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.

Synchronous response of marine plankton ecosystems to climate in the Northeast Atlantic and the North Sea

Over the last few decades, global warming has accelerated both the rate and magnitude of changes observed in many functional units of the Earth System. In this context, plankton are sentinel organisms because they are sensitive to subtle levels of changes in temperature and might help in identifying the current effects of climate change on pelagic ecosystems. In this paper, we performed a comparative approach in two regions of the North Atlantic (i.e. the Northeast Atlantic and the North Sea) to explore the relationships between changes in marine plankton, the regional physico-chemical environment and large-scale hydro-climatic forcing using four key indices: the North Atlantic Oscillation (NAO), the Atlantic Multidecadal Oscillation (AMO), the East Atlantic (EA) pattern and Northern Hemisphere Temperature (NHT) anomalies. Our analyses suggest that long-term changes in the states of the two ecosystems were synchronous and correlated to the same large-scale hydro-climatic variables: NHT anomalies, the AMO and to a lesser extent the EA pattern. No significant correlation was found between long-term ecosystem modifications and the state of the NAO. Our results suggest that the effect of climate on these ecosystems has mainly occurred in both regions through the modulation of the thermal regime.

Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales

Pronounced changes in fauna, extending from the English Channel in the south to the Barents Sea in the north-east and off Greenland in the north-west, have occurred in the late 1920s, the late 1960s and again in the late 1990s. We attribute these events to exchanges of subarctic and subtropical water masses in the north-eastern North Atlantic Ocean, associated with changes in the strength and extent of the subpolar gyre. These exchanges lead to variations in the influence exerted by the subarctic or Lusitanian biomes on the intermediate faunistic zone in the north-eastern Atlantic. This strong and persistent bottom-up bio-physical link is demonstrated using a numerical ocean general circulation model and data on four trophically connected levels in the food chain – phytoplankton, zooplankton, blue whiting, and pilot whales. The plankton data give a unique basin-scale depiction of these changes, and a long pilot whale record from the Faroe Islands offers an exceptional temporal perspective over three centuries. Recent advances in simulating the dynamics of the subpolar gyre suggests a potential for predicting the distribution of the main faunistic zones in the north-eastern Atlantic a few years into the future, which might facilitate a more rational management of the commercially important fisheries in this region.

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.

The CPR Survey (1948-1997): a gridded database browser of plankton abundance in the North Sea.

The Continuous Plankton Recorder (CPR) survey provides a unique multi- decadal dataset on the abundance of plankton in the North Sea and North Atlantic and is one of only a few monitoring programmes operating at a large spatio- temporal scale. The results of all samples analysed from the survey since 1946 are stored on an Access Database at the Sir Alister Hardy Foundation for Ocean Science (SAHFOS) in Plymouth. The database is large, containing more than two million records (~80 million data points, if zero results are added) for more than 450 taxonomic entities. An open data policy is operated by SAHFOS. However, the data are not on-line and so access by scientists and others wishing to use the results is not interactive. Requests for data are dealt with by the Database Manager. To facilitate access to the data from the North Sea, which is an area of high research interest, a selected set of data for key phytoplankton and zooplankton species has been processed in a form that makes them readily available on CD for research and other applications. A set of MATLAB tools has been developed to provide an interpolated spatio-temporal description of plankton sampled by the CPR in the North Sea, as well as easy and fast access to users in the form of a browser. Using geostatistical techniques, plankton abundance values have been interpolated on a regular grid covering the North Sea. The grid is established on centres of 1 degree longitude x 0.5 degree latitude (~32 x 30 nautical miles). Based on a monthly temporal resolution over a fifty-year period (1948-1997), 600 distribution maps have been produced for 54 zooplankton species, and 480 distribution maps for 57 phytoplankton species over the shorter period 1958-1997. The gridded database has been developed in a user-friendly form and incorporates, as a package on a CD, a set of options for visualisation and interpretation, including the facility to plot maps for selected species by month, year, groups of months or years, long-term means or as time series and contour plots. This study constitutes the first application of an easily accessed and interactive gridded database of plankton abundance in the North Sea. As a further development the MATLAB browser is being converted to a user- friendly Windows-compatible format (WinCPR) for release on CD and via the Web in 2003.

Arctic boreal plankton species in the Northwest Atlantic

The Continuous Plankton Recorder (CPR) survey has sampled regularly in the Northwest Atlantic since the early 1960s. Over the last decade there has been a dramatic increase in the abundance of a number of arctic boreal plankton species, notably Calanus hyperboreus (Kroyer), Calanus glacialis (Jaschnov), and Ceratium arcticum, and a southerly shift of the copepod C. hyperboreus in the CPR survey. In 1998, C. hyperboreus was recorded at its farthest position south in the survey, 39 degrees N, off the Georges Bank shelf edge. Other studies have reported similar parallel biological responses on three trophic levels. During the late 1990s, production of Labrador Sea Water (LSW) has been at a high, a direct response to the phase of the North Atlantic Oscillation (NAO). The increase in abundance of these species, up to four standard deviations from the long-term mean, is linked to variability in the hydrography of the area and the driving climatic processes of the North Atlantic.

