Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. “Gut flux” for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.

Ecological Investigations with the Continuous Plankton Recorder: The Zooplankton (other than Copepoda and Young Fish) in the Southern North Sea, 1932-37.

I. The report describes the main monthly changes in the distribution and abundance of the zooplankton, other than Copepoda and young fish (dealt with in separate reports), over the southern part of the North Sea from 1932 to 1937. The work is part of the survey carried out by Continuous Plankton Recorders towed at a depth of 10 metres on regular steamship lines between England and the Continent. 2. The limitations to the sampling method are discussed, and it is shown to be unsuitable for recording Mysidacea and Euphausiacea on account of their marked diurnal variation due presumably to vertical migration; they are omitted from the report. 3. The changing distribution of Sagitta, Limacina, Clione, Lamellibranch larvae, Cladocera, Caprellid Amphipoda, Decapod larvae, Echinoderm larvae and Oikopleura are shown in a series of monthly charts while their seasonal fluctuations are compared in time-chart histograms. 4. The Alima larvae of Squilla are recorded on a few occasions in the regions where the Channel opens into the North Sea. 5. The distributional characteristics of the different forms, i.e. their tendencies to even or " patchy " production, are compared.

The Impact of Polystyrene Microplastics on Feeding, Function and Fecundity in the Marine CopepodCalanus helgolandicus

Microscopic plastic debris, termed “microplastics”, are of increasing environmental concern. Recent studies have demonstrated that a range of zooplankton, including copepods, can ingest microplastics. Copepods are a globally abundant class of zooplankton that form a key trophic link between primary producers and higher trophic marine organisms. Here we demonstrate that ingestion of microplastics can significantly alter the feeding capacity of the pelagic copepod Calanus helgolandicus. Exposed to 20 μm polystyrene beads (75 microplastics mL–1) and cultured algae ([250 μg C L–1) for 24 h, C. helgolandicus ingested 11% fewer algal cells (P = 0.33) and 40% less carbon biomass (P < 0.01). There was a net downward shift in the mean size of algal prey consumed (P < 0.001), with a 3.6 fold increase in ingestion rate for the smallest size class of algal prey (11.6–12.6 μm), suggestive of postcapture or postingestion rejection. Prolonged exposure to polystyrene microplastics significantly decreased reproductive output, but there were no significant differences in egg production rates, respiration or survival. We constructed a conceptual energetic (carbon) budget showing that microplastic-exposed copepods suffer energetic depletion over time. We conclude that microplastics impede feeding in copepods, which over time could lead to sustained reductions in ingested carbon biomass.

Next Generation Sequencing Reveals the Hidden Diversity of Zooplankton Assemblages

Zooplankton play an important role in our oceans, in biogeochemical cycling and providing a food source for commercially important fish larvae. However, difficulties in correctly identifying zooplankton hinder our understanding of their roles in marine ecosystem functioning, and can prevent detection of long term changes in their community structure. The advent of massively parallel next generation sequencing technology allows DNA sequence data to be recovered directly from whole community samples. Here we assess the ability of such sequencing to quantify richness and diversity of a mixed zooplankton assemblage from a productive time series site in the Western English Channel. Methodology/Principle Findings Plankton net hauls (200 µm) were taken at the Western Channel Observatory station L4 in September 2010 and January 2011. These samples were analysed by microscopy and metagenetic analysis of the 18S nuclear small subunit ribosomal RNA gene using the 454 pyrosequencing platform. Following quality control a total of 419,041 sequences were obtained for all samples. The sequences clustered into 205 operational taxonomic units using a 97% similarity cut-off. Allocation of taxonomy by comparison with the National Centre for Biotechnology Information database identified 135 OTUs to species level, 11 to genus level and 1 to order, <2.5% of sequences were classified as unknowns. By comparison a skilled microscopic analyst was able to routinely enumerate only 58 taxonomic groups. Conclusions Metagenetics reveals a previously hidden taxonomic richness, especially for Copepoda and hard-to-identify meroplankton such as Bivalvia, Gastropoda and Polychaeta. It also reveals rare species and parasites. We conclude that Next Generation Sequencing of 18S amplicons is a powerful tool for elucidating the true diversity and species richness of zooplankton communities. While this approach allows for broad diversity assessments of plankton it may become increasingly attractive in future if sequence reference libraries of accurately identified individuals are better populated.

