Articles | Volume 16, issue 22
https://doi.org/10.5194/gmd-16-6701-2023
© Author(s) 2023. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/gmd-16-6701-2023
© Author(s) 2023. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Model description paper
| 
21 Nov 2023
Model description paper |
| 21 Nov 2023
| 21 Nov 2023
Implementation and assessment of a model including mixotrophs and the carbonate cycle (Eco3M_MIX-CarbOx v1.0) in a highly dynamic Mediterranean coastal environment (Bay of Marseille, France) – Part 1: Evolution of ecosystem composition under limited light and nutrient conditions
Lucille Barré
CORRESPONDING AUTHOR
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
Frédéric Diaz
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
deceased
Thibaut Wagener
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
France Van Wambeke
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
Camille Mazoyer
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
Christophe Yohia
Aix Marseille Univ., Université de Toulon, CNRS, IRD, OSU Institut Pythéas, 13288 Marseille, France
Christel Pinazo
CORRESPONDING AUTHOR
Aix Marseille Univ., Université de Toulon, CNRS, IRD, MIO, UM 110, 13288 Marseille, France
Related authors
Lucille Barré, Frédéric Diaz, Thibaut Wagener, Camille Mazoyer, Christophe Yohia, and Christel Pinazo
Geosci. Model Dev., 17, 5851–5882, https://doi.org/10.5194/gmd-17-5851-2024, https://doi.org/10.5194/gmd-17-5851-2024, 2024
Short summary
Short summary
The carbonate system is typically studied using measurements, but modeling can contribute valuable insights. Using a biogeochemical model, we propose a new representation of total alkalinity, dissolved inorganic carbon, pCO2, and pH in a highly dynamic Mediterranean coastal area, the Bay of Marseille, a useful addition to measurements. Through a detailed analysis of pCO2 and air–sea CO2 fluxes, we show that variations are strongly impacted by the hydrodynamic processes that affect the bay.
Gaëlle Capitaine, Samir Alliouane, Thierry Cariou, Jonathan Fin, Paola Fisicaro, and Thibaut Wagener
EGUsphere, https://doi.org/10.5194/egusphere-2025-3588, https://doi.org/10.5194/egusphere-2025-3588, 2025
Short summary
Short summary
Measuring total alkalinity in seawater is essential for understanding and monitoring the ocean carbonate system. To improve the reliability of these measurements, we developed reference materials and tested them in an inter-laboratory comparison. We also thoroughly quantified, for the first time, the uncertainty of the standard measurement method. These results, as well as the key metrological tools developed, support more accurate long-term monitoring of the ocean carbonate system.
Nicolas Metzl, Jonathan Fin, Claire Lo Monaco, Claude Mignon, Samir Alliouane, Bruno Bombled, Jacqueline Boutin, Yann Bozec, Steeve Comeau, Pascal Conan, Laurent Coppola, Pascale Cuet, Eva Ferreira, Jean-Pierre Gattuso, Frédéric Gazeau, Catherine Goyet, Emilie Grossteffan, Bruno Lansard, Dominique Lefèvre, Nathalie Lefèvre, Coraline Leseurre, Sébastien Petton, Mireille Pujo-Pay, Christophe Rabouille, Gilles Reverdin, Céline Ridame, Peggy Rimmelin-Maury, Jean-François Ternon, Franck Touratier, Aline Tribollet, Thibaut Wagener, and Cathy Wimart-Rousseau
Earth Syst. Sci. Data, 17, 1075–1100, https://doi.org/10.5194/essd-17-1075-2025, https://doi.org/10.5194/essd-17-1075-2025, 2025
Short summary
Short summary
This work presents a new synthesis of 67 000 total alkalinity and total dissolved inorganic carbon observations obtained between 1993 and 2023 in the global ocean, coastal zones, and the Mediterranean Sea. We describe the data assemblage and associated quality control and discuss some potential uses of this dataset. The dataset is provided in a single format and includes the quality flag for each sample.
Lucille Barré, Frédéric Diaz, Thibaut Wagener, Camille Mazoyer, Christophe Yohia, and Christel Pinazo
Geosci. Model Dev., 17, 5851–5882, https://doi.org/10.5194/gmd-17-5851-2024, https://doi.org/10.5194/gmd-17-5851-2024, 2024
Short summary
Short summary
The carbonate system is typically studied using measurements, but modeling can contribute valuable insights. Using a biogeochemical model, we propose a new representation of total alkalinity, dissolved inorganic carbon, pCO2, and pH in a highly dynamic Mediterranean coastal area, the Bay of Marseille, a useful addition to measurements. Through a detailed analysis of pCO2 and air–sea CO2 fluxes, we show that variations are strongly impacted by the hydrodynamic processes that affect the bay.
France Van Wambeke, Pascal Conan, Mireille Pujo-Pay, Vincent Taillandier, Olivier Crispi, Alexandra Pavlidou, Sandra Nunige, Morgane Didry, Christophe Salmeron, and Elvira Pulido-Villena
Biogeosciences, 21, 2621–2640, https://doi.org/10.5194/bg-21-2621-2024, https://doi.org/10.5194/bg-21-2621-2024, 2024
Short summary
Short summary
Phosphomonoesterase (PME) and phosphodiesterase (PDE) activities over the epipelagic zone are described in the eastern Mediterranean Sea in winter and autumn. The types of concentration kinetics obtained for PDE (saturation at 50 µM, high Km, high turnover times) compared to those of PME (saturation at 1 µM, low Km, low turnover times) are discussed in regard to the possible inequal distribution of PDE and PME in the size continuum of organic material and accessibility to phosphodiesters.
Nicolas Metzl, Jonathan Fin, Claire Lo Monaco, Claude Mignon, Samir Alliouane, David Antoine, Guillaume Bourdin, Jacqueline Boutin, Yann Bozec, Pascal Conan, Laurent Coppola, Frédéric Diaz, Eric Douville, Xavier Durrieu de Madron, Jean-Pierre Gattuso, Frédéric Gazeau, Melek Golbol, Bruno Lansard, Dominique Lefèvre, Nathalie Lefèvre, Fabien Lombard, Férial Louanchi, Liliane Merlivat, Léa Olivier, Anne Petrenko, Sébastien Petton, Mireille Pujo-Pay, Christophe Rabouille, Gilles Reverdin, Céline Ridame, Aline Tribollet, Vincenzo Vellucci, Thibaut Wagener, and Cathy Wimart-Rousseau
Earth Syst. Sci. Data, 16, 89–120, https://doi.org/10.5194/essd-16-89-2024, https://doi.org/10.5194/essd-16-89-2024, 2024
Short summary
Short summary
This work presents a synthesis of 44 000 total alkalinity and dissolved inorganic carbon observations obtained between 1993 and 2022 in the Global Ocean and the Mediterranean Sea at the surface and in the water column. Seawater samples were measured using the same method and calibrated with international Certified Reference Material. We describe the data assemblage, quality control and some potential uses of this dataset.
