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

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

Intraspecific differences in short- and long-term foraging strategies of reef manta ray (Mobula alfredi) in the Chagos Archipelago

Assessing the foraging ecology of a threatened species is necessary to understand their movement behaviour and habitat use patterns, which are essential for developing effective protection strategies. Here, the foraging ecology of reef manta rays (Mobula alfredi) in the Chagos Archipelago, a region encompassed by a vast no-take marine protected area (MPA), was investigated using stable isotope analysis of skin and muscle tissue. Enriched δ13C values suggest the population predominantly forages in nearshore environments. Skin δ13C values increased with increased rainfall, likely associated with the boosts in primary production and zooplankton biomass due to the coastal advection of seabird guano. Annual variations in δ13C values of skin and muscle were observed and are consistent with reduced nutrient transport associated with the effects of Indian Ocean Dipole oscillations, including a deepening of the thermocline, a suppression of cold-water upwelling, and reduced rainfall. Short- and long-term foraging strategies and locations were identified by applying hierarchical clustering, isotopic niche analysis, and Bayesian stable isotope mixing models to δ13C and δ15N of paired skin and muscle tissue samples. Two isotopically distinct groups of M. alfredi were identified, employing either local foraging strategies restricted to specific locations or wide-ranging strategies that likely mean they engage in regular migrations throughout the archipelago. Ninety-eight percent of M. alfredi were estimated to switch between strategies utilising and connecting multiple discrete nearshore habitats, emphasising their role in ecosystem functioning by facilitating the transport of nutrients across ecosystem boundaries. However, illegal, unreported, and unregulated fishing and lost or abandoned fishing gear commonly occur within the MPA. Locations of particular concern are Egmont Atoll as it is a highly active aggregation location and Peros Banhos Atoll where IUU frequently occurs and M. alfredi are estimated to be heavily reliant upon for foraging. Frequent migrations between atolls by M. alfredi also raises concern over their vulnerability to these activities along migration corridors. This research bridges current knowledge gaps in this population's foraging ecology and concomitant movement patterns, which should inform conservation strategies in the region