Reprint of: High prey-predator size ratios and unselective feeding in copepods: A seasonal comparison of five species with contrasting feeding modes

There has been an upsurge of interest in trait-based approaches to zooplankton, modelling the seasonal changes in the feeding modes of zooplankton in relation to phytoplankton traits such as size or motility. We examined this link at two English Channel plankton monitoring sites south of Plymouth (L4 and E1). At L4 there was a general transition from diatoms in spring to motile microplankton in summer and autumn, but this was not mirrored in the succession of copepod feeding traits; for example the ambushing Oithona similis dominated during the spring diatom bloom. At nearby E1 we measured seasonality of food and grazers, finding strong variation between 2014 and 2015 but overall low mesozooplankton biomass (median 4.5 mg C m−3). We also made a seasonal grazing study of five copepods with contrasting feeding modes (Calanus helgolandicus, Centropages typicus, Acartia clausi, Pseudocalanus elongatus and Oithona similis), counting the larger prey items from the natural seston. All species of copepod fed on all food types and differences between their diets were only subtle; the overriding driver of diet was the composition of the prey field. Even the smaller copepods fed on copepod nauplii at significant rates, supporting previous suggestions of the importance of intra-guild predation. All copepods, including O. similis, were capable of tackling extremely long (>500 µm) diatom chains at clearance rates comparable to those on ciliates. Maximum observed prey:predator length ratios ranged from 0.12 (C. helgolandicus) up to 0.52 (O. similis). Unselective feeding behaviour and the ability to remove highly elongated cells have implications for how copepod feeding is represented in ecological and biogeochemical models

Disentangling the counteracting effects of water content and carbon mass on zooplankton growth

Abstract Zooplankton vary widely in carbon percentage (carbon mass as a percentage of wet mass), but are often described as either gelatinous or non-gelatinous. Here we update datasets of carbon percentage and growth rate to investigate whether carbon percentage is a continuous trait, and whether its inclusion improves zooplankton growth models. We found that carbon percentage is continuous, but that species are not distributed homogenously along this axis. To assess variability of this trait in situ, we investigated the distribution of biomass across the range of carbon percentage for a zooplankton time series at station L4 off Plymouth, UK. This showed separate biomass peaks for gelatinous and crustacean taxa, however, carbon percentage varied 8-fold within the gelatinous group. Species with high carbon mass had lower carbon percentage, allowing separation of the counteracting effects of these two variables on growth rate. Specific growth rates, g (d−1) were negatively related to carbon percentage and carbon mass, even in the gelatinous taxa alone, suggesting that the trend is not driven by a categorical difference between these groups. The addition of carbon percentage doubled the explanatory power of growth models based on mass alone, demonstrating the benefits of considering carbon percentage as a continuous trait

Seasonal variation of zooplankton community structure and trophic position in the Celtic Sea: a stable isotope and biovolume spectrum approach

Zooplankton on continental shelves represent an important intermediary in the transfer of energy and matter from phytoplankton to the wider ecosystem. Their taxonomic composition and trophic interactions with phytoplankton vary in space and time, and interpreting the implications of this constantly evolving landscape remains a major challenge. Here we combine plankton taxonomic data with the analysis of biovolume spectra and stable isotopes to provide insights into the trophic interactions that occur in a shelf sea ecosystem (Celtic Sea) across the spring-summer-autumn transition. Biovolume spectra captured the seasonal development of the zooplankton community well, both in terms of total biomass and trophic positioning, and matched trophic positions estimated by stable isotope analysis. In early April, large microplankton (63-200 µm) occupied higher trophic positions than mesozooplankton (>200 µm), likely reflecting the predominance of nanoplankton (2-20 µm) that were not readily available to mesozooplankton grazers. Biomass and number of trophic levels increased during the spring bloom as elevated primary production allowed for a higher abundance of predatory species. During July, the plankton assemblage occupied relatively high trophic positions, indicating important links to the microbial loop and the recycling of organic matter. The strong correlation between biomass and community trophic level across the study suggests that the Celtic Sea is a relatively enclosed and predominantly energy-limited ecosystem. The progression of the zooplankton biomass and community structure within the central shelf region was different to that at the shelf-break, potentially reflecting increased predatory control of copepods by macrozooplankton and pelagic fishes at the shelf break. We suggest that the combination of size spectra and stable isotope techniques are highly complementary and useful for interpreting the seasonal progression of trophic interactions in the plankton

