Zooplankton biomass, size structure, and associated metabolic fluxes with focus on its roles at the chlorophyll maximum layer during the plankton-contaminant MERITE-HIPPOCAMPE cruise.

Recent studies have demonstrated that plankton can be a key pathway for the uptake and transfer of contaminants entering the marine environment up to top predators. The plankton-contaminant MERITE-HIPPOCAMPE cruise was devoted to quantifying contaminants in water and the whole plankton size range (10 size fractions) at 10 stations along a north-south transect in the western Mediterranean Sea from the French to the Tunisian coasts through the Provençal and Algerian basins. Pumping and filtering devices and net sampling have been used for collecting very high amounts of small particles and planktonic organisms in the chlorophyll maximum layer (CML). The present paper characterizes the zooplankton components for which the contaminant measurements were carried out. At each station, a horizontal towed Hydro-Bios net with a 60 µm mesh-size net was used to discriminate 5 size-fractions from 60 µm to a few mm. For each size-fraction, one part of the sample was used for dry weight measurements and the other one for estimating the contribution to biomass of detritus, phytoplankton, and among zooplankton of the major taxonomic groups based on the imagery tools ZOOSCAN and FLOWCAM. In each zooplankton size fraction, metabolic rates were calculated from the size spectrum to estimate trophic and excretion fluxes flowing through this fraction. These observations were compared to a similar analysis of tows in the upper layer (vertical) and the surface layer (horizontal). The total sampled biomass concentration at the CML was higher than in the water column (COL) and much higher than at the surface (SURF) in most of the stations, but in the CML and COL a substantial contribution was due to detritus mostly concentrated in the smallest size-fractions (60-200 µm and 200-500 µm). Absolute values of zooplankton biomass show neither a clear spatial pattern nor a significant difference between strata. The CML layer was dominated by copepods similarly to COL and SURF, but presented a higher contribution of nauplii and a near absence of appendicularians. At some stations, crustaceans and gelatinous plankton could be important contributors to CML. The zooplankton biomass composition of the two smallest fractions (<500 µm) was dominated by nauplii, small copepods and, occasionally, by small miscellaneous organisms (mostly pteropodes). In contrast, clear differences between stations appeared for the largest fractions (>500 µm) due to large crustaceans, gelatinous organisms, and chaetognaths. These changes in biomass composition according to size fractions suggest a progressive trophic shift from dominant herbivory in the smallest fractions to more contrasted trophic structure (including carnivory) in the largest fractions. The daily carbon demand and the N and P excretion of zooplankton were on average higher at the CML but with no significant difference with COL. The zooplankton grazing represented 2.7 to 22.7 % of the phytoplankton stock per day, whereas its excretion represented a daily N and P recycling compared to dissolved inorganic nitrogen and phosphorus stocks ranging respectively from 0.2 to 19 % and from 0 to 21 %. This information should help in the interpretation of the content of various contaminants in zooplankton fractions.

Investigation of dynamic change in microplastics vertical distribution patterns: The seasonal effect on vertical distribution.

This paper analyzes the variability of microplastics vertical distributions in the oceanic water column. Data were obtained from targeted sampling in the Bay of Marseille (France) and from a numerical simulation forced by realistic physical forcings. By fitting model and in-situ data in a simplified vertical dimension, three microplastics classes may be deduced: settling, buoyant and winter neutrally-buoyant microplastics. Buoyant microplastics are mainly concentrated at the surface but they can be mixed throughout the whole water column during episodes with strong winds and no water stratification, inducing an implicit underestimation of buoyant microplastics in surface sampling. Almost symmetrical to the distribution of buoyant microplastics, settling microplastics are mainly found at the bottom but they can sometimes reach the surface under the mixing conditions cited above. They could thus contribute to surface sampling. Winter neutrally-buoyant microplastics are more homogenously mixed during the winter but are under the stratified layers during summer.

Microplastics in the maximum chlorophyll layer along a north-south transect in the Mediterranean Sea in comparison with zooplankton concentrations.

