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Abstract In the Mediterranean Sea, seaweed marine forests, one of the most important benthic assemblages for the coastal ecosystem functioning, are shaping the rocky reefs. Among the Fucales, Gongolaria barbata (syn. Cystoseira barbata) is a several years living species usually growing in very shallow, sheltered and well-lit marine reef habitats and in coastal lagoons. The long-term change in its distribution in relation to a variety of disturbances has been assessed along the French Mediterranean coast, including Corsica and brackish lagoons, thanks to historical data dating back to the 19th century. The current distribution was established through an extensive survey conducted by snorkelling, encompassing approximately 3 000 km of coastline. A GIS analysis indicates that Gongolaria barbata can currently be considered as regionally extinct in French Catalonia and Western Provence and functionally extinct in the French Riviera. In Languedoc, the species is extinct in the open sea and only present in certain brackish lagoons despite severe repeated anoxic crises (malaïgues) and competition with a rich exotic flora introduced from the NE Atlantic and NW Pacific, especially the invasive Sargassum muticum. In contrast, the populations of G. barbata have remained stable in Eastern Provence and Corsica. The main possible causes of decline are uprooting, overgrazing by herbivores, habitat destruction and competition with introduced seaweeds. The relevance of ecological restoration of G. barbata populations was assessed. more...

Early-life stages play a key role in the dynamics of bipartite life cycle marine fish populations. Difficult to monitor, observations of these stages are often scattered in space and time. While Mediterranean coastlines have often been surveyed, no effort has been made to assemble historical observations. Here we build an exhaustive compilation of dispersal traits for coastal fish species, considering in situ observations and growth models (Di Stefano et al., 2023; https://doi.org/10.17882/91148). Our database contains over 110 000 entries collected from 1993 to 2021 in various subregions. All observations are harmonized to provide information on dates and geolocations of both spawning and settlement, along with pelagic larval durations. When applicable, missing data and associated confidence intervals are reconstructed from dynamic energy budget theory. Statistical analyses allow traits’ variability to be revisited and sampling biases to be revealed across taxa, space and time, hence providing recommendations for future studies and sampling. Comparison of observed and modelled entries provides suggestions to improve the feed of observations into models. Overall, this extensive database is a crucial step to investigate how marine fish populations respond to global changes across environmental gradients. more...

The specific identification of three major morphotypes of the tropical holopelagic Sargassum species causing massive strandings on the African and Caribbean coastlines was attempted by morphological characterisation as well as quantitative and qualitative analyses of several metabolites. Of the 25 morphological variables studied on 208 samples from the North Atlantic Ocean, 22 were used to establish a dichotomous identification key, allowing without any doubt the identification of each morphotype based on their morphological criteria alone. We also attempted to differentiate morphotypes using chemical fingerprintings (HR-MAS NMR) and markers by analysing pigment level and composition using High Pressure Liquid Chromatography, terpene profiles by Thin Layer Chromatography, phenolic compound levels by the Folin-Ciocalteu assay and structures by 2D Nuclear Magnetic Resonance spectroscopy, and fatty acid composition by Gas Chromatography. While pigment level and composition, terpene profiles, and phenolic contents were not discriminating, quantification of eight fatty acids enabled the differentiation of the three morphotypes. Furthermore, phlorotannin purification permitted their structural characterisation allowing discrimination between the three morphotypes. Our study highlights the potential of the free fatty acid profile and phlorotannin structure as good chemomarkers in order to discriminate between the three morphotypes of holopelagic Sargassum. more...

Since 2011, massive new strandings of holopelagic Sargassum have been reported on the coasts of the Caribbean, northern Brazil, Guiana, and West Africa, causing severe economic and ecological damage. Three common morphotypes (S. fluitans III, S. natans I, and S. natans VIII) were identified as responsible for these catastrophic events, with dominance shifts between them over time. However, the taxonomic status of these holopelagic Sargassum morphotypes remains unclear. Using an integrative taxonomy framework, combining a morphological study and molecular analyses, this study aimed to clarify their taxonomic status. Morphological analyses of 54 characters revealed no intermediate form between the three morphotypes, with the overall shape, nature of the axis, and size and shape of blades and vesicles being the most discriminating. An analysis of mitochondrial (IGS, cox2, cox3, mt16S rRNA, and nad6) and plastid (rbcL) markers confirmed the genetic divergence among the three morphotypes, with a lower level of divergence between the two S. natans morphotypes. Without additional molecular characterization, these morphotypes cannot be classified as three distinct species. However, due to their distinct morphological characteristics and sympatry within drifting aggregations, a revision of holopelagic species names is proposed, with Sargassum fluitans var. fluitans (for S. fluitans III), Sargassum natans var. natans (for S. natans I), and S. natans var. wingei (for S. natans VIII). This revision provides necessary clarity on the species involved in inundations of the tropical Atlantic. more...

