Long-Term Oceanographic and Ecological Research in the Western English Channel

Long-term research in the western English Channel, undertaken by the marine laboratories in Plymouth, is described and details of survey methods, sites, and time series given in this chapter. Major findings are summarized and their limitations outlined. Current research, with recent reestablishment and expansion of many sampling programmes, is presented, and possible future approaches are indicated. These unique long-term data sets provide an environmental baseline for predicting complex ecological responses to local, regional, and global environmental change. Between 1888 and the present, investigations have been carried out into the physical, chemical, and biological components (ranging from plankton and fish to benthic and intertidal assemblages) of the western English Channel ecosystem. The Marine Biological Association of the United Kingdom has performed the main body of these observations. More recent contributions come from the Continuous Plankton Recorder Survey, now the Sir Alister Hardy Foundation for Ocean Science, dating from 1957; the Institute for Marine Environmental Research, from 1974 to 1987; and the Plymouth Marine Laboratory, which was formed by amalgamation of the Institute for Marine Environmental Research and part of the Marine Biological Association, from 1988. Together, these contributions constitute a unique data series; one of the longest and most comprehensive samplings of environmental and marine biological variables in the world. Since the termination of many of these time series in 1987-1988 during a reorganisation of UK marine research, there has been a resurgence of interest in long-term environmental change. Many programmes have been restarted and expanded with support from several agencies. The observations span significant periods of warming (1921-1961; 1985-present) and cooling (1962-1980). During these periods of change, the abundance of key species underwent dramatic shifts. The first period of warming saw changes in zooplankton, pelagic fish, and larval fish, including the collapse of an important herring fishery. During later periods of change, shifts in species abundances have been reflected in other assemblages, such as the intertidal zone and the benthic fauna. Many of these changes appear to be related to climate, manifested as temperature changes, acting directly or indirectly. The hypothesis that climate is a forcing factor is widely supported today and has been reinforced by recent studies that show responses of marine organisms to climatic attributes such as the strength of the North Atlantic Oscillation. The long-term data also yield important insights into the effects of anthropogenic disturbances such as fisheries exploitation and pollution. Comparison of demersal fish hauls over time highlights fisheries effects not only on commercially important species but also on the entire demersal community. The effects of acute ("Torrey Canyon" oil spill) and chronic (tributyltin [TBT] antifoulants) pollution are clearly seen in the intertidal records. Significant advances in diverse scientific disciplines have been generated from research undertaken alongside the long-term data series.

Marine biodiversity and climate change: assessing and predicting the influence of climatic change using intertidal rocky shore biota. Occasional Publication of the Marine Biological Association 20

In the last 60 years climate change has altered the distribution and abundance of many seashore species. Below is a summary of the findings of this project. The MarClim project was a four year multi-partner funded project created to investigate the effects of climatic warming on marine biodiversity. In particular the project aimed to use intertidal species, whose abundances had been shown to fluctuate with changes in climatic conditions, as indicator species of likely responses of species not only on rocky shores, but also those found offshore. The project used historic time series data, from in some cases the 1950s onwards, and contemporary data collected as part of the MarClim project (2001-2005), to provide evidence of changes in the abundance, range and population structure of intertidal species and relate these changes to recent rapid climatic warming. In particular quantitative counts of barnacles, limpets and trochids were made as well as semi-quantitative surveys of up to 56 intertidal taxa.Historic and contemporary data informed experiments to understand the mechanisms behind these changes and models to predict future species ranges and abundances.

Creating a modern database from archive material - The Norman Holme archive: Interpretation of marine field research notebooks in the historical context of the MBA, 1958-83

This document provides details of the transfer of the Norman Holme archive data held in the National Marine Biological Library onto a modern database, specifically Marine Recorder. A key part in the creation of the database was the retrieval of a large amount of information recorded in field notebooks and on loosely-bound sheets of paper. As this work involved amending, interpreting and updating the available information, it was felt that an accurate record of this process should exist to allow scientists of the future to be able to clearly link the modern database to the archive material. This document also provides details of external information sources that were used to enhance and qualify the historical interpretation, such as estimating volumes and species abundances.