Macroecology of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas

Global climate change is expected to modify the spatial distribution of marine organisms. However, projections of future changes should be based on robust information on the ecological niche of species. This paper presents a macroecological study of the environmental tolerance and ecological niche (sensu Hutchinson 1957, i.e. the field of tolerance of a species to the principal factors of its environment) of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas. Biological data were collected by the Continuous Plankton Recorder (CPR) Survey, which samples plankton in the North Atlantic and adjacent seas at a standard depth of 7 m. Eleven parameters were chosen including bathymetry, temperature, salinity, nutrients, mixed-layer depth and an index of turbulence compiled from wind data and chlorophyll a concentrations (used herein as an index of available food). The environmental window and the optimum level were determined for both species and for each abiotic factor and chlorophyll concentration. The most important parameters that influenced abundance and spatial distribution were temperature and its correlates such as oxygen and nutrients. Bathymetry and other water-column-related parameters also played an important role. The ecological niche of C. finmarchicus was larger than that of C. helgolandicus and both niches were significantly separated. Our results have important implications in the context of global climate change. As temperature (and to some extent stratification) is predicted to continue to rise in the North Atlantic sector, changes in the spatial distribution of these 2 Calanus species can be expected. Application of this approach to the 1980s North Sea regime shift provides evidence that changes in sea temperature alone could have triggered the substantial and rapid changes identified in the dynamic regimes of these ecosystems. C. finmarchicus appears to be a good indicator of the Atlantic Polar Biome (mainly the Atlantic Subarctic and Arctic provinces) while C. helgolandicus is an indicator of more temperate waters (Atlantic Westerly Winds Biome) in regions characterised by more pronounced spatial changes in bathymetry.

Particulate and dissolved aluminum and titanium in the upper water column of the Atlantic Ocean

This study investigates the oceanic behavior of the lithogenic trace elements Al and Ti in the upper 200 m of the Atlantic Ocean. The distribution of both metals in the dissolved and particulate phases was assessed along an E-W transect in the eastern tropical North Atlantic (December 2009) and along a meridional Atlantic transect (April-May 2010). The surface water concentrations of particulate and dissolved Al and Ti reflected the previously observed pattern of atmospheric inputs into the Atlantic Ocean. Subsurface minima at stations with pronounced fluorescence maxima were observed, suggesting a link between biological productivity and the removal of both dissolved and particulate Al and Ti. This may include uptake mechanisms, adsorption and aggregation processes on biogenic particle surfaces and the formation of large, fast sinking biogenic particles, e.g., fecal pellets. Residence times in the upper water column (100 m) of the tropical and subtropical North Atlantic were estimated to range in the order of days to weeks in the particulate phases (Al: 3-22 days, Ti: 4-37 days) and were 0.9-3.8 years for Al and 10-31 years for Ti in the dissolved phases. Longer residence times in both phases in the South Atlantic are consistent with lower biological productivity and decreased removal rates. In the upper water column, Al was predominantly present in the dissolved form, whereas Ti mostly occurred in the particulate form. Largest deviations in the partition coefficients between the particulate and dissolved phases were found in the surface waters, together with excess dissolved Al over Ti compared to the crustal source. This likely reflects elevated dissolution of Al compared to Ti from atmospheric mineral particles.

Physiological And Biochemical Aspects Of The Valve Snap And Valve Closure Responses In The Giant Scallop Placopecten-Magellanicus .1. Physiology

1. Catabolic processes of the phasic and catch parts of the adductor muscle ofPlacopecten magellanicus have been studied in relation to valve snap and valve closure responses. It is concluded that the snap response is powered by both parts of the adductor muscle and the valve closure response is powered exclusively by the catch part. 2. Both parts of the adductor muscle show a high glycolytic potential, reflected by high levels of glycolytic enzymes (Table 1) and high glycogen levels (Table 2). Lactate dehydrogenase could not be detected. In contrast, octopine dehydrogenase shows high activities in both parts of the adductor muscle. It is therefore concluded that a main anaerobic pathway in both tissues is the breakdown of glycogen to octopine. In the catch part, however, a considerable amount of the pyruvate formed from glycogen may also be converted into alanine (see below). The glycolytic flux in the catch part is much higher during the snap response than during valve closure. 3. The absence of phosphoenolpyruvate carboxykinase in the adductor muscle ofP. magellanicus and the observed changes in aspartate, alanine and succinate demonstrate that the energy metabolism in the catch part during valve closure shows great similarities to that which occurs only in the initial stage of anaerobiosis in the catch adductor muscle of the sea musselMytilus edulis L. 4. Arginine kinase activity and arginine phosphate content of the phasic part are much higher than those of the catch part (Tables 1 and 3). This may explain why in the phasic part during the snap response most ATP equivalents are derived from arginine phosphate, and in the catch part during both valve responses most are derived from glycolysis (Table 6). Despite the limited contribution of glycolysis in the phasic part during the snap response, the glycolytic flux increases by a factor of at least 75. 5. Evidence is obtained that octopine is neither transported from one part of the adductor muscle to the other, nor from the adductor muscle to other tissues.