Stéphanie Barrillon, Robin Fuchs, Anne A. Petrenko, Caroline Comby, Anthony Bosse, Christophe Yohia, Jean-Luc Fuda, Nagib Bhairy, Frédéric Cyr, Andrea M. Doglioli, Gérald Grégori, Roxane Tzortzis, Francesco d'Ovidio, and Melilotus Thyssen
Biogeosciences, 20, 141–161, https://doi.org/10.5194/bg-20-141-2023, https://doi.org/10.5194/bg-20-141-2023, 2023
Short summary
Short summary
Extreme weather events can have a major impact on ocean physics and biogeochemistry, but their study is challenging. In May 2019, an intense storm occurred in the north-western Mediterranean Sea, during which in situ multi-platform measurements were performed. The results show a strong impact on the surface phytoplankton, highlighting the need for high-resolution measurements coupling physics and biology during these violent events that may become more common in the context of global change.
Julie Dinasquet, Estelle Bigeard, Frédéric Gazeau, Farooq Azam, Cécile Guieu, Emilio Marañón, Céline Ridame, France Van Wambeke, Ingrid Obernosterer, and Anne-Claire Baudoux
Biogeosciences, 19, 1303–1319, https://doi.org/10.5194/bg-19-1303-2022, https://doi.org/10.5194/bg-19-1303-2022, 2022
Short summary
Short summary
Saharan dust deposition of nutrients and trace metals is crucial to microbes in the Mediterranean Sea. Here, we tested the response of microbial and viral communities to simulated dust deposition under present and future conditions of temperature and pH. Overall, the effect of the deposition was dependent on the initial microbial assemblage, and future conditions will intensify microbial responses. We observed effects on trophic interactions, cascading all the way down to viral processes.
Céline Ridame, Julie Dinasquet, Søren Hallstrøm, Estelle Bigeard, Lasse Riemann, France Van Wambeke, Matthieu Bressac, Elvira Pulido-Villena, Vincent Taillandier, Fréderic Gazeau, Antonio Tovar-Sanchez, Anne-Claire Baudoux, and Cécile Guieu
Biogeosciences, 19, 415–435, https://doi.org/10.5194/bg-19-415-2022, https://doi.org/10.5194/bg-19-415-2022, 2022
Short summary
Short summary
We show that in the Mediterranean Sea spatial variability in N2 fixation is related to the diazotrophic community composition reflecting different nutrient requirements among species. Nutrient supply by Saharan dust is of great importance to diazotrophs, as shown by the strong stimulation of N2 fixation after a simulated dust event under present and future climate conditions; the magnitude of stimulation depends on the degree of limitation related to the diazotrophic community composition.
Matthieu Bressac, Thibaut Wagener, Nathalie Leblond, Antonio Tovar-Sánchez, Céline Ridame, Vincent Taillandier, Samuel Albani, Sophie Guasco, Aurélie Dufour, Stéphanie H. M. Jacquet, François Dulac, Karine Desboeufs, and Cécile Guieu
Biogeosciences, 18, 6435–6453, https://doi.org/10.5194/bg-18-6435-2021, https://doi.org/10.5194/bg-18-6435-2021, 2021
Short summary
Short summary
Phytoplankton growth is limited by the availability of iron in about 50 % of the ocean. Atmospheric deposition of desert dust represents a key source of iron. Here, we present direct observations of dust deposition in the Mediterranean Sea. A key finding is that the input of iron from dust primarily occurred in the deep ocean, while previous studies mainly focused on the ocean surface. This new insight will enable us to better represent controls on global marine productivity in models.
Stéphanie H. M. Jacquet, Christian Tamburini, Marc Garel, Aurélie Dufour, France Van Vambeke, Frédéric A. C. Le Moigne, Nagib Bhairy, and Sophie Guasco
Biogeosciences, 18, 5891–5902, https://doi.org/10.5194/bg-18-5891-2021, https://doi.org/10.5194/bg-18-5891-2021, 2021
Short summary
Short summary
We compared carbon remineralization rates (MRs) in the western and central Mediterranean Sea in late spring during the PEACETIME cruise, as assessed using the barium tracer. We reported higher and deeper (up to 1000 m depth) MRs in the western basin, potentially sustained by an additional particle export event driven by deep convection. The central basin is the site of a mosaic of blooming and non-blooming water masses and showed lower MRs that were restricted to the upper mesopelagic layer.
Elvira Pulido-Villena, Karine Desboeufs, Kahina Djaoudi, France Van Wambeke, Stéphanie Barrillon, Andrea Doglioli, Anne Petrenko, Vincent Taillandier, Franck Fu, Tiphanie Gaillard, Sophie Guasco, Sandra Nunige, Sylvain Triquet, and Cécile Guieu
Biogeosciences, 18, 5871–5889, https://doi.org/10.5194/bg-18-5871-2021, https://doi.org/10.5194/bg-18-5871-2021, 2021
Short summary
Short summary
We report on phosphorus dynamics in the surface layer of the Mediterranean Sea. Highly sensitive phosphate measurements revealed vertical gradients above the phosphacline. The relative contribution of diapycnal fluxes to total external supply of phosphate to the mixed layer decreased towards the east, where atmospheric deposition dominated. Taken together, external sources of phosphate contributed little to total supply, which was mainly sustained by enzymatic hydrolysis of organic phosphorus.
France Van Wambeke, Vincent Taillandier, Karine Desboeufs, Elvira Pulido-Villena, Julie Dinasquet, Anja Engel, Emilio Marañón, Céline Ridame, and Cécile Guieu
Biogeosciences, 18, 5699–5717, https://doi.org/10.5194/bg-18-5699-2021, https://doi.org/10.5194/bg-18-5699-2021, 2021
Short summary
Short summary
Simultaneous in situ measurements of (dry and wet) atmospheric deposition and biogeochemical stocks and fluxes in the sunlit waters of the open Mediterranean Sea revealed complex physical and biological processes occurring within the mixed layer. Nitrogen (N) budgets were computed to compare the sources and sinks of N in the mixed layer. The transitory effect observed after a wet dust deposition impacted the microbial food web down to the deep chlorophyll maximum.