Predicted levels of future ocean acidification and temperature rise could alter community structure and biodiversity in marine benthic communities

A mesocosm experiment was conducted to quantify the effects of reduced pH and elevated temperature on an intact marine invertebrate community. Standardised faunal communities, collected from the extreme low intertidal zone using artificial substrate units, were exposed to one of eight nominal treatments (four pH levels: 8.0, 7.7, 7.3 and 6.7, crossed with two temperature levels: 12 and 16°C). After 60 days exposure communities showed significant changes in structure and lower diversity in response to reduced pH. The response to temperature was more complex. At higher pH levels (8.0 and 7.7) elevated temperature treatments contained higher species abundances and diversity than the lower temperature treatments. In contrast, at lower pH levels (7.3 and 6.7), elevated temperature treatments had lower species abundances and diversity than lower temperature treatments. The species losses responsible for these changes in community structure and diversity were not randomly distributed across the different phyla examined. Molluscs showed the greatest reduction in abundance and diversity in response to low pH and elevated temperature, whilst annelid abundance and diversity was mostly unaffected by low pH and was higher at the elevated temperature. The arthropod response was between these two extremes with moderately reduced abundance and diversity at low pH and elevated temperature. Nematode abundance increased in response to low pH and elevated temperature, probably due to the reduction of ecological constraints, such as predation and competition, caused by a decrease in macrofaunal abundance. This community-based mesocosm study supports previous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical groups. This study also illustrates the importance of considering indirect effects that occur within multispecies assemblages when attempting to predict the consequences of ocean acidification and global warming on marine communities.

Marine microplankton diversity database

Many macroecological theories have been developed to study the diversity on our planet. All these theories require the existence of consistent databases to test their predictions. In this work, we compiled a data set of marine microplankton species abundances at 788 stations with an extensive geographical coverage. Data were collected on different oceanographic cruises between 1992 and 2002. This database consists of abundances (cells/mL) for each species at each station and depth, together with estimates of the biomass and biovolume for each species. One of the key strengths in this database is that species identifications were made by the same taxonomist, which provides greater strength to the collection and ensures that estimates of species diversity are reliable. Environmental information has also been compiled at each station (chlorophyll, temperature, photosynthetically active radiation [PAR], nutrients) in order to have a characterization of the study area and to be used in studies on the environmental and biological controls of marine biodiversity.

Radiolaria: Major exporters of organic carbon to the deep ocean

Very large pulses of particulate organic matter intermittently sink to the deep waters of the open ocean in the Northeast Atlantic. These pulses, measured by moored sediment traps since 1989, can contribute up to 60% of the organic flux to 3000 m in a particular year and are thus a major cause of the variability in carbon sequestration from the atmosphere in the region. Pulses occur in the late summer and are characterized by material that is very rich in organic carbon but with low concentrations of the biominerals opal and calcite. A number of independent lines of evidence have been examined to determine the causes of these pulses: (1) Data from the Continuous Plankton Recorder (CPR) survey show that in this region, radiolarian protozoans intermittently reach high abundances in the late summer just preceding organic pulses to depth. (2) CPR data also show that the interannual variability in radiolarian abundance since 1997 mirrors very closely the variability of deep ocean organic deposition. (3) The settling material collected in the traps displays a strong correlation between fecal pellets produced by radiolaria and the measured organic carbon flux. These all suggest that the pulses are mediated by radiolarians, a group of protozoans found throughout the world’s oceans and which are widely used by paleontologists to determine past climate conditions. Changes in the upper ocean community structure (between years and on longer timescales) may have profound effects on the ability of the oceans to sequester carbon dioxide from the atmosphere.

Transient dynamics of an altered large marine ecosystem

Overfishing of large-bodied benthic fishes and their subsequent population collapses on the Scotian Shelf of Canada’s east coast1, 2 and elsewhere3, 4 resulted in restructuring of entire food webs now dominated by planktivorous, forage fish species and macroinvertebrates. Despite the imposition of strict management measures in force since the early 1990s, the Scotian Shelf ecosystem has not reverted back to its former structure. Here we provide evidence of the transient nature of this ecosystem and its current return path towards benthic fish species domination. The prolonged duration of the altered food web, and its current recovery, was and is being governed by the oscillatory, runaway consumption dynamics of the forage fish complex. These erupting forage species, which reached biomass levels 900% greater than those prevalent during the pre-collapse years of large benthic predators, are now in decline, having outstripped their zooplankton food supply. This dampening, and the associated reduction in the intensity of predation, was accompanied by lagged increases in species abundances at both lower and higher trophic levels, first witnessed in zooplankton and then in large-bodied predators, all consistent with a return towards the earlier ecosystem structure. We conclude that the reversibility of perturbed ecosystems can occur and that this bodes well for other collapsed fisheries.