Contrasting transcriptome response to thermal stress in two key zooplankton species, Calanus finmarchicus and C. glacialis

Climate change has already led to the range expansion of warm-water plankton assemblages in the northeast Atlantic and the corresponding range contraction of colder-water species. The temperate copepod Calanus finmarchicus is predicted to shift farther northward into polar waters traditionally dominated by the arctic copepod C. glacialis. To identify temperaturemediated changes in gene expression that may be critical for the thermal acclimation and resilience of the 2 Calanus spp., we conducted a whole transcriptome profiling using RNA-seq on an Ion Torrent platform. Transcriptome responses of C. finmarchicus and C. glacialis from Disko Bay, west Greenland, were investigated under realistic thermal stresses (at + 5, +10 and +15°C) for 4 h and 6 d. C. finmarchicus showed a strong response to temperature and duration of stress, involving up-regulation of genes related to protein folding, transcription, translation and metabolism. In sharp contrast, C. glacialis displayed only low-magnitude changes in gene expression in response to temperature and duration of stress. Differences in the thermal responses of the 2 species, particularly the lack of thermal stress response in C. glacialis, are in line with laboratory and field observations and suggest a vulnerability of C. glacialis to climate change.

Plastic and marine turtles: a review and call for research

Plastic debris is now ubiquitous in the marine environment affecting a wide range of taxa, from microscopic zooplankton to large vertebrates. Its persistence and dispersal throughout marine ecosystems has meant that sensitivity toward the scale of threat is growing, particularly for species of conservation concern, such as marine turtles. Their use of a variety of habitats, migratory behaviour, and complex life histories leave them subject to a host of anthropogenic stressors, including exposure to marine plastic pollution. Here, we review the evidence for the effects of plastic debris on turtles and their habitats, highlight knowledge gaps, and make recommendations for future research. We found that, of the seven species, all are known to ingest or become entangled in marine debris. Ingestion can cause intestinal blockage and internal injury, dietary dilution, malnutrition, and increased buoyancy which in turn can result in poor health, reduced growth rates and reproductive output, or death. Entanglement in plastic debris (including ghost fishing gear) is known to cause lacerations, increased drag—which reduces the ability to forage effectively or escape threats—and may lead to drowning or death by starvation. In addition, plastic pollution may impact key turtle habitats. In particular, its presence on nesting beaches may alter nest properties by affecting temperature and sediment permeability. This could influence hatchling sex ratios and reproductive success, resulting in population level implications. Additionally, beach litter may entangle nesting females or emerging hatchlings. Lastly, as an omnipresent and widespread pollutant, plastic debris may cause wider ecosystem effects which result in loss of productivity and implications for trophic interactions. By compiling and presenting this evidence, we demonstrate that urgent action is required to better understand this issue and its effects on marine turtles, so that appropriate and effective mitigation policies can be developed.

Island-coastal and oceanic epipelagic zooplankton biodiversity in the southwestern Indian Ocean

The oceanic Indian Ocean zooplankton species and their distributions have been well described, but the zooplankton of coastal regions, particularly around the oceanic islands, has not been well researched, either taxonomically or experimentally. The environment of the Mascarene region in the southwestern Indian Ocean and zooplankton research that has been carried out there is detailed, along with gaps in our knowledge. Suggestions are given for future research, particularly on the zooplankton species adapted to live in the fluctuating environment of inshore waters, including studies on taxonomy and biodiversity, life cycles, dispersion and genetics. Problems of carrying out taxonomic research are highlighted.