Frédéric Gazeau, France Van Wambeke, Emilio Marañón, Maria Pérez-Lorenzo, Samir Alliouane, Christian Stolpe, Thierry Blasco, Nathalie Leblond, Birthe Zäncker, Anja Engel, Barbara Marie, Julie Dinasquet, and Cécile Guieu
Biogeosciences, 18, 5423–5446, https://doi.org/10.5194/bg-18-5423-2021, https://doi.org/10.5194/bg-18-5423-2021, 2021
Short summary
Short summary
Our study shows that the impact of dust deposition on primary production depends on the initial composition and metabolic state of the tested community and is constrained by the amount of nutrients added, to sustain both the fast response of heterotrophic prokaryotes and the delayed one of phytoplankton. Under future environmental conditions, heterotrophic metabolism will be more impacted than primary production, therefore reducing the capacity of surface waters to sequester anthropogenic CO2.
Frédéric Gazeau, Céline Ridame, France Van Wambeke, Samir Alliouane, Christian Stolpe, Jean-Olivier Irisson, Sophie Marro, Jean-Michel Grisoni, Guillaume De Liège, Sandra Nunige, Kahina Djaoudi, Elvira Pulido-Villena, Julie Dinasquet, Ingrid Obernosterer, Philippe Catala, and Cécile Guieu
Biogeosciences, 18, 5011–5034, https://doi.org/10.5194/bg-18-5011-2021, https://doi.org/10.5194/bg-18-5011-2021, 2021
Short summary
Short summary
This paper shows that the impacts of Saharan dust deposition in different Mediterranean basins are as strong as those observed in coastal waters but differed substantially between the three tested stations, differences attributed to variable initial metabolic states. A stronger impact of warming and acidification on mineralization suggests a decreased capacity of Mediterranean surface communities to sequester CO2 following the deposition of atmospheric particles in the coming decades.
Evelyn Freney, Karine Sellegri, Alessia Nicosia, Leah R. Williams, Matteo Rinaldi, Jonathan T. Trueblood, André S. H. Prévôt, Melilotus Thyssen, Gérald Grégori, Nils Haëntjens, Julie Dinasquet, Ingrid Obernosterer, France Van Wambeke, Anja Engel, Birthe Zäncker, Karine Desboeufs, Eija Asmi, Hilkka Timonen, and Cécile Guieu
Atmos. Chem. Phys., 21, 10625–10641, https://doi.org/10.5194/acp-21-10625-2021, https://doi.org/10.5194/acp-21-10625-2021, 2021
Short summary
Short summary
In this work, we present observations of the organic aerosol content in primary sea spray aerosols (SSAs) continuously generated along a 5-week cruise in the Mediterranean. This information is combined with seawater biogeochemical properties also measured continuously along the ship track to develop a number of parametrizations that can be used in models to determine SSA organic content in oligotrophic waters that represent 60 % of the oceans from commonly measured seawater variables.
Matthieu Roy-Barman, Lorna Foliot, Eric Douville, Nathalie Leblond, Fréderic Gazeau, Matthieu Bressac, Thibaut Wagener, Céline Ridame, Karine Desboeufs, and Cécile Guieu
Biogeosciences, 18, 2663–2678, https://doi.org/10.5194/bg-18-2663-2021, https://doi.org/10.5194/bg-18-2663-2021, 2021
Short summary
Short summary
The release of insoluble elements such as aluminum (Al), iron (Fe), rare earth elements (REEs), thorium (Th) and protactinium (Pa) when Saharan dust falls over the Mediterranean Sea was studied during tank experiments under present and future climate conditions. Each element exhibited different dissolution kinetics and dissolution fractions (always lower than a few percent). Changes in temperature and/or pH under greenhouse conditions lead to a lower Th release and a higher light REE release.
France Van Wambeke, Elvira Pulido, Philippe Catala, Julie Dinasquet, Kahina Djaoudi, Anja Engel, Marc Garel, Sophie Guasco, Barbara Marie, Sandra Nunige, Vincent Taillandier, Birthe Zäncker, and Christian Tamburini
Biogeosciences, 18, 2301–2323, https://doi.org/10.5194/bg-18-2301-2021, https://doi.org/10.5194/bg-18-2301-2021, 2021
Short summary
Short summary
Michaelis–Menten kinetics were determined for alkaline phosphatase, aminopeptidase and β-glucosidase in the Mediterranean Sea. Although the ectoenzymatic-hydrolysis contribution to heterotrophic prokaryotic needs was high in terms of N, it was low in terms of C. This study points out the biases in interpretation of the relative differences in activities among the three tested enzymes in regard to the choice of added concentrations of fluorogenic substrates.
Emilio Marañón, France Van Wambeke, Julia Uitz, Emmanuel S. Boss, Céline Dimier, Julie Dinasquet, Anja Engel, Nils Haëntjens, María Pérez-Lorenzo, Vincent Taillandier, and Birthe Zäncker
Biogeosciences, 18, 1749–1767, https://doi.org/10.5194/bg-18-1749-2021, https://doi.org/10.5194/bg-18-1749-2021, 2021
Short summary
Short summary
The concentration of chlorophyll is commonly used as an indicator of the abundance of photosynthetic plankton (phytoplankton) in lakes and oceans. Our study investigates why a deep chlorophyll maximum, located near the bottom of the upper, illuminated layer develops in the Mediterranean Sea. We find that the acclimation of cells to low light is the main mechanism involved and that this deep maximum represents also a maximum in the biomass and carbon fixation activity of phytoplankton.
Katixa Lajaunie-Salla, Frédéric Diaz, Cathy Wimart-Rousseau, Thibaut Wagener, Dominique Lefèvre, Christophe Yohia, Irène Xueref-Remy, Brian Nathan, Alexandre Armengaud, and Christel Pinazo
Geosci. Model Dev., 14, 295–321, https://doi.org/10.5194/gmd-14-295-2021, https://doi.org/10.5194/gmd-14-295-2021, 2021
Short summary
Short summary
A biogeochemical model of planktonic food webs including a carbonate balance module is applied in the Bay of Marseille (France) to represent the carbon marine cycle expected to change in the future owing to significant increases in anthropogenic emissions of CO2. The model correctly simulates the ranges and seasonal dynamics of most variables of the carbonate system (pH). This study shows that external physical forcings have an important impact on the carbonate equilibrium in this coastal area.