Spatial and inter-decadal variability in plankton abundance and composition in the Northwest Atlantic (1958-2006)

The results of Continuous Plankton Recorder sampling in the NW Atlantic between 1958 and 2006 are presented for 11 plankton taxa in eight shelf and deep ocean regions. For shelf regions, phytoplankton abundances increased in the early 1990s, mainly in winter, as the contribution of Arctic-derived freshwater to the Newfoundland (NLS) and Scotian shelves (SS) increased. Farther east, in the sub-polar gyre, phytoplankton levels increased with rising temperatures during the 1990s and 2000s. In both areas, the changes can be explained by increased stratification. The increased influx of arctic water to the NLS in the 1990s was also probably directly responsible for the increased abundances of two arctic Calanus species (C. glacialis and C. hyperboreus) and indirectly responsible for the decreased abundance of Calanus I–IV (mainly C. finmarchicus), perhaps via changes in food composition. On the SS the arctic Calanus species increased in abundance in the 2000s, likely as the result of increased transport from the Arctic via the Gulf of St Lawrence. In the deep ocean, plankton seasonal cycles changed little over the decades and increasing phytoplankton levels in the 2000s were accompanied by increases in zooplankton abundance, suggesting bottom-up control. In shelf regions, phytoplankton increases in the 1990s were in winter and Calanus I–IV appeared earlier in spring than in previous decades. Zooplankton levels generally did not change overall however, perhaps because the species examined were mainly inactive during winter.

A synthesis of large-scale patterns in the planktonic prey of larval and juvenile cod (Gadus morhua)

Data from 40 published studies of the diet composition of larval and juvenile cod (Gadus morhua) from around the northern North Atlantic were summarized to assess generic patterns in ontogenetic and regional variability in the key prey. The results showed that larvae at the northern edge of the latitudinal range of cod depend primarily on development stages of the copepod Calanus finmarchicus, whilst those at the southern edge depend on Para- and Pseudocalanus species. Juvenile cod preyed on a wider range of taxa than larvae, but euphausiids were the main target prey. Analysis of regional variations in the relative abundances of C. finmarchicus and Para/Pseudocalanus spp. in the plankton, as estimated by the continuous plankton recorder (CPR) surveys, showed a similar geographical pattern to the larval cod stomach contents. Comparison of CPR data from the 1960s and 70s with data from the 1990s showed that the boundary between C. finmarchicus and Para/Pseudocalanus spp. dominance has shifted northwards on both sides of the Atlantic, whilst the abundance of euphausiids in the southern cod stock regions has declined. The results are discussed in relation to regional differences in the response of cod stocks to climate variability.

Spatial variability of the plankton trophic interaction in the North Sea: a new feature after the early 1970s

Traditionally, marine ecosystem structure was thought to be bottom-up controlled. In recent years, a number of studies have highlighted the importance of top-down regulation. Evidence is accumulating that the type of trophic forcing varies temporally and spatially, and an integrated view – considering the interplay of both types of control – is emerging. Correlations between time series spanning several decades of the abundances of adjacent trophic levels are conventionally used to assess the type of control: bottom-up if positive or top-down if this is negative. This approach implies averaging periods which might show time-varying dynamics and therefore can hide part of this temporal variability. Using spatially referenced plankton information extracted from the Continuous Plankton Recorder, this study addresses the potential dynamic character of the trophic structure at the planktonic level in the North Sea by assessing its variation over both temporal and spatial scales. Our results show that until the early-1970s a bottom-up control characterized the base of the food web across the whole North Sea, with diatoms having a positive and homogeneous effect on zooplankton filter-feeders. Afterwards, different regional trophic dynamics were observed, in particular a negative relationship between total phytoplankton and zooplankton was detected off the west coast of Norway and the Skagerrak as opposed to a positive one in the southern reaches. Our results suggest that after the early 1970s diatoms remained the main food source for zooplankton filter-feeders east of Orkney–Shetland and off Scotland, while in the east, from the Norwegian Trench to the German Bight, filter-feeders were mainly sustained by dinoflagellates.

