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AC is funded by the Ministry of Science and Innovation of Spain through project CTM2009-08783. AM was part funded by NERC UK project NE/K001345/1. KJF was funded by NERC UK through projects NE/H01750X/1 and NE/F003455/1. This review was supported in part by project EURO-BASIN (Ref. 264933, 7FP, European Union), and by a Leverhulme International Network ‘‘Placing marine mixotrophs in context: modelling mixotrophy in a changing world’’., 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., FP7, MINECO, NERC, Leverhulme International Network, 3,2690

Species of the genus Pseudocalanus (Copepoda: Calanoida) are widespread in the North Atlantic Ocean and provide an opportunity to explore ocean-basin-scale genetic relationships of closely related holozooplanktonic species. A phylogeographic analysis of five species of Pseudocalanus, using two mitochondrial DNA markers and one nuclear DNA marker, was conducted. The results show that there are differences in the overall genetic structure among the five species. Three species, P. minutus, P. moultoni and P. elongatus displayed no or little genetic structuring, while P. acuspes and P. newmani showed comparatively higher genetic structure. These results, combined with tree-based and demographic-history analyses, may indicate that the five copepod species investigated herein could represent two diverging evolutionary branches.

Recent evidence suggests that diatoms may induce reproductive failure in planktonic copepods, either through ingestion of diatoms by females or contact of embryos with diatom extracts. Therefore, the co-location of copepods and diatoms in situ may affect copepod reproductive success. As part of a study to determine the effect of diatoms on copepod reproductive effort, we measured the abundance and nighttime vertical distribution of two species of copepods, Calanus pacificus and Pseudocalanus newmani , as well as vertical distribution of chlorophyll a , during winter–spring in Dabob Bay, Washington State, USA, in 2002 and 2003, with limited observations in the summer of both years. Populations of both species declined throughout the winter–spring but rebounded by summer. While C. pacificus females, males, and CV copepodites were not strongly co-located with chlorophyll a concentrations, those same stages of P. newmani were consistently found within the layer containing the chlorophyll maximum. Our limited data on vertical migration show that female and CI–CII copepodites of C. pacificus exhibited normal diel vertical migration behavior, but adult P. newmani exhibited reverse diel migration. Overall, these data suggest that the patterns of reproductive success observed for the two species may be related to their different vertical distributions and may affect their population dynamics.

Copepod secondary production has traditionally been linked to the spring diatom bloom in temperate and high latitudes, but laboratory studies have recently challenged this view and have shown either reduced fecundity or viability of offspring when copepods were fed high concentrations of – mostly unialgal – diatoms. However, field evidence that diatoms affect copepod reproduction is still scarce. We analyzed the reproductive response of a common, small calanoid copepod of the boreal Pacific, Pseudocalanus newmani, to spring diatom blooms in Dabob Bay, a semi-enclosed fjord of Puget Sound, Washington, USA. Abundance patterns, egg production rates, egg hatching success, and naupliar viability of the egg-carrying copepod were examined between February and early May in the years 2002–2004. The population underwent strong variations in abundance during both years, with high abundance of all stages from February to mid-March, but dramatically decreasing individual numbers later in spring. A recovery to higher numbers occurred in July. While egg production rates were independent of chlorophyll concentrations, the reproductive success of P. newmani was negatively affected by certain phytoplankton bloom conditions. Hatching success and – more markedly – naupliar survival were reduced following peaks of Thalassiosira species that were producing anti-mitotic aldehydes, but were high during periods when phytoplankton blooms were more diverse or dominated by other prey taxa including diatoms. As a consequence, recruitment of the naupliar population was considerably affected by the Thalassiosira blooms. This study shows for the first time that the so-called diatom effect operates in nature when all prerequisites – (1) high concentration of aldehyde producers, (2) few prey alternatives, and (3) feeding of copepods on these algae – are given. However, the effect was transient in Dabob Bay and may be so in other pelagic ecosystems. It remains to discern the potential sources of mortality, such as toxicity, malnutrition, predation, and/or advection, that may have caused the strong decline of the Pseudocalanus population during April.

