A genetic marker to separate Emiliania huxleyi (Prymnesiophyceae) morphotypes

Emiliania huxleyi (Lohm.) Hay and Mohler is a ubiquitous unicellular marine alga surrounded by an elaborate covering of calcite platelets called coccoliths. It is an important primary producer involved in oceanic biogeochemistry and climate regulation. Currently, E. huxleyi is separated into five morphotypes based on morphometric, physiological, biochemical, and immunological differences. However, a genetic marker has yet to be found to characterize these morphotypes. With the use of sequence analysis and denaturing gradient gel electrophoresis, we discovered a genetic marker that correlates significantly with the separation of the most widely recognized A and B morphotypes. Furthermore, we reveal that the A morphotype is composed of a number of distinct genotypes. This marker lies within the 3' untranslated region of a coccolith associated protein mRNA, which is implicated in regulating coccolith calcification. Consequently, we tentatively termed this marker the coccolith morphology motif.

Relating sardine recruitment in the Northern Benguela to satellite-derived sea surface height using a neural network pattern recognition approach

Processes of enrichment, concentration and retention are thought to be important for the successful recruitment of small pelagic fish in upwelling areas, but are difficult to measure. In this study, a novel approach is used to examine the role of spatio-temporal oceanographic variability on recruitment success of the Northern Benguela sardine Sardinops sagax. This approach applies a neural network pattern recognition technique, called a self-organising map (SOM), to a seven-year time series of satellite-derived sea level data. The Northern Benguela is characterised by quasi-perennial upwelling of cold, nutrient-rich water and is influenced by intrusions of warm, nutrient-poor Angola Current water from the north. In this paper, these processes are categorised in terms of their influence on recruitment success through the key ocean triad mechanisms of enrichment, concentration and retention. Moderate upwelling is seen as favourable for recruitment, whereas strong upwelling, weak upwelling and Angola Current intrusion appear detrimental to recruitment success. The SOM was used to identify characteristic patterns from sea level difference data and these were interpreted with the aid of sea surface temperature data. We found that the major oceanographic processes of upwelling and Angola Current intrusion dominated these patterns, allowing them to be partitioned into those representing recruitment favourable conditions and those representing adverse conditions for recruitment. A marginally significant relationship was found between the index of sardine recruitment and the frequency of recruitment favourable conditions (r super(2) = 0.61, p = 0.068, n = 6). Because larvae are vulnerable to environmental influences for a period of at least 50 days after spawning, the SOM was then used to identify windows of persistent favourable conditions lasting longer than 50 days, termed recruitment favourable periods (RFPs). The occurrence of RFPs was compared with back-calculated spawning dates for each cohort. Finally, a comparison of RFPs with the time of spawning and the index of recruitment showed that in years where there were 50 or more days of favourable conditions following spawning, good recruitment followed (Mann-Whitney U-test: p = 0.064, n = 6). These results show the value of the SOM technique for describing spatio-temporal variability in oceanographic processes. Variability in these processes appears to be an important factor influencing recruitment in the Northern Benguela sardine, although the available data time series is currently too short to be conclusive. Nonetheless, the analysis of satellite data, using a neural network pattern-recognition approach, provides a useful framework for investigating fisheries recruitment problems.

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.

Increased blooms of a dinoflagellate in the NW Atlantic

Sampling by the Continuous Plankton Recorder (CPR) over the NW Atlantic from 1960 to 2000 has enabled long-term studies of the larger components of the phytoplankton community, highlighting various changes, particularly during the 1990s. Analysis of an index of phytoplankton biomass, the Phytoplankton Colour Index (PCI) has revealed an increase over the past decade, most marked during the winter (December to February) months. Examination of the structure of the community using multiple linear-regression models indicates that the winter phytoplankton community composition has changed markedly in the 1990s compared to the 1960s. One phytoplankter, the dinoflagellate Ceratium arcticum (Cleve), has undergone dramatic changes in abundance during this period, with pronounced large winter blooms and decreased autumnal levels, and its contribution to the Phytoplankton Colour index values has increased significantly. Other dominant species in the phytoplankton community, both diatoms and dinoflagellates, did not show the same variations over the examined time period. It is suggested that the response of C. arcticum is probably a result of previously reported changes in stratification in the NW Atlantic, due to dynamic hydro-climatic (freshening and cooling) events.

What determines the likelihood of species discovery in marine holozooplankton: is size, range or depth important?

The relationship between date of first description and size, geographic range and depth of occurrence is investigated for 18 orders of marine holozooplankton (comprising over 4000 species). Results of multiple regression analyses suggest that all attributes are linked, which reflects the complex interplay between them. Partial correlation coefficients suggest that geographic range is the most important predictor of description date, and shows an inverse relationship. By contrast, size is generally a poor indicator of description date, which probably mirrors the size-independent way in which specimens are collected, though there is clearly a positive relationship between both size and depth (for metabolic/trophic reasons), and size and geographic range. There is also a positive relationship between geographic range and depth that probably reflects the near constant nature of the deep-water environment and the wide-ranging currents to be found there. Although we did not explicitly incorporate either abundance or location into models predicting the date of first description, neither should be ignored.