Role of zooplankton dynamics for Southern Ocean phytoplankton biomass and global biogeochemical cycles

Global ocean biogeochemistry models currently employed in climate change projections use highly simplified representations of pelagic food webs. These food webs do not necessarily include critical pathways by which ecosystems interact with ocean biogeochemistry and climate. Here we present a global biogeochemical model which incorporates ecosystem dynamics based on the representation of ten plankton functional types (PFTs); six types of phytoplankton, three types of zooplankton, and heterotrophic bacteria. We improved the representation of zooplankton dynamics in our model through (a) the explicit inclusion of large, slow-growing zooplankton, and (b) the introduction of trophic cascades among the three zooplankton types. We use the model to quantitatively assess the relative roles of iron vs. grazing in determining phytoplankton biomass in the Southern Ocean High Nutrient Low Chlorophyll (HNLC) region during summer. When model simulations do not represent crustacean macrozooplankton grazing, they systematically overestimate Southern Ocean chlorophyll biomass during the summer, even when there was no iron deposition from dust. When model simulations included the developments of the zooplankton component, the simulation of phytoplankton biomass improved and the high chlorophyll summer bias in the Southern Ocean HNLC region largely disappeared. Our model results suggest that the observed low phytoplankton biomass in the Southern Ocean during summer is primarily explained by the dynamics of the Southern Ocean zooplankton community rather than iron limitation. This result has implications for the representation of global biogeochemical cycles in models as zooplankton faecal pellets sink rapidly and partly control the carbon export to the intermediate and deep ocean.

Microplastics Alter the Properties and Sinking Rates of Zooplankton Faecal Pellets

Plastic debris is a widespread contaminant, prevalent in aquatic ecosystems across the globe. Zooplankton readily ingest microscopic plastic (microplastic, < 1 mm), which are later egested within their faecal pellets. These pellets are a source of food for marine organisms, and contribute to the oceanic vertical flux of particulate organic matter as part of the biological pump. The effects of microplastics on faecal pellet properties are currently unknown. Here we test the hypotheses that (1) faecal pellets are a vector for transport of microplastics, (2) polystyrene microplastics can alter the properties and sinking rates of zooplankton egests and, (3) faecal pellets can facilitate the transfer of plastics to coprophagous biota. Following exposure to 20.6 μm polystyrene microplastics (1000 microplastics mL–1) and natural prey (∼1650 algae mL–1) the copepod Calanus helgolandicus egested faecal pellets with significantly (P < 0.001) reduced densities, a 2.25-fold reduction in sinking rates, and a higher propensity for fragmentation. We further show that microplastics, encapsulated within egests of the copepod Centropages typicus, could be transferred to C. helgolandicus via coprophagy. Our results support the proposal that sinking faecal matter represents a mechanism by which floating plastics can be vertically transported away from surface waters.

Modelling the future biogeography of North Atlantic zooplankton communities in response to climate change

Advances in habitat and climate modelling allow us to reduce uncertainties of climate change impacts on species distribution. We evaluated the impacts of future climate change on community structure, diversity, distribution and phenology of 14 copepod species in the North Atlantic. We developed and validated habitat models for key zooplankton species using continuous plankton recorder (CPR) survey data collected at mid latitudes of the North Atlantic. Generalized additive models (GAMs) were applied to relate the occurrence of species to environmental variables. Models were projected to future (2080–2099) environmental conditions using coupled hydroclimatix–biogeochemical models under the Intergovernmental Panel on Climate Change (IPCC) A1B climate scenario, and compared to present (2001–2020) conditions. Our projections indicated that the copepod community is expected to respond substantially to climate change: a mean poleward latitudinal shift of 8.7 km per decade for the overall community with an important species range variation (–15 to 18 km per decade); the species seasonal peak is expected to occur 12–13 d earlier for Calanus finmarchicus and C. hyperboreus; and important changes in community structure are also expected (high species turnover of 43–79% south of the Oceanic Polar Front). The impacts of the change expected by the end of the century under IPCC global warming scenarios on copepods highlight poleward shifts, earlier seasonal peak and changes in biodiversity spatial patterns that might lead to alterations of the future North Atlantic pelagic ecosystem. Our model and projections are supported by a temporal validation undertaken using the North Atlantic climate regime shift that occurred in the 1980s: the habitat model built in the cold period (1970–1986) has been validated in the warm period (1987–2004).

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.