The aim of this study was to characterize and quantify microplastics (MPs) at the chlorophyll maximum layer (CML), around 30 to 60 m depth, during a cruise dedicated to the study of contaminants in plankton, the MERITE-HIPPOCAMPE project, along a north-south transect in the western Mediterranean Sea (Tedetti et al., 2023). Plankton were collected by horizontal net tows in this layer using a multinet Hydrobios Midi equipped with 60 µm mesh-size nets. The collected plankton were fractionated through a sieve column for various later contaminant measurements and for zooplankton analysis (Fierro-González et al., 2023). For all stations, samples were also fully examined for microplastics (MPs) for fractions >300 µm. MPs were found at all stations in the CML layer (mean: 42.9 ± 45.4 MPs m-3), of which 96 ± 4 % were fibers. The ratios of mesozooplankton/MPs and detritus/MPs in this CML were respectively 223 ± 315 and 2544 ± 2268. These data are analyzed together with MPs concentrations from sea- surface sampled with a 300 µm net-size Manta net at the same stations. Overall, our observations highlight the very high density of fibers at the CML, mainly associated with aggregates, raising the hypothesis of their interactions with marine snow. Therefore, the importance of marine snow and vertical layering will have to be considered in future MP distribution modelling efforts.

Spatial variations of biochemical content and stable isotope ratios of size-fractionated plankton in the Mediterranean Sea (MERITE-HIPPOCAMPE campaign).

Plankton represents the main source of carbon in marine ecosystems and is consequently an important gateway for contaminants into the marine food webs. During the MERITE- HIPPOCAMPE campaign in the Mediterranean Sea (April-May 2019), plankton was sampled from pumping and net tows at 10 stations from the French coast to the Gulf of Gabès (Tunisia) to obtain different size fractions in contrasted regions. This study combines various approaches, including biochemical analyses, analyses of stable isotope ratios (δ13C, δ15N), cytometry analyses and mixing models (MixSiar) on size-fractions of phyto- and zooplankton from 0.7 to >2000 µm. Pico- and nanoplankton represented a large energetic resource at the base of pelagic food webs. Proteins, lipids, and stable isotope ratios increased with size in zooplankton and were higher than in phytoplankton. Stable isotope ratios suggest different sources of carbon and nutrients at the base of the planktonic food webs depending on the coast and the offshore area. In addition, a link between productivity and trophic pathways was shown, with high trophic levels and low zooplankton biomass recorded in the offshore area. The results of our study highlight spatial variations of the trophic structure within the plankton size-fractions and will contribute to assess the role of the plankton as a biological pump of contaminants.

Hydrocarbons in size-fractionated plankton of the Mediterranean Sea (MERITE-HIPPOCAMPE campaign).

Aliphatic and polycyclic aromatic hydrocarbons (AHs and PAHs, respectively) were analyzed in the dissolved fraction (<0.7 µm) of surface water and in various particulate/planktonic size fractions (0.7-60, 60-200, 200-500 and 500-1000 µm) collected at the deep chlorophyll maximum, along a North-South transect in the Mediterranean Sea in spring 2019 (MERITE-HIPPOCAMPE campaign). Suspended particulate matter, biomass, total chlorophyll a, particulate organic carbon, C and N isotopic ratios, and lipid biomarkers were also determined to help characterizing the size-fractionated plankton and highlight the potential link with the content in AHs and PAHs in these size fractions. Æ©28AH concentrations ranged 18-489 ng L-1 for water, 3.9-72 µg g-1 dry weight (dw) for the size fraction 0.7-60 µm, and 3.4-55 µg g-1 dw for the fractions 60-200, 200-500 and 500-1000 µm. AH molecular profiles revealed that they were mainly of biogenic origin. Æ©14PAH concentrations were 0.9-16 ng L-1 for water, and Æ©27PAH concentrations were 53-220 ng g-1 dw for the fraction 0.7-60 µm and 35-255 ng g-1 dw for the three higher fractions, phenanthrene being the most abundant compound in planktonic compartment. Two processes were evidenced concerning the PAH patterns, the bioreduction, i.e., the decrease in concentrations from the small size fractions (0.7-60 and 60-200 µm) to the higher ones (200-500 µm and 500-1000 µm), and the biodilution, i.e., the decrease in concentrations in plankton at higher suspended matter or biomass, especially for the 0.7-60 and 60-200-µm size fractions. We estimated the biological pump fluxes of Æ©27PAHs below 100-m depth in the Western Mediterranean Sea at 15 ± 10 ng m-2 day-1, which is comparable to those previously reported in the South Pacific and Indian Ocean.

