Lipid-storing copepods are fundamental to the functioning of marine ecosystems, transferring energy from primary producers to higher trophic levels and sequestering atmospheric carbon (C) in the deep ocean. Quantifying trophic transfer and biogeochemical cycling by copepods requires improved understanding of copepod metabolic rates in both surface waters and during lipid-fueled metabolism over winter. Here we present new biomass turnover rates of C and nitrogen (N) in Calanoides acutus, Calanoides natalis, Calanus glacialis and Calanus hyperboreus alongside published data for Calanus finmarchicus and Calanus pacificus. Turnover rates in metabolically active animals, normalised to 10°C, ranged between 0.007 – 0.105 d-1 and 0.004 – 0.065 d-1 for C and N, respectively. Turnover rates of C were typically faster than those for N, supporting the understanding that non-protein C, e.g. lipid, is catabolised faster than protein. Re-analysis of published data indicates that inactive, overwintering C. finmarchicus turn over wax ester lipids at a rate of 0.0016 d-1. These and other basal rate data will facilitate the mechanistic representation of copepod physiology in global biogeochemical models, thereby reducing uncertainties in our predictions of future ocean ecosystem functioning and C sequestration.

The changing Arctic environment is affecting zooplankton that support its abundant wildlife. We examined how these changes are influencing a key zooplankton species, Calanus finmarchicus, principally found in the North Atlantic but expatriated to the Arctic. Close to the ice-edge in the Fram Strait, we identified areas that, since the 1980s, are increasingly favourable to C. finmarchicus. Field-sampling revealed part of the population there to be capable of amassing enough reserves to overwinter. Early developmental stages were also present in early summer, suggesting successful local recruitment. This extension to suitable C. finmarchicus habitat is most likely facilitated by the long-term retreat of the ice-edge, allowing phytoplankton to bloom earlier and for longer and through higher temperatures increasing copepod developmental rates. The increased capacity for this species to complete its life-cycle and prosper in the Fram Strait can change community structure, with large consequences to regional food-webs.

The boreal copepod Calanus finmarchicus sequesters substantial amounts of carbon (C) in the deep layers of the North Atlantic Ocean through their contribution to the “lipid pump.” This pump is driven by these zooplankton descending from the surface layers to spend prolonged periods at depth during which time they metabolise substantial lipid reserves and a fraction suffer mortality. C. finmarchicus is principally a boreal species but is expatriated by currents flowing northwards into Arctic regions such as the Fram Strait, where it is now able to complete its life cycle. We considered how this expansion to its distributional range adds to the estimated magnitude of the lipid pump. Field sampling in the Fram Strait found C. finmarchicus abundance to be spatially variable with high values, equivalent to those reported for core distributional areas further south, found mainly in the eastern region. Lipid reserve levels were sufficient for many individuals to survive the overwintering period and reproduce the following spring. In accordance with abundance patterns, lipid pump magnitude was greater in the Eastern Fram Strait (2.04 g C m−2 year−1) compared to the Western Fram Strait (0.33 g C m−2 year−1). At least for the eastern region, these rates are similar to those reported for this species elsewhere (average of 4.35 g C m−2 year−1). When extrapolated to the wider spatial area of the Fram Strait, the lipid pump generated by this species in this ocean sector amounts to 0.3 Mt C year−1. Although constituting a modest proportion of the total C. finmarchicus lipid pump of 19.3 Mt C year−1, it indicates that the continued northwards expansion of this species will act to increase the size of its lipid pump, which may counteract that lost through the northwards retreat of its Arctic congeners, Calanus glacialis and Calanus hyperboreus.

