Temporal changes in zooplankton indicators highlight a bottom-up process in the Bay of Marseille (NW Mediterranean Sea).

Sixteen years (2005-2020) of zooplankton monitoring in the Bay of Marseille (N-W Mediterranean Sea) are analyzed in relation to physical, meteorological, climatic and biotic data. Samples were collected every two weeks by a vertical haul (0-55 m) of a 200 µm plankton net. Different indices characterizing the mesozooplankton are compared: biomass dry weight of four size fractions between 200 and 2000 µm; abundances of the whole of the mesozooplankton and of 13 main taxonomic groups defined from plankton imagery; seasonal onset timing of each zooplankton group; and two other types of indices: the first characterized diversity based on abundance data, and the second was derived from zooplankton size spectra shape. The clearest pattern in the environmental compartment was an overall decreasing trend in nutrients, shifts in phytoplankton metrics (i.e. size structure and particulate organic matter), and changes in winter conditions (i.e. increasing temperatures, precipitation and NAO). Interannual patterns in the mesozooplankton community were: (i) a decrease of total abundance (ii) a decrease in biomass for the four size fractions, with an earlier decrease for the 1000-2000 µm size fraction (in 2008); (iii) a reduced dominance of copepods (calanoids and oithonoids) and a concomitant increase in abundance of other taxonomic groups (crustaceans, pteropods, chaetognaths, salps) which induced higher diversity; (iv) a first shift in size spectra towards smaller sizes in 2009, when the 1000-2000 µm size fraction biomass decreased, and a second shift towards larger sizes in 2013 along with increased diversity; and (iv) a later onset in the phenology for some zooplankton variables and earlier onset for salps. Concomitant changes in the phytoplankton compartment, winter environmental conditions, zooplankton community structure (in size and diversity) and zooplankton phenology marked by a shift in 2013 suggest bottom-up control of the pelagic ecosystem.

Isotopic and biochemical trophic markers reveal the complexity of interactions at the base of pelagic food webs (Mediterranean sea).

To gain insight into the impact of bottom-up changes in the plankton community on planktivorous fish in the context of the decline of small pelagic fisheries in the Northwestern Mediterranean Sea, we have conducted an extensive year-long study. The investigation combined biochemical analyses (proteins, carbohydrates, and lipids) with C and N stable isotope analyses (SIA) to simultaneously study phytoplankton, zooplankton, and eight planktivorous fish species (Engraulis encrasicolus, Sardina pilchardus, Sardinella aurita, Sprattus sprattus, Cepola macrophthalma, Chromis chromis, Boops boops, and Spicara maena). This study is the first to analyze both stable isotope and biochemical compositions in coastal particulate organic matter (POM) size classes (0.7-2.7 µm, 2.7-20 µm, and 20-200 µm), zooplankton size classes (200-300 µm, 300-500 µm, 500-1000 µm, 1000-2000 µm, and >2000 µm), and taxonomic groups. We demonstrated that: (1) POM stable isotope compositions varied based on its spatial origin, the taxonomic composition of its biota, and its biochemical content; (2) δ15N values increased with zooplankton size classes and groups, indicating different trophic levels; (3) Phytoplankton exhibited a lipid-rich composition (∼55%), while zooplankton and fish muscles were protein-rich (∼61% and ∼66%, respectively). Bayesian stable isotope mixing models revealed that, on average: (1) POM from oceanic waters contributed the most to the POM in the bay (>51%), with a dominance of pico-POM (∼43%); (2) The 200-1000 µm zooplankton primarily consumed nano-POM, the 1000-2000 µm zooplankton mostly consumed micro-POM (∼64%), and the >2000 µm zooplankton also mostly consumed micro-POM; (3) Mesozooplankton (200-2000 µm) constituted the main portion (∼42%) of the diet for planktivorous fish species, while macrozooplankton organisms (>2000 µm) were the primary food resource (∼43%) for both B. boops and S. sprattus. Our study underscores the complexity of the pelagic food web and highlights the bottom-up transfer of organic matter from the smallest phytoplankton size fractions to planktivorous fish.

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.

Bioconcentration, bioaccumulation and biomagnification of mercury in plankton of the Mediterranean Sea.

