Influence of climate change and trophic coupling across four trophic levels in the Celtic Sea

Climate change has had profound effects upon marine ecosystems, impacting across all trophic levels from plankton to apex predators. Determining the impacts of climate change on marine ecosystems requires understanding the direct effects on all trophic levels as well as indirect effects mediated by trophic coupling. The aim of this study was to investigate the effects of climate change on the pelagic food web in the Celtic Sea, a productive shelf region in the Northeast Atlantic. Using long-term data, we examined possible direct and indirect ‘bottom-up’ climate effects across four trophic levels: phytoplankton, zooplankton, mid-trophic level fish and seabirds. During the period 1986–2007, although there was no temporal trend in the North Atlantic Oscillation index (NAO), the decadal mean Sea Surface Temperature (SST) in the Celtic Sea increased by 0.66±0.02°C. Despite this, there was only a weak signal of climate change in the Celtic Sea food web. Changes in plankton community structure were found, however this was not related to SST or NAO. A negative relationship occurred between herring abundance (0- and 1-group) and spring SST (0-group: p = 0.02, slope = −0.305±0.125; 1-group: p = 0.04, slope = −0.410±0.193). Seabird demographics showed complex species–specific responses. There was evidence of direct effects of spring NAO (on black-legged kittiwake population growth rate: p = 0.03, slope = 0.0314±0.014) as well as indirect bottom-up effects of lagged spring SST (on razorbill breeding success: p = 0.01, slope = −0.144±0.05). Negative relationships between breeding success and population growth rate of razorbills and common guillemots may be explained by interactions between mid-trophic level fish. Our findings show that the impacts of climate change on the Celtic Sea ecosystem is not as marked as in nearby regions (e.g. the North Sea), emphasizing the need for more research at regional scales.

Microbial spatial variability: An example from the Celtic Sea

In July 2004, dominant populations of microbial ultraplankton (<5 μm), in the surface of the Celtic Sea (between UK and Eire), were repeatedly mapped using flow cytometry, at 1.5 km resolution over a region of diameter 100 km. The numerically dominant representatives of all basic functional types were enumerated including one group of phototrophic bacteria (Syn), two groups of phytoplankton (PP, NP), three groups of heterotrophic bacterioplankton (HB) and the regionally dominant group of heterotrophic protists (HP). The distributions of all organisms showed strong spatial variability with little relation to variability in physical fields such as salinity and temperature. Furthermore, there was little agreement between distributions of different organisms. The only linear correlation consistently explaining more than 50% of the variance between any pairing of the organism groups enumerated is between two different groups of HB. Specifically, no linear, or non-linear, relationship is found between any pairings of SYB, PP or HB groups with their protist predators HP. Looking for multiple dependencies, factor analysis reveals three groupings: Syn, PP and low nucleic acid content HB (LNA); high nucleic acid content HB (HNA); HP and NP. Even the manner in which the spatial variability of Syn, PP and HB abundance varies as a function of lengthscale (represented by a semivariogram) differs significantly from that for HP. In summary, although all microbial planktonic groups enumerated are present and numerically dominant throughout the region studied, at face value the relationships between them seem weak. Nevertheless, the behaviour of a simple, illustrative ecological model, with strongly interacting phototrophs and heterotrophs, with stochastic forcing, is shown to be consistent with the observed poor correlations and differences in how spatial variability varies with lengthscale. Thus, our study suggests that a comparison of microbial abundances alone may not discern strong underlying trophic interactions. Specific knowledge of these processes, in particular grazing, will be required to explain the causes of the observed microbial spatial variability and its resulting consequences for the functioning of the ecosystem.