Calanoid copepods comprise around 90% of Arctic zooplankton biomass and are fundamental to the ecological and biogeochemical functioning of high-latitude pelagic ecosystems. They accumulate lipid reserves during the productive months and represent an energy-rich food source for higher trophic levels. Rapidly changing climate in the Arctic may alter the quantity and composition of the food environment for one of the key copepod species, Calanus finmarchicus, with as yet unquantified effects on its production. Here we present rates of feeding and egg production in female C. finmarchicus exposed to the range of feeding conditions encountered across the Fram Strait in May/June 2018. Carbon (C) budgets were constructed and used to examine the relationship between feeding and growth (= egg production) in these animals. C-specific ingestion rates (mean ± standard deviation) were highly variable, ranging from 0.015 ± 0.004 to 0.645 ± 0.017 day-1 (mean = 0.295 ± 0.223 day-1), and were positively correlated with food availability. C-specific egg production rates ranged from 0.00 to 0.049 day-1 (mean = 0.012 ± 0.011) and were not correlated with either food availability or ingestion rate. Calculated gross growth efficiencies (GGE: growth/ingestion) were low, 0.12 ± 0.13 (range = 0.01 to 0.39). The assembled C budgets indicate that the average fraction of ingested food that was surplus to the requirements for egg production, respiration and losses to faecal pellets was 0.17 ± 0.42. We suggest that this excess occurred, at least in part, because many of the incubated females were still undergoing the energetically (C-) expensive process of gonad maturation at the time of sampling, an assertion that is supported by the relatively high C:N (nitrogen) ratios of the incubated females, the typically low egg production rates, and gonad maturation status. Ontogenetic development may thus explain the large variability seen in the relationship between egg production and ingestion. The apparently excessive ingestion rates may additionally indicate that recently moulted females must acquire additional N via ingestion to complete the maturation process and begin spawning. Our results highlight the need for improved fundamental understanding of the physiology of high-latitude copepods and its response to environmental change.

Mesopelagic fish are an important component of marine ecosystems, and their contribution to marine biogeochemical cycles is becoming increasingly recognized. However, major uncertainties remain in the rates at which they remineralize organic matter. We present respiration rate estimates of mesopelagic fish from two oceanographically contrasting regions: the Scotia Sea and the Benguela Current. Respiration rates were estimated by measuring the enzyme activities of the electron transport system. Regression analysis of respiration with wet mass highlights regional and inter-specific differences. The mean respiration rates of all mesopelagic fish sampled were 593.6 and 354.9 ml O2 individual-1 h-1 in the Scotia Sea and Benguela Current, respectively. Global allometric models performed poorly in colder regions compared with our observations, underestimating respiratory flux in the Scotia Sea by 67–88%. This may reflect that most data used to fit such models are derived from temperate and subtropical regions. We recommend caution when applying globally derived allometric models to regional data, particularly in cold (

Blooms of harmful algae are increasing globally, yet their impacts on copepods, an important link between primary producers and higher trophic levels, remain largely unknown. Algal toxins may have direct, negative effects on the survival of copepods. They may also indirectly affect copepod survival by deterring feeding and thus decreasing the availability of energy and nutritional resources. Here we present a series of short-term (24 h) experiments in which the cosmopolitan marine copepod, Acartia tonsa, was exposed to a range of concentrations of the toxic dinoflagellate, Alexandrium catenella (strain 1119/27, formerly Alexandrium tamarense), with and without the presence of alternative, non-toxic prey (Rhodomonas sp.). We also present the toxin profile concentrations for A. catenella. The survival and feeding of A. tonsa were not affected across the range of concentrations recorded for A. catenella in the field; increased mortality of A. tonsa was only discernible when A. catenella was present at concentrations that exceed their reported environmental concentrations by two orders of magnitude. The observed lethal median concentration (LC50) for A. tonsa exposed to A. catenella was 12.45 ng STX eq L-1. We demonstrate that A. tonsa is capable of simultaneously ingesting both toxic and non-toxic algae, but increases clearance rates towards non-toxic prey as the proportional abundance of toxic A. catenella increases. The ability to actively select non-toxic algae whilst also ingesting toxic algae suggests that consumption of the latter does not cause physical incapacitation and thus does not affect ingestion in A. tonsa. This work shows that short-term exposure to toxic A. catenella is unlikely to elicit major effects on the grazing or survival of A. tonsa. However, more work is needed to understand the longer-term and sub-lethal effects of toxic algae on marine copepods.