Plankton plays a prominent role in the bioaccumulation of mercury (Hg). The MERITE-HIPPOCAMPE campaign was carried out in spring 2019 along a north-south transect including coastal and offshore areas of the Mediterranean Sea. Sampling of sea water and plankton by pumping and nets was carried out in the chlorophyll maximum layer. Two size-fractions of phytoplankton (0.7-2.7 and 2.7-20 µm) and five of zooplankton (between 60 and >2000 µm) were separated, and their total mercury (THg) and monomethylmercury (MMHg) contents were measured. Bioconcentration of THg was significantly higher in the smallest phytoplankton size-fraction dominated by Synechococcus spp. The bioaccumulation and biomagnification of MMHg in zooplankton was influenced by size, food sources, biochemical composition and trophic level. MMHg was biomagnified in the plankton food web, while THg decreased toward higher trophic levels. Higher MMHg concentrations were measured in oligotrophic areas. Plankton communities in the Southern Mediterranean Sea had lower MMHg concentrations than those in the Northern Mediterranean Sea. These results highlighted the influence of environmental conditions and trophodynamics on the transfer of Hg in Mediterranean plankton food webs, with implications for higher trophic level consumers.

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.

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.

Trace elements, dioxins and PCBs in different fish species and marine regions: Importance of the taxon and regional features.

Chemical contaminant concentrations in wild organisms are used to assess environmental status under the European Marine Strategy Framework Directive. However, this approach is challenged by the complex intra- and inter-species variability, and the different regional features. In this study, concentrations in trace elements (As, Cd, Hg and Pb), polychlorinated biphenyls (PCBs), polychlorodibenzo-para-dioxines (PCDDs) and polychlorodibenzofuranes (PCDFs) were monitored in 8 fish species sampled on the continental shelf of three French regions: the Eastern English Channel (EEC) and Bay of Biscay (BoB) in the Northeast Atlantic Ocean, and the Gulf of Lions (GoL) in Western Mediterranean Sea. Our objectives were to identify species or regions more likely to be contaminated and to assess how to take this variability into account in environmental assessment. While concentrations were higher in benthic and demersal piscivores, PCB and PCDD/F concentrations (lipid-weight) were similar in most teleost species. For Cd, Hg and Pb, the trophic group accumulating the highest concentrations depended on the contaminant and region. Concentrations in Hg, PCBs and PCDD/Fs were higher in the EEC and/or GoL than in BoB. Cadmium and Pb concentrations were highest in the BoB. Lipid content accounted for 35%-84% of organic contaminant variability. Lipid normalisation was employed to enhance robustness in the identification of spatial patterns. Contaminant patterns in chondrichthyans clearly differed from that in teleosts. In addition, trophic levels accounted for ≤1% and ≤33% of the contaminant variability in teleost fishes in the EEC and BoB, respectively. Therefore, developing taxa-specific thresholds might be a more practical way forward for environmental assessment than normalisation to trophic levels.

Mediterranean Mercury Assessment 2022: An Updated Budget, Health Consequences, and Research Perspectives.

Mercury (Hg) and especially its methylated species (MeHg) are toxic chemicals that contaminate humans via the consumption of seafood. The most recent UNEP Global Mercury Assessment stressed that Mediterranean populations have higher Hg levels than people elsewhere in Europe. The present Critical Review updates current knowledge on the sources, biogeochemical cycling, and mass balance of Hg in the Mediterranean and identifies perspectives for future research especially in the context of global change. Concentrations of Hg in the Western Mediterranean average 0.86 ± 0.27 pmol L-1 in the upper water layer and 1.02 ± 0.12 pmol L-1 in intermediate and deep waters. In the Eastern Mediterranean, Hg measurements are in the same range but are too few to determine any consistent oceanographical pattern. The Mediterranean waters have a high methylation capacity, with MeHg representing up to 86% of the total Hg, and constitute a source of MeHg for the adjacent North Atlantic Ocean. The highest MeHg concentrations are associated with low oxygen water masses, suggesting a microbiological control on Hg methylation, consistent with the identification of hgcA-like genes in Mediterranean waters. MeHg concentrations are twice as high in the waters of the Western Basin compared to the ultra-oligotrophic Eastern Basin waters. This difference appears to be transferred through the food webs and the Hg content in predators to be ultimately controlled by MeHg concentrations of the waters of their foraging zones. Many Mediterranean top-predatory fish still exceed European Union regulatory Hg thresholds. This emphasizes the necessity of monitoring the exposure of Mediterranean populations, to formulate adequate mitigation strategies and recommendations, without advising against seafood consumption. This review also points out other insufficiencies of knowledge of Hg cycling in the Mediterranean Sea, including temporal variations in air-sea exchange, hydrothermal and cold seep inputs, point sources, submarine groundwater discharge, and exchanges between margins and the open sea. Future assessment of global change impacts under the Minamata Convention Hg policy requires long-term observations and dedicated high-resolution Earth System Models for the Mediterranean region.

