The importance of zooplankton in reducing levels of atmospheric CO sub(2) via the biological pump

Zooplankton play a key role in climate change through the transfer of large quantities of CO sub(2) to the deep ocean by a process known as the biological pump. Plankton composition is crucial as associated mineral material facilitates sinking of carbon rich debris and some taxa package faecal and detrital material. Ocean acidification may impact calcareous groups. Zooplankton have also been shown to be highly sensitive indicators of environmental change. Results will be presented to show that ocean temperature, circulation and planktonic ecosystems (using data from the Continuous Plankton Recorder, CPR survey) in the North Atlantic are changing rapidly in concert and that there is evidence to suggest that the changes are an ocean wide response to global warming with potential feedback effects. Given the importance of the oceans to the carbon cycle, even a minor change in the flux of carbon to the deep ocean would have a big impact increasing growth of atmospheric CO sub(2). We have virtually no understanding of the spatial and temporal variability in the efficiency of the biological pump for most of the world's ocean. Establishing new plankton monitoring programmes backed up by appropriate research to help understand processes is needed to address this gap in knowledge. There is little doubt within a global change context and the future of mankind that a potential acceleration in the growth of atmospheric carbon due to a reduction in the efficiency of the biological pump is a key issue for future research in zooplankton ecology.

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.

An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 C water, with maximum abundances from 13-17 C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 C, with peak abundances from 0 to 9 C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.

A change in the zooplankton of the central North Sea (55 degree to 58 degree N): A possible consequence of changes in the benthos

The mesozooplankton taken in continuous plankton recorder samples from the Central North Sea has changed from being numerically dominated by holoplanktonic calanoid copepod species from 1958 to the late 1970s to a situation where pluteus larvae of echinoid and ophiuroid echinoderms have been more abundant than any single holoplanktonic species in the 1980s and early 1990s. The abundance of the echinoderm larvae as a proportion of the zooplankton taken in the samples has followed a continuous increasing trend over the Dogger Bank, but off the eastern coast of northern England and southern Scotland the increase did not become obvious until the 1980s. This trend is consistent with reported increases in abundance of the macrobenthos. It is proposed that changes in the benthos have influenced the composition of the plankton.

Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea

Analyses of long-term time series of North Sea plankton and sea surface temperature (SST) data reveal that the annual planktonic larval abundance of three benthic phyla, Echinodermata, Arthropoda, and Mollusca, responds positively and immediately to SST. Long-term outcomes for the planktonic abundance of these three phyla are different, however. The planktonic larvae of echinoderms and decapod crustaceans have increased in abundance from 1958 to 2005, and especially since the mid-1980s, as North Sea SST has increased. In contrast, the abundance of bivalve mollusc larvae has declined, despite the positive year-to-year relationship between temperature and bivalve larval abundance continuing to hold. We argue that the changes in meroplankton abundance, coincident with increased phytoplankton and declining holoplankton, reflect the synchronous effect of rising SST and related changes in the pelagic community on the reproduction and recruitment of many benthic marine invertebrates. Under this scenario, the long-term decline in bivalve mollusc larvae will reflect increased predation on the settled larvae and adults by benthic decapods. These alterations in the zooplankton may therefore describe an ecosystem-wide restructuring of North Sea trophic interactions.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

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.

Macro-ecology of plankton–seabird associations in the North Pacific Ocean

In conjunction with the North Pacific Continuous Plankton Recorder program, we conducted surveys of seabirds from June 2002 to June 2007. Here, we tested the hypotheses of (i) east–west variations in coupled plankton and seabird abundance, and (ii) that surface-feeding and diving seabirds vary in their relationships to primary productivity and mesozooplankton species abundance and diversity. To test these hypotheses, we developed statistical models for 20 species of seabirds and 12 zooplankton taxonomic groups. Seabird density was highly variable between seasons, but was consistently higher in the western than eastern North Pacific. Seabird diversity was greater in the east. Zooplankton abundance did not differ between regions. We found associations at the “bulk” level between seabird density and net primary productivity, but only one association between seabirds and total zooplankton abundance or diversity. However, we found many relationships between seabird species and the abundance of different zooplankton summarized at the genus or family level. Some of these taxonomic relationships reflect direct predator–prey interactions, while others may reflect zooplankton that serve as ecological indicators of other prey, such as micronekton, upon which the birds may feed. Surface or near-surface feeding, mostly piscivorous seabirds, did not differ systematically from diving, mainly planktivorous seabirds in their zooplankton associations. Seabirds apparently respond to zooplankton taxonomic groupings more so than bulk zooplankton characteristics, such as abundance or diversity. Macro-ecological studies of remote marine ecosystems using zooplankton and seabirds as ecological indicators provide a framework for understanding and assessing spatial and temporal variations in these difficult-to-study pelagic environments.

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.

Have we been underestimating the effects of ocean acidification in zooplankton?

Understanding how copepods may respond to ocean acidification (OA) is critical for risk assessments of ocean ecology and biogeochemistry. The perception that copepods are insensitive to OA is largely based on experiments with adult females. Their apparent resilience to increased carbon dioxide (pCO2 ) concentrations has supported the view that copepods are 'winners' under OA. Here, we show that this conclusion is not robust, that sensitivity across different life stages is significantly misrepresented by studies solely using adult females. Stage-specific responses to pCO2 (385-6000 μatm) were studied across different life stages of a calanoid copepod, monitoring for lethal and sublethal responses. Mortality rates varied significantly across the different life stages, with nauplii showing the highest lethal effects; nauplii mortality rates increased threefold when pCO2 concentrations reached 1000 μatm (year 2100 scenario) with LC50 at 1084 μatm pCO2 . In comparison, eggs, early copepodite stages, and adult males and females were not affected lethally until pCO2 concentrations ≥3000 μatm. Adverse effects on reproduction were found, with >35% decline in nauplii recruitment at 1000 μatm pCO2 . This suppression of reproductive scope, coupled with the decreased survival of early stage progeny at this pCO2 concentration, has clear potential to damage population growth dynamics in this species. The disparity in responses seen across the different developmental stages emphasizes the need for a holistic life-cycle approach to make species-level projections to climate change. Significant misrepresentation and error propagation can develop from studies which attempt to project outcomes to future OA conditions solely based on single life history stage exposures.