Kahina Djaoudi, France Van Wambeke, Aude Barani, Nagib Bhairy, Servanne Chevaillier, Karine Desboeufs, Sandra Nunige, Mohamed Labiadh, Thierry Henry des Tureaux, Dominique Lefèvre, Amel Nouara, Christos Panagiotopoulos, Marc Tedetti, and Elvira Pulido-Villena
Biogeosciences, 17, 6271–6285, https://doi.org/10.5194/bg-17-6271-2020, https://doi.org/10.5194/bg-17-6271-2020, 2020
Cited articles
Agawin, N. S. R., Duarte, C. M., and Agusti, S.: Nutrient and temperature control of the contribution of picoplankton to phytoplankton biomass and production, Limnol. Oceanogr., 45, 591–600, https://doi.org/10.4319/lo.2000.45.3.0591, 2000.
Allen, J. I., Holt, J. T., Blackford, J., and Proctor, R.: Error quantification of a high-resolution coupled hydrodynamic-ecosystem coastal ocean model: Part 2. Chlorophyll-a, nutrients, and SPM, J. Mar. Syst., 68, 381–404, https://doi.org/10.1016/j-jmarsys.2007.01.005, 2007.
Auger, P. A., Diaz, F., Ulses, C., Estournel, C., Neveux, J., Joux, F., Pujo-Pay, M., and Naudin, J. J.: Functioning of the planktonic ecosystem on the Gulf of Lions shelf (NW Mediterranean) during spring and its impact on the carbon deposition: a field data and 3-D modelling combined approach, Biogeosciences, 8, 3231–3261, https://doi.org/10.5194/bg-8-3231-2011, 2011.
Baklouti, M., Diaz, F., Pinazo, C., Faure, V., and Queguiner, B.: Investigation of mechanistic formulations depicting phytoplankton dynamics for models of marine pelagic ecosystems and description of a new model, Prog. Oceanogr., 71, 1–33, https://doi.org/10.1016/j.pocean.2006.05.002, 2006a.
Baklouti, M., Faure, V., Pawlowski, L., and Sciandra, A.: Investigation and sensitivity analysis of a mechanistic phytoplankton model implemented in a new modular numerical tool (Eco3M) dedicated to biogeochemical modelling, Prog. Oceanogr., 71, 34–58, https://doi.org/10.1016/j.pocean.2006.05.003, 2006b.
Banaru, D., Diaz, F., Verley, P., Campbell, R., Navarro, J., Yohia, C., Oliveros-Ramos, R., Mellon-Duval, C., and Shin, Y. J.: Implementation of an end-to-end model of the Gulf of Lions ecosystem (NW Mediterranean Sea). I. Parametrization, calibration and evaluation, Ecol. Model., 401, 1–19, https://doi.org/10.1016/j.ecolmodel.2019.03.005, 2019.
Barré, L., Diaz, F., Wagener, T., Van Wambeke, F., Mazoyer, C., Yohia, C., and Pinazo, C.: Eco3M_MIX-CarbOx (v1.0), Zenodo [code], https://doi.org/10.5281/zenodo.7669658, 2022.
Barré, L., Diaz, F., Wagener, T., Mazoyer, C., Yohia, C., and Pinazo, C.: Implementation and assessment of a model including mixotrophs and the carbonate cycle (Eco3M_MIX-CarbOx v1.0) in a highly dynamic Mediterranean coastal environment (Bay of Marseille, France) (Part. II): Towards a better representation of total alkalinity when modelling the carbonate system and air-sea CO2 fluxes, Geosci. Model Dev. Discuss. [preprint], https://doi.org/10.5194/gmd-2023-34, in review, 2023.
Barrier, N., Petrenko, A. A., and Ourmières, Y.: Strong intrusions of the Northern Mediterranean Current on the eastern Gulf of Lion: insights from in-situ observations and high-resolution numerical modelling, Ocean Dynam., 66, 313–327, https://doi.org/10.1007/s10236-016-0921-7, 2016.
Bernard, C. and Rassoulzadegan, F.: Seasonal variations of mixotrophic ciliates in the northwest Mediterranean Sea, Mar. Ecol. Prog. Ser., 108, 295–301, 1994.
Burkholder, J. M., Glibert, P. M., and Skelton, H. M.: Mixotrophy, a major mode of nutrition for harmful algal species in eutrophic waters, Harmful Algae, 8, 77–93, https://doi.org/10.1016/j.hal.2008.08.010, 2008.
Campbell, R., Diaz, F., Hu, Z., Doglioli, A., Petrenko, A., and Dekeyser, I.: Nutrients and plankton spatial distributions induced by a coastal eddy in the Gulf of Lion. Insights from a numerical model, Prog. Oceanogr., 109, 47–69, https://doi.org/10.1016/j.pocean.2012.09.005, 2013.
Christaki, U., Van Wambeke, F., and Dolan, J. R.: Nanoflagellates (mixotrophs, heterotrophs and autotrophs) in the oligotrophic eastern Mediterranean: standing stocks, bacterivory and relationships with bacterial production, Mar. Ecol. Prog. Ser., 181, 297–307, 1999.
Christaki, U., Courties, C., Karayanni, H., Giannakourou, A., Maravelias, C., Kormas, K., and Lebaron, P.: Dynamic characteristics of Prochlorococcus and Synechococcus consumption by bacterivorous nanoflagellates, Microb. Ecol., 43, 341–352, 2002.
Christaki, U., Courties, C., Joux, F., Jeffrey, W. H., Neveux, J., and Naudin, J. J.: Community structure and trophic role of ciliates and heterotrophic nanoflagellates in Rhone River diluted mesoscale structures (NW Mediterranean Sea), Aquat. Microb. Ecol., 57, 263–277, https://doi.org/10.3354/ame01339, 2009.
Djaoudi, K., Van Wanbeke, F., Barani, A., Nunige, S. H., Sempere, R., and Pulido-Vilena, E.: Atmospheric fluxes of soluble organic C, N, and P to the Mediterranean Sea: Potential biogeochemical implications in the surface layer, Prog. Oceanogr., 163, 59–69, https://doi.org/10.1016/j.pocean.2017.07.008, 2018.
Dolan, J. R.: Mixotrophy in ciliates: a review of Chlorella symbiosis and chloroplast retention, Marine Microbial Food Webs, 6, 115–132, 1992.
Dolan, J. R. and Perez, M. T.: Costs, benefits, and characteristics of mixotrophy in marine oligotrichs, Freshwater Biol., 45, 227–238, 2000.
Duhamel, S., Van Wambeke, F., Lefevre, D., Benavides, M., and Bonnet, S.: Mixotrophic metabolism by natural communities of unicellular cyanobacteria in the western tropical South Pacific Ocean, Environ. Microbiol., 20, 2743–2756, https://doi.org/10.1111/1462-2920.14111, 2018.