Seasonal movements and behaviour of basking sharks from archival tagging: No evidence of winter hibernation

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer-feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of continental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hibernate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (> 750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete productivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

Using continuous plankton recorder data

The continuous plankton recorder (CPR) survey is the largest multi-decadal plankton monitoring programme in the world. It was initiated in 1931 and by the end of 2004 had counted 207,619 samples and identified 437 phyto- and zooplankton taxa throughout the North Atlantic. CPR data are used extensively by the research community and in recent years have been used increasingly to underpin marine management. Here, we take a critical look at how best to use CPR data. We first describe the CPR itself, CPR sampling, and plankton counting procedures. We discuss the spatial and temporal biases in the Survey, summarise environmental data that have not previously been available, and describe the new data access policy. We supply information essential to using CPR data, including descriptions of each CPR taxonomic entity, the idiosyncrasies associated with counting many of the taxa, the logic behind taxonomic changes in the Survey, the semi-quantitative nature of CPR sampling, and recommendations on choosing the spatial and temporal scale of study. This forms the basis for a broader discussion on how to use CPR data for deriving ecologically meaningful indices based on size, functional groups and biomass that can be used to support research and management. This contribution should be useful for plankton ecologists, modellers and policy makers that actively use CPR data.

Climate change and marine plankton

Understanding how climate change will affect the planet is a key issue worldwide. Questions concerning the pace and impacts of climate change are thus central to many ecological and biogeochemical studies, and addressing the consequences of climate change is now high on the list of priorities for funding agencies. Here, we review the interactions between climate change and plankton communities, focusing on systematic changes in plankton community structure, abundance, distribution and phenology over recent decades. We examine the potential socioeconomic impacts of these plankton changes, such as the effects of bottom-up forcing on commercially exploited fish stocks (i.e. plankton as food for fish). We also consider the crucial roles that plankton might have in dictating the future pace of climate change via feedback mechanisms responding to elevated atmospheric CO sub(2) levels. An important message emerges from this review: ongoing plankton monitoring programmes worldwide will act as sentinels to identify future changes in marine ecosystems.

Continuous Plankton Recorder Database: evolution, current uses and future directions.

The Continuous Plankton Recorder (CPR) survey, operated by the Sir Alister Hardy Foundation for Ocean Science (SAHFOS), is the largest plankton monitoring programme in the world and has spanned >70 yr. The dataset contains information from ~200 000 samples, with over 2.3 million records of individual taxa. Here we outline the evolution of the CPR database through changes in technology, and how this has increased data access. Recent high-impact publications and the expanded role of CPR data in marine management demonstrate the usefulness of the dataset. We argue that solely supplying data to the research community is not sufficient in the current research climate; to promote wider use, additional tools need to be developed to provide visual representation and summary statistics. We outline 2 software visualisation tools, SAHFOS WinCPR and the digital CPR Atlas, which provide access to CPR data for both researchers and non-plankton specialists. We also describe future directions of the database, data policy and the development of visualisation tools. We believe that the approach at SAHFOS to increase data accessibility and provide new visualisation tools has enhanced awareness of the data and led to the financial security of the organisation; it also provides a good model of how long-term monitoring programmes can evolve to help secure their future.

Climate effects and benthic-pelagic coupling in the North Sea

The North Sea is one of the most biologically productive ecosystems in the world and supports important fisheries. Climate-induced changes occurred in the pelagic ecosystems of the North Sea during the 1980s. These changes, which have been observed from phytoplankton to fish and among permanent (holoplankton) and temporary (meroplankton) plankton species, have resulted in alterations in plankton community composition and seasonality. Until now, the effects of climate-driven changes on biological linkages between pelagic and benthic ecosystems have not been examined. The present study indicates that changes in benthic organisms could have a profound effect on the trophodynamics of the pelagos. We demonstrate this by analyses of a long-term time series of North Sea plankton and sea surface temperature data. We discover that pronounced changes in the North Sea meroplankton, mainly related to an increased abundance and spatial distribution of the larvae of a benthic echinoderm, Echinocardium cordatum, result primarily from a stepwise increase in sea temperature after 1987 that has caused warmer conditions to occur earlier in the year than previously. Key stages of reproduction in E. cordatum, gametogenesis and spawning, appear to be influenced by winter and spring sea temperature and their larval development is affected by the quantity and quality of their phytoplankton food. Our analyses suggest that a new thermal regime in the North Sea in winter and spring may have benefited reproduction and survival in this benthic species. As a result, E. cordatum may be altering the trophodynamics of the summer pelagic ecosystem through competition between its larvae and holozooplankton taxa.

From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.