Repeated, long-distance migrations by a philopatric predator targeting highly contrasting ecosystems

Long-distance movements of animals are an important driver of population spatial dynamics and determine the extent of overlap with area-focused human activities, such as fishing. Despite global concerns of declining shark populations, a major limitation in assessments of population trends or spatial management options is the lack of information on their long-term migratory behaviour. For a large marine predator, the tiger shark Galeocerdo cuvier, we show from individuals satellite-tracked for multiple years (up to 1101 days) that adult males undertake annually repeated, round-trip migrations of over 7,500 km in the northwest Atlantic. Notably, these migrations occurred between the highly disparate ecosystems of Caribbean coral reef regions in winter and high latitude oceanic areas in summer, with strong, repeated philopatry to specific overwintering insular habitat. Partial migration also occurred, with smaller, immature individuals displaying reduced migration propensity. Foraging may be a putative motivation for these oceanic migrations, with summer behaviour showing higher path tortuosity at the oceanic range extremes. The predictable migratory patterns and use of highly divergent ecosystems shown by male tiger sharks appear broadly similar to migrations seen in birds, reptiles and mammals, and highlight opportunities for dynamic spatial management and conservation measures of highly mobile sharks.

Repeated, long-distance migrations by a philopatric predator targeting highly contrasting ecosystems

Long-distance movements of animals are an important driver of population spatial dynamics and determine the extent of overlap with area-focused human activities, such as fishing. Despite global concerns of declining shark populations, a major limitation in assessments of population trends or spatial management options is the lack of information on their long-term migratory behaviour. For a large marine predator, the tiger shark Galeocerdo cuvier, we show from individuals satellite-tracked for multiple years (up to 1101 days) that adult males undertake annually repeated, round-trip migrations of over 7,500 km in the northwest Atlantic. Notably, these migrations occurred between the highly disparate ecosystems of Caribbean coral reef regions in winter and high latitude oceanic areas in summer, with strong, repeated philopatry to specific overwintering insular habitat. Partial migration also occurred, with smaller, immature individuals displaying reduced migration propensity. Foraging may be a putative motivation for these oceanic migrations, with summer behaviour showing higher path tortuosity at the oceanic range extremes. The predictable migratory patterns and use of highly divergent ecosystems shown by male tiger sharks appear broadly similar to migrations seen in birds, reptiles and mammals, and highlight opportunities for dynamic spatial management and conservation measures of highly mobile sharks.

Report on trawling and other investigations carried out in the Bays on the South-East Coast of Devon during 1901 and 1902

Section 1. The Brixham fishing-grounds and fishery statistics. Section 2. Distribution and migrations of food-fishes. Section 3. The reproduction of the flat-fishes. Appendix 1. Preliminary Report on the Trawling Experiments in the Bays on the South Coast of Devon.

Impact of climate change on marine pelagic phenology and trophic mismatch

Phenology, the study of annually recurring life cycle events such as the timing of migrations and flowering, can provide particularly sensitive indicators of climate change. Changes in phenology may be important to ecosystem function because the level of response to climate change may vary across functional groups and multiple trophic levels. The decoupling of phenological relationships will have important ramifications for trophic interactions, altering food-web structures and leading to eventual ecosystem-level changes. Temperate marine environments may be particularly vulnerable to these changes because the recruitment success of higher trophic levels is highly dependent on synchronization with pulsed planktonic production. Using long-term data of 66 plankton taxa during the period from 1958 to 2002, we investigated whether climate warming signals are emergent across all trophic levels and functional groups within an ecological community. Here we show that not only is the marine pelagic community responding to climate changes, but also that the level of response differs throughout the community and the seasonal cycle, leading to a mismatch between trophic levels and functional groups.

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.

An individual-based model of the early life history of mackerel (Scomber scombrus) in the eastern North Atlantic, simulating transport, growth and mortality