Epstein, S. S., Burkovsky, I. V., and Shiaris, M. P.: Ciliate grazing on bacteria, flagellates, and microalgae in a temperate zone sandy tidal flat: ingestion rates and food niche partitioning, J. Exp. Mar. Biol. Ecol., 165, 103–123, https://doi.org/10.1016/0022-0981(92)90292-I, 1992.
Esteban, G. F., Fenchel, T., and Finlay, B. J.: Mixotrophy in ciliates, Protists, 161, 621–641, https://doi.org/10.1016/j.protis.2010.08.002, 2010.
Faure, V., Pinazo, C., Torréton, J. P., and Jacquet, S.: Modelling the spatial and temporal variability of the SW lagoon of New Caledonia I: A new biogeochemical model based on microbial loop recycling, Mar. Pollut. Bull., 61, 465–479, https://doi.org/10.1016/j.marpolbul.2010.06.041, 2010a.
Faure, V., Pinazo, C., Torréton, J. P., and Douillet, P.: Modelling the spatial and temporal variability of the SW lagoon of New Caledonia II: Realistic 3D simulations compared with in situ data, Mar. Pollut. Bull., 61, 480–502, https://doi.org/10.1016/j.marpolbul.2010.06.040, 2010b.
Fisher, N. L. and Halsey, K. H.: Mechanisms that increase the growth efficiency of diatoms in low light, Photosynth. Res., 129, 183–197, 2016.
Flynn, K. J. and McGillicuddy, D. J.: Modeling marine harmful algal blooms: Current status and future prospects, Harmful Algal Blooms: A Compendium Desk Reference, 115–134, https://doi.org/10.1002/9781118994672.ch3, 2018.
Flynn, K. J., Stoecker, D. K., Mitra, A., Raven, J., Glibert, P. M., Hansen, P. J., Graneli, E., and Burkholder, J. M.: Misuse of the phytoplankton–zooplankton dichotomy: the need to assign organisms as mixotrophs within plankton functional types, J. Plankton Res., 35, 3–11, https://doi.org/10.1093/plankt/fbs062, 2012.
Fraysse, M., Pinazo, C., Faure, V. M., Fuchs, R., Lazzari, P., Raimbault, P., and Peyraud, I.: Development of a 3D Coupled Physical-Biogeochemical Model for the Marseille Coastal Area (NW Mediterranean Sea): What Complexity Is Required in the Coastal Zone?, PLoS ONE, 8, e80012, https://doi.org/10.1371/journal.pone.0080012, 2013.
Fraysse, M., Pairaud, I., Ross, O. N., Faure, V. M., and Pinazo, C.: Intrusion of Rhone River diluted water into the Bay of Marseille: Generation processes and impacts on ecosystem functioning, J. Geophys. Res.-Oceans, 119, 119, 6535–6556, https://doi.org/10.1002/2014JC010022, 2014.
Garcia, F. and Raimbault, P.: Panache du Rhône: Suivi des eaux du Rhône en zone côtière par mesure haute fréquence de la salinité de surface, MIO UMR 7294 CNRS [data set], https://doi.org/10.34930/79C421C5-9335-4957-88FA-804FB4AE4B43, 2022.
Garcia, N., Gregori, G., Lafont, M., Lagadec, V., Nunige, S., and Raimbault, P.:SOMLIT-Frioul time series (French Research Infrastructure ILICO): long-term core parameter monitoring in the Bay of Marseille, SEANOE [data set], https://doi.org/10.17882/96252, 1997.
Gatti, J., Petrenko, A., Devenon, J. -L., Leredde, Y., and Ulses, C.: The Rhone River dilution zone present in the northeastern shelf of the Gulf of Lion in December 2003, Cont. Shelf Res., 26, 1794–1815, https://doi.org/10.1016/j.csr.2006.05.012, 2006.
Gaudy, R. and Thibault-Botha, D.: Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean, Prog. Oceanogr., 72, 151–163, https://doi.org/10.1016/j.pocean.2007.01.005, 2007.
Gehlen, M., Gangstø, R., Schneider, B., Bopp, L., Aumont, O., and Ethe, C.: The fate of pelagic CaCO3 production in a high CO2 ocean: a model study, Biogeosciences, 4, 505–519, https://doi.org/10.5194/bg-4-505-2007, 2007.
Geider, R.: Algal photosynthesis, Vol. 2, Springer Science & Business Media, ISBN 1475721536, 2013.
Geider, R., MacIntyre, H. L., and Kana T. M.: A dynamic regulatory model of phytoplanktonic acclimation to light, nutrients, and temperature, Limnol. Oceanogr., 43, 679–694, 1998.
Ghyoot, C., Flynn, K. J., Mitra, A., Lancelot, C., and Gypens, N.: Modeling Plankton Mixotrophy: A Mechanistic Model Consistent with the Shuter-Type Biochemical Approach, Frontiers in Ecology and Evolution, 5, 5–78, https://doi.org/10.3389/fevo.2017.00078, 2017.
Gibson, R. N., Atkinson, R. J. A., and Gordon, J. D. M.: Review of three-dimensional ecological modelling related to the North Sea shelf system, Part II: Model validation and data needs, Oceanogr. Mar. Biol., 44, 1–60, 1420006398, 2006.
Glibert, P. M. and Legrand, C., The Diverse Nutrient Strategies of Harmful Algae: Focus on Osmotrophy, Ecology of Harmful Algae, 163–175, https://doi.org/10.1007/978-3-540-32210-8_13, 2006.
Glibert, P. M., Al-Azri, A., Allen, J. I., Bouwman, A. F., Beusen, A. H. W., Burford, M. A., Harrison, P. J., and Zhou, M.: Key Questions and Recent Research Advances on Harmful Algal Blooms in Relation to Nutrients and Eutrophication, Global Ecology and Oceanography of Harmful Algal Blooms, Ecol. Stud., 232, 229–258, https://doi.org/10.1007/978-3-319-70069-4_12, 2018.
Gorsky, G., Dallot, S., Sardou, J., Fenaux, R., Carré, C., and Palazzoli, I.: C and N composition of some northwestern Mediterranean zooplankton and micronekton species, J. Exp. Mar. Biol. Ecol., 124, 133–144, https://doi.org/10.1016/0022-0981(88)90116-5, 1988.
Guillemain, D.: Données de température à la station Planier-Souquet (5m), Regional temperature observation network T-MEDNet [data set], https://t-mednet.org/request-data (last access: 7 November 2023), 2021.