Contamination of planktonic food webs in the Mediterranean Sea: Setting the frame for the MERITE-HIPPOCAMPE oceanographic cruise (spring 2019).

This paper looks at experiential feedback and the technical and scientific challenges tied to the MERITE-HIPPOCAMPE cruise that took place in the Mediterranean Sea in spring 2019. This cruise proposes an innovative approach to investigate the accumulation and transfer of inorganic and organic contaminants within the planktonic food webs. We present detailed information on how the cruise worked, including 1) the cruise track and sampling stations, 2) the overall strategy, based mainly on the collection of plankton, suspended particles and water at the deep chlorophyll maximum, and the separation of these particles and planktonic organisms into various size fractions, as well as the collection of atmospheric deposition, 3) the operations performed and material used at each station, and 4) the sequence of operations and main parameters analysed. The paper also provides the main environmental conditions that were prevailing during the campaign. Lastly, we present the types of articles produced based on work completed by the cruise that are part of this special issue.

Do microbial planktonic communities reflect the ecological changes of Glorieuses coral reefs (Iles Eparses, Western Indian Ocean)?

Ecological baselines for the structure and functioning of ecosystems in the absence of human activity can provide essential information on their health status. The Glorieuses islands are located in the Western Indian Ocean (WIO) and can be considered as "pristine" ecosystems that have not been subjected to anthropogenic pressure. Their nutrient context and the microbial assemblages were assessed by determining the abundance of heterotrophic prokaryotes (archaea and bacteria), picocyanobacteria, picoeukaryotes, microphytoplankton and protozooplankton communities in five stations, during two contrasted periods (November 2015 and May 2016). Chlorophyll-a concentrations were always under 1 µg/L and associated to very low levels in orthophosphates, nitrate and dissolved organic carbon, revealing an ultra-oligotrophic status for the Glorieuses waters. Picocyanobacteria confirmed the ultra-oligotrophic status with a predominance of Synechococcus. Zeaxanthin associated with the presence of picocyanobacteria represented the major pigment in both surveys. Three indices of diversity (species richness, Shannon and Pielou indexes) from microscopy observations highlighted the difference of diversity in microphytoplankton between the surveys. A focus on a 16S metabarcoding approach showed a high dominance of picocyanobacteria, Alpha- and Gammaproteobacteria, regardless of station or period. Multivariate analyses (co-inertia analyses) revealed a strong variability of ecological conditions between the two periods, with (i) high nutrient concentrations and heterotrophic nanoflagellate abundance in November 2015, and (ii) high heterotrophic prokaryote and picoeukaryote abundance in May 2016. The impact of a category 5 tropical cyclone (Fantala) on the regional zone in April 2016 is also advanced to explain these contrasted situations. Relative importance of top-down factors between bacterial and heterotrophic nanoflagellates was observed in November 2015 with an active microbial food web. All the results indicate that three microbial indexes potentially can be considered to assess the ecological change in Glorieuses marine waters.

Responses of planktonic microorganisms from tropical reservoirs to paraquat and deltamethrin exposure.

