The Russell Cycle. An update and review of trends in zooplankton and fish larvae off Plymouth 1924-2009. Occasional Publication of the Marine Biological Association 24

Regular plankton sampling off Plymouth by the Marine Biological Association (MBA) has been carried out from the early 1900s. Much of the sample analysis and description of the results was carried out by Sir Frederick Russell and Professor Alan Southward (AJS), the latter having completed the organisation and transfer of the paper records to digital files. The current authors have transferred the main data files of AJS on zooplankton and fish larvae to the MBA long-term database (including various editing and checking against original analysis records and published data) together with adding the data for 2002-2009. In this report the updated time-series are reviewed in the context of earlier work, particularly with respect to the Russell Cycle. It is not intended as an exhaustive analysis. Brief details of the sampling and comments on data processing are given in an appendix.

Bridging the gap between marine biogeochemical and fisheries sciences; configuring the zooplankton link

Exploring climate and anthropogenic impacts on marine ecosystems requires an understanding of how trophic components interact. However, integrative end-to-end ecosystem studies (experimental and/or modelling) are rare. Experimental investigations often concentrate on a particular group or individual species within a trophic level, while tropho-dynamic field studies typically employ either a bottom-up approach concentrating on the phytoplankton community or a top-down approach concentrating on the fish community. Likewise the emphasis within modelling studies is usually placed upon phytoplankton-dominated biogeochemistry or on aspects of fisheries regulation. In consequence the roles of zooplankton communities (protists and metazoans) linking phytoplankton and fish communities are typically under-represented if not (especially in fisheries models) ignored. Where represented in ecosystem models, zooplankton are usually incorporated in an extremely simplistic fashion, using empirical descriptions merging various interacting physiological functions governing zooplankton growth and development, and thence ignoring physiological feedback mechanisms. Here we demonstrate, within a modelled plankton food-web system, how trophic dynamics are sensitive to small changes in parameter values describing zooplankton vital rates and thus the importance of using appropriate zooplankton descriptors. Through a comprehensive review, we reveal the mismatch between empirical understanding and modelling activities identifying important issues that warrant further experimental and modelling investigation. These include: food selectivity, kinetics of prey consumption and interactions with assimilation and growth, form of voided material, mortality rates at different age-stages relative to prior nutrient history. In particular there is a need for dynamic data series in which predator and prey of known nutrient history are studied interacting under varied pH and temperature regimes.

Identification of the copepodite developmental stages of twenty-six North Atlantic copepods. Occasional Publication of the Marine Biological Association 21.

In zooplankton copepod studies there is often the requirement to be able to identify the six copepodite developmental stages of different species, or to know their body dimensions. However, this information is not available for many species, or is dispersed through the literature. This guide gathers together both original and previously published information on morphology and measurements for the stages of twenty-six common North Atlantic copepod species and tabulates them in a standard format. For each species additional notes useful in their identification are also given.

Flexible diel vertical migration behaviour of zooplankton in the Irish Sea

The diel vertical migration (DVM) of the whole plankton community was investigated in the central and coastal Irish Sea. Generally, more than 60% of the plankton did not perform significant DVM. A correlation analysis of the weighted mean depths of different organisms and their potential predators suggested relationships between two groups, Oithona spp., copepod nauplii and fish larvae, and between Pseudocalanus elongatus, Calanus spp. and chaetognaths. The organisms showing significant DVM were chaetognaths (Sagitta spp.), Calanus spp. and P. elongatus. Calanus spp. showed clear ontogenic variations in DVM, and along with P. elongatus demonstrated great flexibility both in the amplitude and direction of migration. P. elongatus did not migrate in the coastal area and Calanus spp. showed a clear reverse migration. The direction of migration appeared to be related to the vertical position of the chaetognaths in the water column during the day.

Use of Continuous Plankton Recorder information in support of marine management: applications in fisheries, environmental protection, and in the study of ecosystem response to environmental change

The Continuous Plankton Recorder (CPR) survey was conceived from the outset as a programme of applied research designed to assist the fishing industry. Its survival and continuing vigour after 70 years is a testament to its utility, which has been achieved in spite of great changes in our understanding of the marine environment and in our concerns over how to manage it. The CPR has been superseded in several respects by other technologies, such as acoustics and remote sensing, but it continues to provide unrivalled seasonal and geographic information about a wide range of zooplankton and phytoplankton taxa. The value of this coverage increases with time and provides the basis for placing recent observations into the context of long-term, large-scale variability and thus suggesting what the causes are likely to be. Information from the CPR is used extensively in judging environmental impacts and producing quality status reports (QSR); it has shown the distributions of fish stocks, which had not previously been exploited; it has pointed to the extent of ungrazed phytoplankton production in the North Atlantic, which was a vital element in establishing the importance of carbon sequestration by phytoplankton. The CPR continues to be the principal source of large-scale, long-term information about the plankton ecosystem of the North Atlantic. It has recently provided extensive information about the biodiversity of the plankton and about the distribution of introduced species. It serves as a valuable example for the design of future monitoring of the marine environment and it has been essential to the design and implementation of most North Atlantic plankton research.

Encounter success of free-ranging marine predator movements across a dynamic prey landscape

Movements of wide-ranging top predators can now be studied effectively using satellite and archival telemetry. However, the motivations underlying movements remain difficult to determine because trajectories are seldom related to key biological gradients, such as changing prey distributions. Here, we use a dynamic prey landscape of zooplankton biomass in the north-east Atlantic Ocean to examine active habitat selection in the plankton-feeding basking shark Cetorhinus maximus. The relative success of shark searches across this landscape was examined by comparing prey biomass encountered by sharks with encounters by random-walk simulations of ‘model’ sharks. Movements of transmitter-tagged sharks monitored for 964 days (16754km estimated minimum distance) were concentrated on the European continental shelf in areas characterized by high seasonal productivity and complex prey distributions. We show movements by adult and sub-adult sharks yielded consistently higher prey encounter rates than 90% of random-walk simulations. Behavioural patterns were consistent with basking sharks using search tactics structured across multiple scales to exploit the richest prey areas available in preferred habitats. Simple behavioural rules based on learned responses to previously encountered prey distributions may explain the high performances. This study highlights how dynamic prey landscapes enable active habitat selection in large predators to be investigated from a trophic perspective, an approach that may inform conservation by identifying critical habitat of vulnerable species.

Use of satellite data for modelling food availability and survival of marine fish larvae

A modelling scheme is described which uses satellite retrieved sea-surface temperature and chlorophyll-a to derive monthly zooplankton biomass estimates in the eastern North Atlantic; this forms part of a bio-physical model of inter-annual variations in the growth and survival of larvae and post-larvae of mackerel (Scomber scombrus). The temperature and chlorophyll data are incorporated first to model copepod (Calanus) egg production rates. Egg production is then converted to available food using distribution data from the Continuous Plankton Recorder (CPR) Survey, observed population biomass per unit daily egg production and the proportion of the larval mackerel diet comprising Calanus. Results are validated in comparison with field observations of zooplankton biomass. The principal benefit of the modelling scheme is the ability to use the combination of broad scale coverage and fine scale temporal and spatial variability of satellite data as driving forces in the model; weaknesses are the simplicity of the egg production model and the broad-scale generalizations assumed in the raising factors to convert egg production to biomass.