Graneli, E., Carlsson, P., and Legrand, C.: The role of C, N and P in dissolved and particulate organic matter as a nutrient source for phytoplankton growth, including toxic species, Aquat. Ecol., 33, 17–27, 1999.
Hartmann, M., Grob, C., Tarran, G. A., Martin, A. P., Burkill, P. H., Scanlan, D. J., and Zubkova, M. V.: Mixotrophic basis of Atlantic oligotrophic ecosystems, P. Natl. Acad. Sci. USA, 109, 5756–5760, https://doi.org/10.1073/pnas.1118179109, 2012.
Johansson, M., Gorokhova, E., and Larsson, U. L. F.: Annual variability in ciliate community structure, potential prey and predators in the open northern Baltic Sea proper, J. Plankton Res., 26, 67–80, https://doi.org/10.1093/plankt/fbg115, 2004.
Jost, C., Lawrence, C. A., Campolongo, F., Van de Bund, W., Hill, S., and DeAngelis, D. L.: The effects of mixotrophy on the stability and dynamics of a simple planktonic food web model, Theor. Popul. Biol., 66, 37–51, https://doi.org/10.1016/j.tpb.2004.02.001, 2004.
Kempton, J. W., Lewitus, A. J., Deeds, J. R., Law, J. M., and Place, A. R.: Toxicity of Karlodinium micrum (Dinophyceae) associated with a fish kill in a South Carolina brackish retention pond, Harmful Algae, 1, 233–241, https://doi.org/10.1016/S1568-9883(02)00015-X, 2002.
Kirchman, D. L. and Gasol, J. M.: Microbial Ecology of the Oceans, John Wiley & Sons, ISBN 1119107199, 2018.
Lacroix, G. and Grégoire, M.: Revisited ecosystem model (MOD-ECOGeL) of the Ligurian Sea: seasonal and interannual variability due to atmospheric forcing, J. Marine Syst., 37, 229–258, https://doi.org/10.1016/S0924-7963(02)00190-2, 2002.
Lajaunie-Salla, K., Diaz, F., Wimart-Rousseau, C., Wagener, T., Lefèvre, D., Yohia, C., Xueref-Remy, I., Nathan, B., Armengaud, A., and Pinazo, C.: Implementation and assessment of a carbonate system model (Eco3M-CarbOx v1.1) in a highly dynamic Mediterranean coastal site (Bay of Marseille, France), Geosci. Model Dev., 14, 295–321, https://doi.org/10.5194/gmd-14-295-2021, 2021.
Leblanc, K., Quéguiner, B., Diaz, F., Cornet, V., Michel-Rodriguez, M., Durrieu de Madron, X., Bowler, C., Malviva, S., Thyssen, M., Grégori, G., Rembauville, M., Grosso, O., Poulain, J., de Vargas, C., Pujo-Pay, M., and Conan, P.: Nanoplanktonic diatoms are globally overlooked but play a role in spring blooms and carbon export, Nat. Commun., 9, 953–964, https://doi.org/10.1038/s41467-018-03376-9, 2018.
Leles, S. G., Bruggeman, L. P. J., Blackford, J., Ciavatta, S., Mitra, A., and Flynn, K. J.: Modelling mixotrophic functional diversity and implications for ecosystem function, J. Plankton Res., 40, 627–642, https://doi.org/10.1093/plankt/fby044, 2018.
Lewitus, A. J: Osmotrophy in marine microalgae, Algal cultures, analogues of blooms and applications, Volume 1, Subba Rao D. V., Science publishers Inc., USA, ISBN 1578083923, 2006.
Livanou E., Lagaria, A., Santi, I., Mandalakis, M., Paylidou, A., Lika, K., and Psarra, S.: Pigmented and heterotrophic nanoflagellates: Abundance and grazing on prokaryotic picoplankton in the ultra-oligotrophic Eastern Mediterranean Sea, Deep-Sea Res. Pt. II, 164, 100–111, https://doi.org/10.1016/j.dsr2.2019.04.007, 2019.
Livanou, E., Oikonomou, A., Psarra, S., and Konstadia, L.: Role of mixotrophic nanoflagellates in the Eastern Mediterranean microbial food web, Mar. Ecol. Prog. Ser., 672, 15–32, https://doi.org/10.3354/meps13782, 2021.
Marechal, D.: A soil-based approach to rainfall-runoff modelling in ungauged catchments for England and Wales, PhD thesis, Cranfield University, 157 pp., 2004.
Margalef, R.: Life-forms of phytoplankton as survival alternatives in an unstable environment, edited by: Gauthier-Villars, Oceanol. Acta, 1, 493–509, https://archimer.ifremer.fr/doc/00123/23403/ (last access: July 2023), 1978.
Marty, J.-C., Chiavérini, J., Pizay, M.-D., and Avril, B.: Seasonaland interannual dynamics of nutrients and phytoplankton pigments in the western Mediterranean Sea at the DYFAMED time series station (1991–1999), Deep-Sea Res. Pt. II, 49, 1965–1985, https://doi.org/10.1016/S0967-0645(02)00022-X, 2001.
McGillicuddy Jr., D. J.: Models of harmful algal blooms: conceptual, empirical, and numerical approaches, J. Marine Syst., 83, 105, 2010.
Mella-Flores, D., Mazard, S., Humily, F., Partensky, F., Mahé, F., Bariat, L., Courties, C., Marie, D., Ras, J., Mauriac, R., Jeanthon, C., Mahdi Bendif, E., Ostrowski, M., Scanlan, D. J., and Garczarek, L.: Is the distribution of Prochlorococcus and Synechococcus ecotypes in the Mediterranean Sea affected by global warming?, Biogeosciences, 8, 2785–2804, https://doi.org/10.5194/bg-8-2785-2011, 2011.
Millet, B., Pinazo, C., Banaru, D., Pagès, R., Guiart, P., and Pairaud, I.: 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), Édité par João Miguel Dias, PLoS ONE, 13, e0195257, https://doi.org/10.1371/journal.pone.0195257, 2018.
Millette, N. C., Pierson, J. J., Aceves, A., and Stoecker, D. K.: Mixotrophy in Heterocapsa rotundata: a mechanism for dominating the winter phytoplankton, Limnol. Oceanogr., 62, 836–845, https://doi.org/10.1002/lno.10470, 2017.
Millot, C.: The Golf of Lions' hydrodynamic, Cont. Shelf Res., 10, 885–894, 1990.
Mitra, A. and Flynn, K. J.: Modelling mixotrophy in harmful algal blooms: More or less the sum of the parts?, J. Marine Syst., 83, 158–169, https://doi.org/10.1016/j.jmarsys.2010.04.006, 2010.