Use of the dilution technique to determine microzooplankton herbivory in three contrasting oceanic systems

The dilution technique has been used during both laboratory experiments and in the field in three contrasting oceanic systems: the northeast Atlantic, Bellingshausen Sea and Arabian Sea. Laboratory dilution grazing studies, using Oxyrrhis marina as the predator and Dunaliella tertiolecta as prey, have demonstrated a significant linear relationship between apparent phytoplankton growth and dilution factor. The daily turnover of prey by Oxyrrhis ranged between 45 and 95%. Growth of prey was constant at all dilutions demonstrating that phytoplankton growth was density independent. Studies carried out during summer along a transect in the northeast Atlantic between 60°N and 47°N showed microzooplankton to consume between 0.5 and 17mg phytoplankton C d&quot;\ This was equivalent to between 288 and 589 mg C m ^ d&quot;^ being grazed in the mixed layer. Microzooplankton grazing impact was highest at the southerly end of the transect. During the spring-bloom period in 1990, microzooplankton grazed between 28 and 78% and between 26 and 55% of the daily primary respectively. The microzooplankton community was dominated by small cells (&lt;20|jm). In the Bellingshausen Sea, microzooplankton consumed between 0.04 and 31.21 mg C m ^ d'\ and were estimated to graze between 21 and 3260 mg C m ^ d'^ in the pack ice and open water station respectively. In the Arabian Sea, microzooplankton grazed between 1 and 17 mg C m&quot;^ d'\ In the mixed layer microzooplankton grazed between 161 and 415 mg C m&quot;^ d&quot;^ during the SW monsoon and between 110 and 407 mg C m&quot;^ d&quot;^ during the intermonsoon period. Substantial grazing of phytoplankton biomass by the microzooplankton occurred in all three oceanic systems. Variations in the grazing and phytoplankton growth rates can be attributed to differences in the phytoplankton and microzooplankton communities encountered. Results demonstrate a tight coupling between growth of prey and consumption for the northeast Atlantic and Bellingshausen Sea. production in May and June</p

The herbivorous impact of microzooplankton during two short-term Lagrangian experiments off the NW coast of Galicia in summer 1998

Microzooplankton (heterotrophic microplankton and heterotrophic nanoflagellates) and their herbivorous activity were estimated from dilution experiments in August 1998 during two Lagrangian drift experiments that sampled contrasting conditions—an upwelling/relaxation event along the shelf edge and an oligotrophic offshore filament. During upwelling/relaxation, heterotrophic microplankton were present at mean surface concentrations between 15,000 and 48,000 cells l?1. Heterotrophic nanoflagellate concentrations were between 200 and 700 cells ml?1 and the most abundant component of the heterotrophic microplankton was the aloricate choreotrich ciliates which increased dramatically in concentration from 6,000 to 24,000 cells l?1 during the first 4 days of the study. Total microzooplankton biomass reached a maximum of 39mgC.m?3. In the filament, which developed from the upwelling, cell concentrations were lower and averaged 4,500 cells l?1 for heterotrophic microplankton and 250 cells ml?1 for heterotrophic nanoflagellates. Total microzooplankton biomass was about 10–12mgC.m?3. Microzooplankton turned over between 40 and 85% of the phytoplankton standing stock, thereby consuming between 5 and 78mg phytoplankton carbon.m?3.d?1. The magnitude of this activity was highest during upwelling/relaxation and was positively correlated to heterotrophic nanoflagellate biomass and chlorophyll-a concentration but not heterotrophic microplankton biomass. The proportion of primary production grazed decreased from 160 to 59% d?1 during upwelling/relaxation and ranged between 60 and 90% d?1 in the filament. Microzooplankton herbivory within the euphotic zone increased from 684 to &gt;2000mgC.m?2.d?1 during upwelling/relaxation and was between 327 and 802mgC.m?2.d?1 in the filament. Although microzooplankton herbivory was lower and less variable during the filament study, microzooplankton consumed on average 60% of the phytoplankton standing stocks which was higher than found during upwelling/relaxation. Microzooplankton assimilation efficiency ranged between 3 and 33% during upwelling/relaxation and between 0 and 13% in the filament. Our data demonstrate a close coupling between phytoplankton growth and microzooplankton herbivory in surface waters off the Galician Coast and suggest that microzooplankton may have been a significant sink for phytogenic carbon during August 1998