This study focused on the effects of two pesticides, paraquat (herbicide) and deltamethrin (insecticide), which are two common molecules used intensively in Burkina Faso. Natural bacterial populations, phytoplankton cultures (one cyanobacterium, Cylindrospermopsis raciborskii, and one chlorophycea, Monoraphidium sp.), and two species of zooplankton (Diaphanosoma excisum and Moina micrura) were isolated from aquatic communities and were used as biological targets in the experimental protocols. Paraquat was moderately toxic to bacteria and phytoplankton, whereas deltamethrin was significantly toxic only to the zooplankton species. Paradoxically, the chlorophycea Monoraphidium sp. exhibited a significant increase of in vitro fluorescence after 48 h at the highest doses. Preliminary tests were also performed from natural water extract of the main drinking water supply of the country (Loumbila Reservoir) by using solid-phase extraction. Obviously, the natural extract proved to be toxic to the same biological targets. Despite the absence of any determination of pesticides in the natural extract, the question of contamination and toxicity of these waters affects concerns about the safety of water supply and the effect of human pressure on the dynamics of planktonic communities of freshwater reservoirs in arid regions of western Africa.

Trophic relationships between metazooplankton communities and their plankton food sources in the Iles Eparses (Western Indian Ocean).

Coral reef and atoll lagoons are among the most diversified marine ecosystems but also the most affected by the combined effects of climate change and human activities. The Iles Eparses (Scattered Islands) in the Western Indian Ocean have been little affected by human pressure and can be considered to be "pristine" ecosystems. Metazooplankton plays a major role in the functioning and productivity of aquatic ecosystems, and this study was undertaken: (i) to determine the spatial abundance, distribution and species composition of metazooplankton, (ii) to assess the effect of metazooplankton grazing on pico- and nanophytoplankton and (iii) to analyze the trophic positions of metazooplankton by using the stable isotope signatures of a wide variety of taxa and particulate organic matter from the Iles Eparses and Mayotte. Tromelin Island (which is not located in the Mozambique Channel) had the lowest metazooplankton abundance with no cyanobacteria Trichodesmium spp. or mollusks (pteropods) presence, and with δ(15)N signatures of organisms that were higher than for the islands in the Mozambique Channel. Trichodesmium spp. was found in the Mozambique Channel and the plankton food web was probably based preferentially on these cyanobacteria with lower δ(15)N signatures indicating direct or indirect trophic transfer of diazotrophic nitrogen to metazooplankton. Three of the islands were distinct: Europa had the highest proportion of copepods, with oithonids being dominant, which is typical of rich mangrove systems, while Juan de Nova and Mayotte seemed to be the sites most affected by human activity with a high abundance of appendicularians and distinct particulate organic matter ∂(13)C signatures. Grazing experiments showed that food could be a limiting factor for metazooplankton in the Iles Eparses. However, the effect of metazooplankton grazing on phytoplankton appeared to be very low (0.01-2.32% of the total phytoplankton per day).

Identifying the drivers of abundance and size of the invasive ctenophore Mnemiopsis leidyi in Northwestern Mediterranean lagoons.

Acknowledged as among the worst invasive marine species, Mnemiopsis leidyi has spread through European Seas since the mid-1980's. Here we report a bimonthly survey conducted in 2010-11 in three lagoons (Bages-Sigean, Thau and Berre) and at two adjacent coastal stations (Sète and SOMLIT-Marseille) along the French Mediterranean coast. M. leidyi was present only in Berre and Bages-Sigean with maximum abundances observed in late summer. M. leidyi adults were present year round in Berre with the largest organisms (∼6 cm) observed in April. In Bages-Sigean, they occurred in sufficient abundance to be recorded by fishermen between August and November. Multiple linear regressions highlighted that abundance in both lagoons was mainly influenced by direct effects of salinity and chlorophyll-a, and temperature to a lesser extent. While M. leidyi has not yet been recorded in Thau, the lagoon is continually monitored to detect the potential establishment of M. leidyi.

Copepod feeding in a tuna fishery area of the tropical Atlantic Ocean.

Biomass, feeding and metabolic rates of planktonic copepods were studied in an oligotrophic area of the tropical Atlantic Ocean during an instability wave period (boreal summer) and a stratified period (boreal winter). In summer, zooplankton biomass was higher than in winter, showing a positive effect of the instability wave. Moreover, feeding equilibrated metabolic expenditures of copepods in most cases during the instability. In contrast, in stratified conditions copepods did not equilibrate their metabolic budget. Our results suggest that the microbial loop was the dominant trophic pathway during both periods but with a quicker cycling during the instability.