Mitra, A., Flynn, K. J., Burkholder, J. M., Berge, T., Calbet, A., Raven, J. A., Granéli, E., Glibert, P. M., Hansen, P. J., Stoecker, D. K., Thingstad, F., Tillmann, U., Våge, S., Wilken, S., and Zubkov, M. V.: The role of mixotrophic protists in the biological carbon pump, Biogeosciences, 11, 995–1005, https://doi.org/10.5194/bg-11-995-2014, 2014.
Mitra, A., Flynn, K. J., Tillmann, U., Raven, J. A., Caron, D., Stoecker, D. K., Not, F., Hansen, P. J., Hallegraeff, G., Sanders, R., Wilken, S., McManus, G., Johnson, M., Pitta, P., Våge, S., Berge, T., Calbet, A., Thingstad, F., Jin Jeong, H., Burkholder, J. -A., Glibert, P. M., Granéli, E., and Lundgren, V.: Defining planktonic protist functional groups on mechanisms for energy and nutrient acquisition: Incorporation of diverse mixotrophic strategies, Protist, 167, 106–120, https://doi.org/10.1016/j.protis.2016.01.003, 2016.
Montagnes, D. J.: Growth responses of planktonic ciliates in the genera Strobilidium and Strombidium, Mar. Ecol. Prog. Ser., 130, 241–254, https://doi.org/10.3354/meps130241, 1996.
Morel, A. and André, J. -M.: Pigment distribution and primary production in the western Mediterranean as derived and modelled from coastal zone colour scanner observations, J. Geophys. Res.-Oceans, 96, 12685–12698, https://doi.org/10.1029/91JC00788, 1991.
Nielsen, L. P., Christensen, P. B., Revsbech, N, P., and Sorensen, I.: Denitrification and photosynthesis in stream sediment studied with microsensor and wholecore techniques, Limnol. Oceanogr., 35, 1135–1144, https://doi.org/10.4319/lo.1990.35.5.1135, 1990.
Oikomonou, A., Livanou, E., Mandalakis, M., Ligaria, A., and Psarra, S.: Grazing effect of flagellates on bacteria in response to phosphate addition in the oligotrophic Cretan Sea, NE Mediterranean, FEMS Microbiol. Ecol., 96, fiaa086, https://doi.org/10.1093/femsec/fiaa086, 2020.
Oursel, B., Garniera, C., Zebracki, M., Durrieu, G., Pairaud, I., Omanovic, D., Cossab, D., and Lucas, Y.: Flood inputs in a Mediterranean coastal zone impacted by a large urban area: Dynamic and fate of trace metals, Mar. Chem., 167, 44–56, https://doi.org/10.1016/j.marchem.2014.08.005, 2014.
Pitta, P. and Giannakourou, A.: Planktonic ciliate in the oligotrophic Eastern Mediterranean: vertical, spatial distribution and mixotrophy, Mar. Ecol. Prog. Ser., 194, 269–282, 2000.
Platt, T. G. C. L., Gallegos, C. L., and Harrison, W. G.: Photoinhibition of photosynthesis in natural assemblages of marine phytoplankton, J. Mar. Res., 38, 1525, 1980.
Polovina, J. J., Howell, E. A., and Abecassis, A.: Ocean's least productive waters are expanding, Geophys. Res. Lett., 35, L03618, https://doi.org/10.1029/2007GL031745, 2008.
Pratt, J. R. and Cairns, J.: Functional groups in the protozoa, Roles in differing ecosystems, J. Protozool. Res., 32, 415–423, 1985.
Price, R. W. and Turner, J. T.: Ecology of planktonic ciliates in marine food webs, Rev. Aquat. Sci., 6, 139–181, 1992.
Ptacnik, R., Sommer, U., Hansen, T., and Martens, V.: Effects of microzooplankton and mixotrophy in an experimental planktonic food web, Limnol. Oceanogr., 49, 1435–1445, 2004.
Ptacnik, R., Gomes, A., Royer, S-J., Berger, S. A., Calbet, A., Nejstgaard, J. C., Gasol, J. M., Isari, S., Moorthi, S. D., Ptacnikova, R., Striebel, M., Sazhin, A. F., Tsagaraki, T. M., Zervoudaki, S., Altoja, K., Dimitriou, P. D., Laas, P., Gazihan, A., Martinez, R. A., Schabhuttl, S., Santi, I., Sousoni, D., and Pitta, P.: A light-induced shortcut in the planktonic microbial loop, Sci. Rep., 6, 29286, https://doi.org/10.1038/srep29286, 2016.
Pujo-Pay, M., Conan, P., Oriol, L., Cornet-Barthaux, V., Falco, C., Ghiglione, J.-F., Goyet, C., Moutin, T., and Prieur, L.: Integrated survey of elemental stoichiometry (C, N, P) from the western to eastern Mediterranean Sea, Biogeosciences, 8, 883–899, https://doi.org/10.5194/bg-8-883-2011, 2011.
Putt, M.: Metabolism of photosynthate in the chloroplast-retaining ciliate Laboea strobila, Mar. Ecol. Prog. Ser., Oldendorf, 60, 271–282, 1990.
Raven, J. A.: Phagotrophy in phototrophs, Limnol. Oceanogr., 42, 198–205, 1997.
Razzak, S. A., Ilyas, M., Ali, S. A. M., and Hossain, M. M.: Effects of CO2 Concentration and pH on Mixotrophic Growth of Nannochloropsis oculate, Appl. Biochem. Biotech., 176, 1290–1302, https://doi.org/10.1007/s12010-015-1646-7, 2015.
Riemann, B., Havskum, H., Thingstad, F., and Bernard, C.: The role of mixotrophy in pelagic environments, Molecular Ecology of Aquatic Microbes, NATO ASI Series, 38, Joint, I. Springer, Berlin, Heidelberg, https://doi.org/10.1007/978-3-642-79923-5_6, 1995.
Ross, O. N., Fraysse, M., Pinazo, C., and Pairaud, I.: Impact of an intrusion by the Northern Current on the biogeochemistry in the Eastern Gulf of Lion, NW Mediterranean, Estuar. Coast. Shelf S., 170, 1–9, 2016.
Sanders, R.: Mixotrophic Protists in Marine and Freshwater Ecosystems, J. Protozool., 38, 76–81, 1991.
Sarthou, G., Timmerman, K. R., Blain, S., and Tréguer, P.: Growth physiology and fate of diatoms in the ocean: a review, J. Sea Res., 53, 25–42, https://doi.org/10.1016/j.seares.2004.01.007, 2005.