The main purpose of this paper is to provide the core description of the modelling exercise within the Shelf Edge Advection Mortality And Recruitment (SEAMAR) programme. An individual-based model (IBM) was developed for the prediction of year-to-year survival of the early life-history stages of mackerel (Scomber scombrus) in the eastern North Atlantic. The IBM is one of two components of the model system. The first component is a circulation model to provide physical input data for the IBM. The circulation model is a geographical variant of the HAMburg Shelf Ocean Model (HAMSOM). The second component is the IBM, which is an i-space configuration model in which large numbers of individuals are followed as discrete entities to simulate the transport, growth and mortality of mackerel eggs, larvae and post-larvae. Larval and post-larval growth is modelled as a function of length, temperature and food distribution; mortality is modelled as a function of length and absolute growth rate. Each particle is considered as a super-individual representing 10 super(6) eggs at the outset of the simulation, and then declining according to the mortality function. Simulations were carried out for the years 1998-2000. Results showed concentrations of particles at Porcupine Bank and the adjacent Irish shelf, along the Celtic Sea shelf-edge, and in the southern Bay of Biscay. High survival was observed only at Porcupine and the adjacent shelf areas, and, more patchily, around the coastal margin of Biscay. The low survival along the shelf-edge of the Celtic Sea was due to the consistently low estimates of food availability in that area.

Exceptional abundance of the snake pipefish (Entelurus aequoreus) in the north-eastern Atlantic Ocean

Pipefish (Syngnathidae) have occurred with unprecedented frequency in Continuous Plankton Recorder (CPR) samples to the west of the British Isles from 2003 to 2005. Identification by mtDNA sequencing established that they were snake pipefish, Entelurus aequoreus. The geographical range of the records were from the outer continental shelf of the Celtic Sea and north-west of Ireland to the mid-Atlantic Ridge between 40° and 57°N, with the greatest abundance near the shelf edge and adjacent oceanic waters south of Ireland and west of Brittany. There were records in every month from February to November but most were in late spring and summer. A proposed mechanism for the increase in abundance of the species is that recent climate change has had beneficial impacts on the reproduction of adults and the survival of larvae and juveniles.

Variations in the distributions of the Centropages chierchiae and Temora stylifera (Copepoda: Calanoida) in the north-eastern Atlantic Ocean and western european shelf waters.

Centropages chierchiae and Temora stylifera occurred rarely in the Continuous Plankton Recorder (CPR) survey in the Bay of Biscay, Celtic Sea, and English Channel before 1988. By 2000 they were found frequently and in abundance. The seasonal cycles of abundance of these species differ, C. chierchiae occurring mainly in the summer while T. stylifera was found most frequently in late autumn or winter towards the northern limits of its distribution. The increase in abundance of both species is related to temperature. However, in the years when it was found in the samples, the frequency of occurrence of C. chierchiae was correlated positively with the strength of the shelf edge current and negatively with the North Atlantic Oscillation (NAO) while the reverse was true for T. stylifera.

An overview of the literature concerning the oceanography of the eastern North Atlantic region

An overview of the main oceanographic features of the eastern North Atlantic boundary, with emphasis toward the upper layers, is presented. The principal features discussed are: water mass boundaries; forcing by wind, density and tides; topographic features and effects; fronts; upwelling and downwelling; poleward flows; coastal currents; eddies. The occurrence and spatial and seasonal variability of these features is described in five regional sections: Celtic Sea and western English Channel; Bay of Biscay; western Iberia; Gulf of Cadiz; northwest Africa. This paper is intended to provide a base of physical oceanographic knowledge in support of research in fisheries, biological and chemical oceanography, and marine biology.

Relating plankton assemblages to environmental variables using instruments towed by ships-of-opportunity

Undulating Oceanographic Recorders (UORs) and Continuous Plankton Recorders (CPRs) equipped with a suite of sensors were towed by merchant vessels in the North Sea between 1988 and 1991, recording a range of environmental variables. These were used to interpret the results of analyses of the plankton taken on CPR tows off the northeast coast of the UK in 1989 and in the Skagerrak and Kattegat in July 1988 and through 1989. Correlations were found between the biota and the environmental variables. The tidal front off the northeast coast of the UK and the front between the low salinity water in the Kattegat and the higher salinity water in the Skagerrak were dominant factors correlating with the distribution of the plankton assemblages. Discontinuities, defining the positions of the fronts, in the values of physical variables (temperature and, where measured, salinity and turbidity) were closely identified with geographical divisions between plankton assemblages. Measures of irradiance were found to be important on several occasions, presumably due to diel migrations of the zooplankton.