A study of trophic structure, physiological condition and mercury biomagnification in swordfish (Xiphias gladius): Evidence of unfavourable conditions for the swordfish population in the Western Mediterranean.

Studies integrating trophic ecology, physiological condition and accumulation of heavy metals in top predators, such as swordfish, are needed to better understand the links between them and the risk to humans associated with consumption of these fish. This research focuses on the swordfish of the Catalan Sea and follows a multi method approach that considers their diet, their liver lipid content, and mercury accumulation in their bodies as well as in their prey. The aim is to highlight the links between trophic ecology, physiology (fish condition), and eco-toxicology. Results indicate that poor condition of swordfish based on size and the levels of lipid in the liver, and the high Hg levels accumulated to the trophic web (particularly from cephalopods) may indicate potential unfavourable feeding and reproduction conditions for swordfish in the NW Mediterranean and this warrants further investigation.

Systematics of European coastal anchovies (genus Engraulis Cuvier).

Reports of morphological differences between European anchovy (Engraulis cf. encrasicolus) from coastal and marine habitats have long existed in the ichthyologic literature and have given rise to a long-standing debate on their taxonomic status. More recently, molecular studies have confirmed the existence of genetic differentiation between the two anchovy ecotypes. Using ancestry-informative markers, we show that coastal anchovies throughout the Mediterranean share a common ancestry and that substantial genetic differentiation persists in different pairs of coastal/marine populations despite the presence of limited gene flow. On the basis of genetic and ecological arguments, we propose that coastal anchovies deserve a species status of their own (E. maeoticus) and argue that a unified taxonomical framework is critical for future research and management.

Multi-trophic markers illuminate the understanding of the functioning of a remote, low coral cover Marquesan coral reef food web.

We studied the food web structure and functioning of a coral reef ecosystem in the Marquesas Islands, French Polynesia, characterized by low coral cover, high sea surface temperature and meso- to eutrophic waters. The Marquesas constitute a relevant ecosystem to understand the functioning of low diversity reefs that are also subject to global change. A multi-tracer assessment of organic matter pathways was run to delineate ecosystem functioning, using analysis of fatty acids, bulk and compound specific stable isotope analysis and stable isotopes mixing models. Macroalgae and phytoplankton were the two major food sources fueling this food web with, however, some marked seasonal variations. Specifically, zooplankton relied on phytoplankton-derived organic matter and herbivorous fishes on macroalgae-derived organic matter to a much higher extent in summer than in winter (~ 75% vs. ~ 15%, and ~ 70 to 75% vs. ~ 5 to 15%, respectively) . Despite remarkably high δ15N values for all trophic compartments, likely due to local dynamics in the nitrogen stock, trophic levels of consumers were similar to those of other coral reef ecosystems. These findings shed light on the functioning of low coral cover systems, which are expected to expand worldwide under global change.

Persistent Organic Pollutants Burden, Trophic Magnification and Risk in a Pelagic Food Web from Coastal NW Mediterranean Sea.

The storage capacity, trophic magnification and risk of sixty-two POPs have been evaluated in a well-characterized pelagic food web (including phytoplankton, zooplankton, six fish, and two cephalopods species) from an impacted area in NW Mediterranean Sea. Our results show the high capacity of the planktonic compartment for the storage of polybrominated diphenyl ethers (PBDEs) and polychlorinated dibenzo-p-dioxins and dibenzofurans (PCDD/Fs), consistent with their estimated low trophic magnification factors (TMF) of 0.2-2.0 (PBDEs) and of 0.3-1.1 (PCDD/Fs). ∑PBDEs dominated in the zooplankton size-class 200-1000 µm (∼330 ng g-1 lw, median), whereas ∑PCDD/Fs accumulated preferentially in phytoplankton size-class 0.7-200 µm (875 pg g-1 lw, median). In contrast, polychlorinated biphenyls (PCBs) were preferentially bioaccumulated in the higher trophic levels (six fish species and two cephalopods) with TMFs = 0.8-3.9, reaching median concentrations of 4270 and 3140 ng g-1 lw (∑PCBs) in Atlantic bonito (Sarda sarda) and chub mackerel (Scomber colias), respectively. For these edible species, the estimated weekly intakes of dioxin-like POPs for humans based on national consumption standards overpassed the EU tolerable weekly intake. Moreover, the concentrations of nondioxin-like PCBs in S. sarda were above the EU maximum levels in foodstuffs, pointing to a risk. No risk evidence was found due to consumption of all other edible species studied, neither for PBDEs. The integrated burden of POPs in the food web reached ∼18 µg g-1 lw, representing a dynamic stock of toxic organic chemicals in the study area. We show that the characterized food web could be a useful and comprehensive "bioindicator" of the chemical pollution status of the study area, opening new perspectives for the monitoring of toxic chemicals in Mediterranean coastal waters.