During 24, three-day cruises to Dabob Bay, Washington State, USA, from February 4 to April 26, 2002, and February 4 to May 1 2003, we examined the relative growth and grazing rates of phytoplankton and microzooplankton using dilution experiments. Experiments were conducted over two time intervals: 8–10 h during the nighttime only, or 24 h from noon to noon. We used water from two depths during each cruise: from the surface mixed layer, and from a deep layer below the seasonal thermocline. During 2002, there was one mid-sized bloom consisting mainly of Thalassiosira spp. in early February, and a larger bloom in April comprised of two Chaetoceros spp. and Phaeocystis sp. During 2003, there were also two blooms, one in early February, which was again dominated by Thalassiosira spp., and a second larger bloom in mid-April, comprised mainly of Thalassiosira spp. and Chaetoceros spp. During all four of these blooms, and for both water source depths, specific grazing rates of microzooplankton were most often as high or higher than the calculated phytoplankton specific growth rates. The major microzooplankton categories that could have accounted for this were (1) a large Gyrodinium spp., (2) a group of fusiform-shaped mid-sized Protoperidinium species, and (3) three loosely defined taxonomic groups consisting of naked ciliates, tintinnids, and unidentified heterotrophic dinoflagellates. Based on our measurements, it appears that the microzooplankton community grazing pressure can often exert significant control on phytoplankton biomass, even during the extremely productive spring bloom periods and under several different diatom-dominated bloom types. These results suggest that even in highly productive estuarine ecosystems, which are often nurseries to economically important fisheries species, microzooplankton play a critical role and may significantly alter the availability and efficiency of transfer of energy to higher trophic levels.

During late winter and spring of 2002 and 2003, 24 two- to three-day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani . Here, we discuss the results of our grazing experiments for P. newmani . Each week, we conducted traditional microzooplankton dilution experiments and “copepod dilution” experiments, each from two different layers. Grazing was measured by changes in chlorophyll concentration and direct cell counts. Clearance rates on individual prey species, as calculated by cell counts, showed that Pseudocalanus are highly selective in their feeding, and may have much higher grazing rates on individual taxa than calculated from bulk chlorophyll disappearance. The grazing rates of the copepods, however, are typically an order of magnitude lower than the grazing rates of the microzooplankton community, or the growth rates of the phytoplankton. P. newmani ingested diatoms, but, at certain times fed preferentially on microzooplankton, such as ciliates, tintinnids, and larger dinoflagellates. Removal of the microzooplankton may have released the other phytoplankton species from grazing pressure, allowing those species’ abundance to increase, which was measured as an apparent “negative” grazing on those phytoplankton species. The net result of grazing on some phytoplankton species, while simultaneously releasing others from grazing pressure resulted in bulk chlorophyll-derived estimates of grazing which were essentially zero or slightly negative; thus bulk chlorophyll disappearance is a poor indicator of copepod grazing. Whether copepods can significantly release phytoplankton from the grazing pressure by microzooplankton in situ, thus causing a trophic cascade, remains to be verified, but is suggested by our study.

During late winter and spring of 2002 and 2003, 24, 2–3 day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. The results of the copepod grazing experiments for C. pacificus are discussed here. Each week, copepod grazing incubation experiments from two different depth layers were conducted. Grazing was measured by both changes in chlorophyll concentration and cell counts. In 2002, there was one moderate bloom consisting mainly of Thalassiosira spp. in early February, and a larger bloom in April comprised of two Chaetoceros species and Phaeocystis sp. Similarly, in 2003, there were two blooms, an early one dominated by Thalassiosira spp., and a later one consisting of Chaetoceros spp. and Thalassiosira spp. Clearance rates on individual prey species, as calculated by cell counts, showed that C. pacificus are highly selective in their feeding, and may have much higher clearance rates on individual taxa than rates calculated from bulk chlorophyll disappearance. During weeks of high phytoplankton concentration, the copepods generally ate phytoplankton. However, they often rejected the most abundant phytoplankton species, particularly certain Thalassiosira spp., even though the rejected prey were often of the same genus and similar size to the preferred prey. It is speculated that this avoidance may be related to the possible deleterious effects that certain of these diatom species have on the reproductive success of these copepods. During weeks of medium to low phytoplankton concentration, the copepods selectively ate certain species of phytoplankton, and often had high electivity for microzooplankton. The selection mechanism must consist of active particle rejection most likely based on detection of surface chemical properties, since the diatoms that were selected were of the same genus, nearly the same size, and at lower numerical abundance than those cells that were avoided. The grazing choices made by these copepods may have important consequences for the overall ecosystem function within coastal and estuarine systems through changes in the transfer efficiency of energy to higher trophic levels.