Effects Of Preservation On Wet Weight Dry-Weight Nitrogen And Carbon Contents Of Calanus-Helgolandicus (Crustacea Copepoda)

The lengths, wet and dry weights, nitrogen and carbon contents of fresh, frozen and formaldehyde-fixed specimens of Calanus helgolandicus (Claus) were determined. Samples were collected during May 1980 in the Celtic Sea. Individual Copepodite Stages 3, 4, 5, and Adult Male and Female Stage 6 were measured and analysed, and 36 linear regression equations derived for these variables together with mean values, standard deviations and 95% confidence limits. The range of nitrogen values in the fresh material, expressed as a percentage of dry weight, ranged from 8.08%±0.80 (Copepodite Stage 3) to 10.89%±0.27 (adult female); carbon values changed from 41.6%±3.05 (mean ±95% confidence limits) for Copepodite Stage 3 to 50.97%±2.63 in Copepodite Stage 5. The adult females had a high nitrogen and relatively low carbon content, while the converse was true for Stage 5 copepodites. There was a loss of dry weight from the frozen samples (57%) and the fixed samples (38%) compared with the mean of the fresh dry weight of all stages. The material lost from the copepods was rich in nitrogen, thus, artificially high percentage carbon values were determined from the frozen and fixed samples (52.0 to 60.3% and 44.7 to 58.5%, respectively).

Zooplankton Of The Bristol Channel And Severn Estuary - The Distribution Of 4 Copepods In Relation To Salinity

The seasonal variations in distribution and abundance of the common zooplankton species in the Bristol Channel and Severn Estuary were related to the salinity regimes observed over the period November 1973 to February 1975. The dominant constituents in all regions were the calanoid copepods, which reached maximum densities in July: approximately 100 times their winter levels. Four zooplankton assemblages were recognised using an objective classification program which computed similarity coefficients and used group-average sorting. The assemblages existed along the salinity gradient observed from the Severn Estuary to the Celtic Sea. The assemblages were classified as true estuarine, estuarine and marine, euryhaline marine and stenohaline marine and were characterized by the copepods Eurytemora affinis (Poppe) (<30‰S), Acartia bifilosa var. inermis (rose) (27 to 33.5‰S), Centropages hamatus (Lilljeborg) (31 to 35‰S) and Calanus helgolandicus (Claus) (>33‰S), respectively.

Oceanic controls on the primary production of the northwest European continental shelf: model experiments under recent past conditions and a potential future scenario

In this paper we clearly demonstrate that changes in oceanic nutrients are a first order factor in determining changes in the primary production of the northwest European continental shelf on time scales of 5–10 yr. We present a series of coupled hydrodynamic ecosystem modelling simulations, using the POLCOMS-ERSEM system. These are forced by both reanalysis data and a single example of a coupled ocean-atmosphere general circulation model (OA-GCM) representative of possible conditions in 2080–2100 under an SRES A1B emissions scenario, along with the corresponding present day control. The OA-GCM forced simulations show a substantial reduction in surface nutrients in the open-ocean regions of the model domain, comparing future and present day time-slices. This arises from a large increase in oceanic stratification. Tracer transport experiments identify a substantial fraction of on-shelf water originates from the open-ocean region to the south of the domain, where this increase is largest, and indeed the on-shelf nutrient and primary production are reduced as this water is transported on-shelf. This relationship is confirmed quantitatively by comparing changes in winter nitrate with total annual nitrate uptake. The reduction in primary production by the reduced nutrient transport is mitigated by on-shelf processes relating to temperature, stratification (length of growing season) and recycling. Regions less exposed to ocean-shelf exchange in this model (Celtic Sea, Irish Sea, English Channel, and Southern North Sea) show a modest increase in primary production (of 5–10%) compared with a decrease of 0–20% in the outer shelf, Central and Northern North Sea. These findings are backed up by a boundary condition perturbation experiment and a simple mixing model.