Effects of the sandbar breaching on hydrobiological parameters and zooplankton communities in the Senegal River Estuary (West Africa).

This study describes the changes in hydrology, zooplankton communities and abundance in the Senegal River Estuary (SRE) before and after the breaching of the sandbar in October 2003. Samples were taken in 2003 at 3 stations located upstream (DI), in mid estuary (HY) and downstream (RM), and in 2005 at the same stations (RM becoming Old River Mouth: ORM), plus the new river mouth (NRM) resulting from the morphological evolution of the SRE. The study showed marked seasonal variations that affected the structure and distribution of zooplankton as well as major changes caused by the sandbar opening: increased marine influence throughout the whole SRE, changes in the horizontal gradients, arrival of euryhaline species and increase in meroplankton, in particular decapod larvae, transformation of the ORM area into a slackwater area with limited exchanges and the highest zooplankton numbers during high waters.

Do human activities affect the picoplankton structure of the Ahe atoll lagoon (Tuamotu Archipelago, French Polynesia)?

The spatial variations of the picoplankton (photoautotrophic and heterotrophic microorganisms) in the Ahe atoll lagoon were studied in May and October 2008 to assess whether they were affected by human activities along the atoll. Spatial patterns were studied using 10 sampling stations chosen according to the location of the anthropogenic activities (pearl farming, harbor). Experiments were also carried out to determine whether bacterial growth, with or without predators, was limited by inorganic (N and P) substrates. The results showed that heterotrophic bacterioplankton abundance was superior to the photoautotrophic organisms, especially in May. Significant increases in bacterial abundance were observed in May after 24 h incubation with +P and +N (but not in October). All samples complied with the quality levels for fecal indicator bacteria (FIB) defined by the European Union and there was no evidence that human sewage had any impact on picoplankton over the whole atoll.

Man-induced hydrological changes, metazooplankton communities and invasive species in the Berre Lagoon (Mediterranean Sea, France).

The Berre Lagoon has been under strong anthropogenic pressure since the early 1950s. The opening of the hydroelectric EDF power plant in 1966 led to large salinity drops. The zooplankton community was mainly composed of two common brackish species: Acartia tonsa and Brachionus plicatilis. Since 2006, European litigation has strongly constrained the input of freshwater, maintaining the salinity above 15. A study was performed between 2008 and 2010 to evaluate how these modifications have impacted the zooplankton community. Our results show that the community is more diverse and contains several coastal marine species (i.e., Centropages typicus, Paracalanus parvus and Acartia clausi). A. tonsa is still present but is less abundant, whereas B. plicatilis has completely disappeared. Strong predatory marine species, such as chaetognaths, the large conspicuous autochtonous jellyfish Aurelia aurita and the invasive ctenophore Mnemiopsis leidyi, are now very common as either seasonal or permanent features of the lagoon.

Metazooplankton communities in the Ahe atoll lagoon (Tuamotu Archipelago, French Polynesia): spatiotemporal variations and trophic relationships.

Metazooplankton abundance, biomass (<80 µm, 200-500 µm and >500 µm) and community structure in the Ahe atoll were studied together with their relationships with environmental factors (temperature, salinity, wind) and trophic factors (phytoplankton, bacteria, heterotrophic nanoflagellates (HNF) and ciliates) during three periods in 2008-2009. Meroplankton, mainly bivalve and gastropod larvae, was dominant. Holoplankton was dominated by copepods, the main species being Oithona spp., Paracalanus parvus, Clausocalanus spp., Corycaeus spp., Acartia fossae and Undinula vulgaris. The results suggest a clear wind influence on the structure and horizontal distribution of the zooplankton communities. The metazooplankton appeared to be controlled mainly by food resources, suggesting a bottom-up control. The low nanophytoplankton biomass in contrast to the high abundance of picophytoplankton, HNF and nano-particle grazers (mainly Oithona spp., Paracalanus and bivalve larvae) highlighted the importance of the microbial loop in the food web.