Schaeffer, A., Molcard, A., Forget, P., Fraunié, P., and Garreau, P.: Generation mechanisms for mesoscale eddies in the Gulf of Lions: radar observation and modelling, Ocean Dynam., 61, 1587–1609, https://doi.org/10.1007/s10236-011-0482-8, 2011.
Sherr, B. F., Sherr, E. B., Caron, D. A., Vaulot, D., and Worden, A. Z.: Oceanic protists, Oceanography, 20, 130–134, https://doi.org/10.5670/oceanog.2007.57, 2007.
Stickney, H. L., Hood, R. R., and Stoecker, D. K.: The impact of mixotrophy on planktonic marine ecosystems, Ecol. Model., 125, 203–230, https://doi.org/10.1016/S0304-3800(99)00181-7, 2000.
Stoecker, D. K.: Mixotrophy among Dinoflagellates 1, J. Eukaryot. Microbiol., 46, 397–401, 1999.
Stoecker, D. K. and Capuzzo, J. M.: Predation on protozoa: its importance to zooplankton, J. Plankton Res., 12, 891–908, 1990.
Stoecker, D. K., Silver, M. W., Michaels, A. E., and Davis, L. H.: Obligate mixotrophy in Laboea strobila, a ciliate which retains chloroplasts, Mar. Biol., 99, 415–423, 1988.
Stoecker, D. K., Li, A., Coats, D. W., Gustafson, D. E., and Nannen, M. K.: Mixotrophy in the dinoflagellate Prorocentrum minimum, Mar. Ecol. Prog. Ser., 152, 1–12, 1997.
Stoecker, D. K.: Conceptual models of mixotrophy in planktonic protists and some ecological and evolutionary implications, Eur. J. Protistol., 34, 281–290, https://doi.org/10.1016/S0932-4739(98)80055-2, 1998.
Stoecker, D. K., Hansen, P. J., Caron, D. A., and Mitra, A.: Mixotrophy in marine plankton, Annu. Rev. Mar. Sci., 9, 11–35, https://doi.org/10.1146/annurev-marine-010816-060617, 2017.
Tett, P.: A three-layer vertical and microbiological processes model for shelf seas, Report No. 14, Proudman Oceanographic Laboratory, Birkenhead, UK, 85, https://nora.nerc.ac.uk/id/eprint/3877/1/ir14.pdf (last access: 7 November 2023), 1990.
Thingstad, T. F., Hagström, A., and Rassoulzadegan, F.: Accumulation of degradable DOC in surface waters: Is it caused by a malfunctioning microbialloop?, Limnol. Oceanogr., 42, 398–404, https://doi.org/10.4319/lo.1997.42.2.0398, 1997.
Timmermans, K. R., van der Wagt, B., Veldhuis, M. J. W., Maatman, A., and de Baar, H. J. W.: Physiological responses of three species of marine pico-phytoplankton to ammonium, phosphate, iron and light limitation, J. Sea Res., 53, 109–120, https://doi.org/10.1016/j.seares.2004.05.003, 2005.
Thornley, J. H. M. and Cannell, M. G. R.: Modelling the component of plant respiration: Representation and realism, Ann. Bot.-London, 85, 55–67, https://doi.org/10.1006/anbo.1999.0997, 2000.
Unrein, F., Gasol, J. M., and Massana, R.: Dinobryon faculiferum (Chrysophyta) in coastal Mediterranean seawater: presence and grazing impact on bacteria, J. Plankton Res., 32, 559–564, https://doi.org/10.1093/plankt/fbp150, 2010.
Verity, P. G. and Paffenhofer, G. A.: On assessment of prey ingestion by copepods, J. Plankton Res., 18, 1767–1779, 1996.
Vrede, K., Heldal, M., Norland, S., and Bratbak, G.: Elemental Composition (C, N, P) and Cell Volume of Exponentially Growing and Nutrient-Limited Bacterioplankton, American Society for Microbiology Journal, 68, 2965–2971, https://doi.org/10.1128/AEM.68.6.2965-2971.2002, 2002.
Wanninkhof, R.: Relationship between wind speed and gas exchange over the ocean revisited, Limnol. Oceanogr.-Meth., 12, 351–362, https://doi.org/10.4319/lom.2014.12.351, 2014.
Ward, B. A. and Follows M. J.: Marine mixotrophy increases trophic transfer efficiency, mean organism size, and vertical carbon flux, P. Natl. Acad. Sci. USA, 113, 2958–2963, https://doi.org/10.1073/pnas.1517118113, 2016.
Wilken, S., Huisman, J., Naus-Wiezer, S., and Van Donk, E.: Mixotrophic organisms become more heterotrophic with rising temperature, Ecol. Lett., 16, 225–233, https://doi.org/10.1111/ele.12033, 2013.
Xueref-Remy, I., Lefèvre, D., Nathan, B., Milne, M., Piazzola, J., Yohia, C., Armengaud, A., Blanc, P. E. Mevy, J.-P., Reiter, I., Paya, A., Lajaunie-Salla, K., Wimart-Rousseau, C., Diaz, F., Pinazo, C., and Wagener, T.: Aix-Marseille Carbon Pilot Study (AMC), https://www.otmed.fr/research-projects-and-results/result-2449 (last access: 7 November 2023), 2016.
Yacobi, Y. Z., Zohary, T., Kress, N., Hecht, A., Robarts, R. D., Waiser, M., Wood, A. M., and Li, W. K. W.: Chlorophyll distribution throughout the southeastern Mediterranean in relation to the physical structure of the water mass, J. Marine Syst., 6, 179–190, https://doi.org/10.1016/0924-7963(94)00028-A, 1995.
Yohia, C.: Genèse du mistral par interaction barocline et advection du tourbillon potentiel, Climatologie, 13, 24–37, https://doi.org/10.4267/climatologie.1182, 2017.
Zubkov, M. V. and Tarran, G. A.: High bacterivory by the smallest phytoplankton in the North Atlantic Ocean, Nature, 455, 224–226, https://doi.org/10.1038/nature07236, 2008.
Short summary
While several studies have shown that mixotrophs play a crucial role in the carbon cycle, the impact of environmental forcings on their dynamics remains poorly investigated. Using a biogeochemical model that considers mixotrophs, we study the impact of light and nutrient concentration on the ecosystem composition in a highly dynamic Mediterranean coastal area: the Bay of Marseille. We show that mixotrophs cope better with oligotrophic conditions compared to strict auto- and heterotrophs.
While several studies have shown that mixotrophs play a crucial role in the carbon cycle, the...