Microbial plankton communities in the coastal southeastern Black Sea: biomass, composition and trophic interactions

Summary: We investigated biomass and composition of the pico-, nano- and microplankton communities in a coastal station of the southeastern Black Sea during 2011. We also examined trophic interactions within these communities from size-fractionated dilution experiments in February, June and December. Autotrophic and heterotrophic biomasses showed similar seasonal trends, with a peak in June, but heterotrophs dominated throughout the year. Autotrophic biomass was mainly comprised by nanoflagellates and diatoms in the first half of the year, and by dinoflagellates and Synechococcus spp. in the second half. Heterotrophic biomass was mostly dominated by heterotrophic bacteria, followed by nanoflagellates and microzooplankton. Dilution experiments suggest that nano- and microzooplankton were significant consumers of autotrophs and heterotrophic bacteria. More than 100% of bacterial production was consumed by grazers in all experiments, while 46%, 21% and 30% of daily primary production were consumed in February, June and December, respectively. In February, autotrophs were the main carbon source, but in December, it was heterotrophic bacteria. An intermediate situation was observed in June, with similar carbon flows from autotrophs and heterotrophic bacteria. Size-fraction dilution experiments suggested that heterotrophic nanoflagellates are an important link between the high heterotrophic bacterial biomass and microzooplankton. In summary, these results indicate that nano- and microzooplankton were responsible for comprising a significant fraction of total microbial plankton biomass, standing stocks, growth and grazing processes. This suggests that in 2011, the microbial food web was an important compartment of the planktonic food web in the coastal southeastern Black Sea. Keywords: Phytoplankton, Microzooplankton, Carbon biomass, Microbial food web, Grazing, Black Se

Seawater carbonate chemistry and reproduction of the two calanoid copepods Centropages typicus and Temora longicornis in a laboratory experiment

Some planktonic groups suffer negative effects from ocean acidification (OA), although copepods might be less sensitive. We investigated the effect of predicted CO2 levels (range 480-750 ppm), on egg production and hatching success of two copepod species, Centropages typicus and Temora longicornis. In these short-term incubations there was no significant effect of high CO2 on these parameters. Additionally a very high CO2 treatment, (CO2 = 9830 ppm), representative of carbon capture and storage scenarios, resulted in a reduction of egg production rate and hatching success of C. typicus, but not T. longicornis. In conclusion, reproduction of C. typicus was more sensitive to acute elevated seawater CO2 than that of T. longicornis, but neither species was affected by exposure to CO2 levels predicted for the year 2100. The duration and seasonal timing of exposures to high pCO2, however, might have a significant effect on the reproduction success of calanoid copepods

Comment. What drives plankton seasonality in a stratifying shelf sea? Some competing and complementary theories

The Plymouth L4 time plankton series in the Western English Channel is a textbook example of a shallow, stratifying shelf sea system. Over its 30 yr of weekly sampling, this site has provided a diverse and contrasting suite of numerical and conceptual models of plankton bloom formation, phenology, and seasonal succession. The most recent of these papers, Kenitz et al. (2017) has initiated this comment, partly because we feel that it has presented a slightly misleading picture of the plankton composition at this site, and of a robust, recurring seasonal succession. We address this by illustrating the extent of inter-annual variability in phenology that occurs at the site, and which needs to be captured better within models. However our main aim is to foster a much better integration of the variety of top-down and bottom-up processes that have all been suggested to be key in driving seasonal succession. Some of these, particularly the multiple grazing and growth controls contributing to the so-called "loophole hypothesis" may be complementary, but others, such as the role of copepod feeding traits in driving species succession (Kenitz et al. 2017) offer testable competing hypotheses. The basic assumptions and outputs of all these models need to be validated more critically, both against time series data and process studies that include the finding of unselective feeding. We suggest that the variability in plankton phenology (and not just mean timing and amplitude) could be used to diagnose the performance of alternative models of plankton succession