We applied the concept of source-sink dynamics to investigate a recent (1999-2013) increase in the occurrence of the siphonophore Muggiaea atlantica in Scottish coastal waters. Our aim was to determine whether this change represented the establishment of resident populations (i.e. "sources"), or transient populations reliant on immigration (i.e. "sinks"). First, we show that local production was not always sufficient to account for recruitment (a "source" prerequisite), suggesting reliance on immigration (a "sink" prerequisite). Using variation partitioning, we then discriminated between the exclusive effects of immigration [indexed by the European Slope Current (ESC)] and local production (indexed by local sea temperature and food availability). On the west coast (Loch Ewe), interannual variability in the species' abundance was determined by, in order of increasing importance: (i) suitable local environmental conditions (13%); (ii) the role of the ESC in modulating these conditions (20%); and (iii) immigration via the ESC (29%). These results provided a strong indication that Loch Ewe represents a sink habitat for M. atlantica. However, on the east coast (Stonehaven) our results were less conclusive, probably due to the less direct influence of the ESC. For both locations, we suggest that low winter temperatures prevented overwintering, necessitating annual re-colonization via immigration.

Marine copepods are central to the productivity and biogeochemistry of marine ecosystems. Nevertheless, the direct and indirect effects of climate change on their metabolic functioning remain poorly understood. Here, we use metabolomics, the unbiased study of multiple low molecular weight organic metabolites, to examine how the physiology of Calanus spp. is affected by end-of-century global warming and ocean acidification scenarios. We report that the physiological stresses associated with incubation without food over a 5-day period greatly exceed those caused directly by seawater temperature or pH perturbations. This highlights the need to contextualise the results of climate change experiments by comparison to other, naturally occurring stressors such as food deprivation, which is being exacerbated by global warming. Protein and lipid metabolism were up-regulated in the food-deprived animals, with a novel class of taurine-containing lipids and the essential polyunsaturated fatty acids (PUFAs), eicosapentaenoic acid and docosahexaenoic acid, changing significantly over the duration of our experiment. Copepods derive these PUFAs by ingesting diatoms and flagellated microplankton respectively. Climate-driven changes in the productivity, phenology and composition of microplankton communities, and hence the availability of these fatty acids, therefore have the potential to influence the ability of copepods to survive starvation and other environmental stressors.

Photosynthesis in the surface ocean produces approximately 100 gigatonnes of organic carbon per year, of which 5 to 15 per cent is exported to the deep ocean. The rate at which the sinking carbon is converted into carbon dioxide by heterotrophic organisms at depth is important in controlling oceanic carbon storage. It remains uncertain, however, to what extent surface ocean carbon supply meets the demand of water-column biota; the discrepancy between known carbon sources and sinks is as much as two orders of magnitude. Here we present field measurements, respiration rate estimates and a steady-state model that allow us to balance carbon sources and sinks to within observational uncertainties at the Porcupine Abyssal Plain site in the eastern North Atlantic Ocean. We find that prokaryotes are responsible for 70 to 92 per cent of the estimated remineralization in the twilight zone (depths of 50 to 1,000 metres) despite the fact that much of the organic carbon is exported in the form of large, fast-sinking particles accessible to larger zooplankton. We suggest that this occurs because zooplankton fragment and ingest half of the fast-sinking particles, of which more than 30 per cent may be released as suspended and slowly sinking matter, stimulating the deep-ocean microbial loop. The synergy between microbes and zooplankton in the twilight zone is important to our understanding of the processes controlling the oceanic carbon sink.

The performance of a new mesozooplankton sampler, the U-Tow, was compared to that of the traditional WP2 net. The U-Tow significantly underestimated species abundance, but gave a very good representation of species composition and community size structure. WP2 net samples could be used to calibrate the U-Tow, allowing absolute abundance to be determined. It is recommended that the U- Tow, in its current configuration, be used in conjunction with WP2 net samples to give measures of abundance, or as a tool to identify areas of change in plankton communities.