Unexpected spatial impact of treatment plant discharges induced by episodic hydrodynamic events: Modelling Lagrangian transport of fine particles by Northern Current intrusions in the bays of Marseille (France).

Our study highlights the Lagrangian transport of solid particles discharged at the Marseille Wastewater Treatment Plant (WWTP), located at Cortiou on the southern coastline. We focused on episodic situations characterized by a coastal circulation pattern induced by intrusion events of the Northern Current (NC) on the continental shelf, associated with SE wind regimes. We computed, using MARS3D-RHOMA and ICHTHYOP models, the particle trajectories from a patch of 5.104 passive and conservative fine particles released at the WWTP outlet, during 2 chosen representative periods of intrusion of the NC in June 2008 and in October 2011, associated with S-SE and E-SE winds, respectively. Unexpected results highlighted that the amount of particles reaching the vulnerable shorelines of both northern and southern bays accounted for 21.2% and 46.3% of the WWTP initial patch, in June 2008 and October 2011, respectively. Finally, a conceptual diagram is proposed to highlight the mechanisms of dispersion within the bays of Marseille of the fine particles released at the WWTP outlet that have long been underestimated.

Unexpected headless and tailless fish in the stomach content of shortfin mako Isurus oxyrinchus.

The stomach content of 113 individuals of shortfin mako Isurus oxyrinchus was analyzed. Individuals were sampled at landing in Vigo (Spain) and captured by sea-surface long-liners in the vicinity of the Azores Archipelago and between Azores and the Iberian Peninsula, in March and October 2012, and March 2013. Teleosts constituted the dominant item, mainly Atlantic saury Scomberesox saurus (87% of teleost prey). Among them, 94% were deprived of both head and the caudal fin, while the flesh and bones of the body were preserved. The presence of eye's lenses, the number of which was consistent with the number of fish remains, likely rules out the elimination of the heads before ingestion. There is no obvious explanation for this unexpected and unrecorded pattern of digestion.

Man induced change in community control in the north-western Black Sea: The top-down bottom-up balance.

The present study shows how marine commercial fish food webs dramatically changed in the north-western Black Sea on both pelagic and benthic environments. Fisheries landings, diversity and equitability strongly decreased between 1965-1970 and 2001-2005. Fishes adapted their feeding behaviour to the increasingly low species diversity of the Black Sea communities. Their food web became poor and simplified following the loss of many top predator species and their trophic links. Linkage density, connectivity and Lyapunov stability proxy strongly decreased. The north-western Black Sea system switched from a complex top-down and bottom-up functioning pattern to a dominantly bottom-up functioning pattern. This study contributes to a better understanding of these transformations within the Danube-Black Sea system in the last decades. An attempt is made to relate these changes with river inputs, fisheries and coastal pollution.

Influence of the Danube River inputs on C and N stable isotope ratios of the Romanian coastal waters and sediment (Black Sea).

The main aim of this study was to estimate the influence of the seasonal variations of the Danube River particulate organic matter (POM) inputs on the Black Sea surface seawater POM and upper layer of sediments along the Romanian coast. Ratios of carbon ((13)C/(12)C) and nitrogen ((15)N/(14)N) stable isotopes allowed differentiating river and marine organic matter sources. Danube River POM presented significantly lower average values of delta(13)C (-27.52+/-0.88 per thousand) and delta(15)N (4.88+/-1.45 per thousand) than seawater POM (delta(13)C=-24.70+/-2.37 per thousand and delta(15)N=6.75+/-1.96 per thousand), whereas surface sediment presented average values similar to seawater POM (delta(13)C=-24.02+/-2.39 per thousand and delta(15)N=7.29+/-2.16 per thousand). Stable isotope values showed that the Danube River influence on marine ecosystems decreased from the North to the South of the Romanian coast. Strong seasonal variations of C and N isotopic signatures were observed in all compartments studied with generally higher values in spring when the river was flooding.