Zooplankton response to a warmer northern Wadden Sea

Weekly measurements of mesozooplankton (>76 mu m) and hydrographic parameters have been carried out since 1984 in the List Tidal Basin (northern Wadden Sea). Monthly water temperature significantly increased by 0.04 degrees C year. The largest increase by 3 degrees C in 22 years occurred in September, implying, an extension of the warm summer period. Mean annual copepod abundance and length of copepod season correlated significantly with mean temperature from January to May. Except for an increasing Acartia sp. abundance during spring (April-May), no longterm trends in copepod abundance were observed. The percentage of carnivorous zooplankton increased significantly since 1984 mainly due to a sudden increase in the cyclopoid copepod Oithona similis in 1997. We expect that global warming will lead to a longer copepod season and higher copepod abundances in the northern Wadden Sea.

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.

Recruitment of lesser sandeel Ammodytes marinus in relation to density dependence and zooplankton composition

Recent recruitment failure of lesser sandeel Ammodytes marinus, a key prey fish in the North Sea, followed by several years of low spawning stock biomass, prompted us to investigate factors influencing the recruitment of this species. We tested 2 hypotheses that relate to ecological mechanisms of recruitment regulation in lesser sandeel: (1) a positive spawning stock–recruitment relationship is decoupled in years associated with high abundances of age-1 sandeels and (2) the survival success of early larvae depends specifically on the abundance of Calanus finmarchicus and not C. helgolandicus. The findings of the present study supported both hypotheses and resulted in a multiple linear recruitment model with pronounced predictive capabilities. The model includes interactions between age-1 abundance and spawning stock biomass, plus the effect of C. finmarchicus abundance, and it explained around 65% of the inter-annual variation in recruitment in contrast to only 12% by a traditional Ricker curve. We argue that early egg production in C. finmarchicus supports the survival of larvae, and that climate-generated shifts in the Calanus species composition lead to a mismatch in timing between food availability and the early life history of lesser sandeels.

Mediterranean marine copepods: basin-scale trends of the calanoid Centropages typicus

The Mediterranean Sea is located in a crossroad of mid-latitude and subtropical climatic modes that enhance contrasting environmental conditions over both latitudinal and longitudinal ranges. Here, we show that the large-scale environmental forcing is reflected in the basin scale trends of the adult population of the calanoid copepod Centropages typicus. The species is distributed over the whole Mediterranean basin, and maximal abundances were found in the north-western basin associated to oceanic fronts, and in the Adriatic Sea associated to shallow and semi enclosed waters. The peak of main abundances of C. typicus correlates with the latitudinal temperature gradient and the highest seasonal abundances occurred in spring within the 14–18°C temperature window. Such thermal cline may define the latitudinal geographic region where C. typicus seasonally dominates the >200 μm-sized spring copepod community in the Mediterranean Sea. The approach used here is generally applicable to investigate the large-scale spatial patterns of other planktonic organisms and to identify favourable environmental windows for population development.

Using the Continuous Plankton Recorder to investigate the absence of North Atlantic right whales (Eubalaena glacialis) from the Roseway Basin foraging ground

North Atlantic right whales (Eubalaena glacialis) were absent from Roseway Basin, located off southeastern Nova Scotia, for a 7-year period (1993–1999). The objective of this study was to examine the availability of the right whale's main prey, Calanus finmarchicus, in Roseway Basin during those 7 years to determine if the whales’ absence was due to inadequate prey resources. Since we had no historical data on zooplankton abundances at depth on the Scotian Shelf, near-surface zooplankton abundance data from the Continuous Plankton Recorder were used to infer water-column abundances. In addition, environmental parameters that are often correlated with high zooplankton concentrations were examined. The hypotheses tested were that changes in these parameters would be detectable between three time periods: pre-1993, 1993–1999 and post-1999. Calanus finmarchicus abundance was found to be lowest during 1993–1999, suggesting that right whales were not foraging in Roseway Basin because of the near-absence of their main prey species. Decreased in situ salinity and density proved to be indicators of the changes in circulation in the 1990s that may have affected the advection of C. finmarchicus onto the Scotian Shelf.