The Mediterranean Sea, a biodiversity hotspot, faces significant threats from non-indigenous species (NIS), which drive biodiversity changes. Over the past century, the introduction of NIS has accelerated due to maritime traffic, aquaculture, and interoceanic canals, fostering biological invasions. Marine protected areas (MPAs), established to preserve biodiversity, are increasingly impacted. This review quantified and characterized French Mediterranean MPAs, analyzing non-indigenous macroalgae distribution based on the existing literature and the authors’ observations. Results revealed widespread occurrence, with the highest NIS richness in strictly regulated MPAs; their proximity to large harbors highlights the paramount importance of the introduction pathways. In addition, there is a significant knowledge gap regarding the distribution of NIS within MPAs, complicating efforts to monitor and study these species effectively. These findings highlight the challenges in monitoring and managing invasions and the urgent need for controlling primary and secondary invasion pathways, within and outside the MPAs, international collaboration to control them, and enhanced funding for NIS monitoring. Without adaptive management, even strictly protected MPAs are vulnerable to the escalating impacts of invasive species. more...

The analysis of biological and ecological traits has a long history in evolutionary and ecological research. However, trait data are often scattered and standardised terminology that transcends taxonomic and biogeographical context are generally missing. As part of the development of a global trait database of marine species, we collated trait information for European seaweeds and structured the data within the standardised framework of the World Register of Marine Species (WoRMS). We collected 45 175 trait records for 21 biologically and ecologically relevant traits of seaweeds. This resulted in a trait database for 1745 European seaweed species of which more than half (56 %) of the records were documented at the species level, while the remaining 44 % were documented at a higher taxonomic level and subsequently inherited at lower levels. The trait database for European seaweeds will serve as a foundation for future research on diversity and evolution of seaweeds and their responses to global changes. The data will contribute to developing detailed trait-based ecosystem models and will be an important tool to inform marine conservation policies. The data are publicly accessible through the AlgaeTraits portal, https://doi.org/10.14284/574 (AlgaeTraits, 2022). more...

Abstract The calcified red macroalga Lithophyllum byssoides, a very common midlittoral species in the western Mediterranean Sea, is a significant ecosystem engineer capable, under exposed and dim light conditions, of building wide and solid endemic bioconstructions near the mean sea level: the L. byssoides rims or 'trottoirs à L. byssoides'. Although the growth of the species is relatively rapid for a calcified alga, the construction of a large rim requires several centuries of near stable or slowly rising sea level. As the time scale of their formation is measured in centuries, L. byssoides bioconstructions constitute valuable and sensitive sea level markers. The health status of L. byssoides rims has been studied at two sites located far apart from each other (Marseille and Corsica), both in areas heavily impacted by humans and in areas with little impact (MPAs and unprotected areas). A health index is proposed: Lithophylum byssoides Rims Health Index. The main and inevitable threat is the rise in the sea level. This ecosystem would be the first case worldwide of marine ecosystem collapse resulting, indirectly, from man-induced global change. more...

The worldwide decline of macroalgal forests is raising major concerns for the potentially negative consequences on biodiversity and ecosystem functions, pushing for the definition of specific conservation and restoration measures. Protecting and restoring these habitats requires detailed information on their distribution, ecological status, and drivers of decline. Here, we provide the most updated available information on the distribution of Mediterranean Cystoseira s.l. forests by conducting a comprehensive bibliographic survey of literature published from 2009 to 2021, complemented by unpublished data. We also provide insights into the ecological status of these forests and the stressors affecting them across the Mediterranean basin. Our results show that most Mediterranean coasts remain un(der)studied and that the available information is concentrated in spatially limited coastal areas, restricted to very few species. When the ecological status is reported, data is highly heterogeneous, making any comparisons problematic, what claims for the description and use of easy and standardized monitoring methods for comparative purposes. Drivers of decline of Cystoseira s.l. forest have been even less investigated and, therefore, still poorly characterized. Our results highlight that our current knowledge is still insufficient to implement effective conservation and restoration strategies at the basin scale but also regionally. We call for the urgent need for mapping and standardized monitoring of Cystoseira s.l. forests to obtain baseline information for future management strategies involving their conservation, the mitigation of the stressors threatening them and the restoration of the degraded forests. more...

Abstract Coastal zones are ecosystems of high economic value but exposed to numerous disturbances, while they represent nurseries for many fish species, raising the issue of the preservation of their functions and services. In this context, the juvenile fish assemblages of all types of habitats present in shallow coastal zones were studied on the south-east coast of France using underwater visual censuses in warm (June–July 2014) and cold (April 2015) periods. A total of fourteen habitat types were characterized, which could be grouped into three broad categories, rocky substrates (natural and artificial), sedimentary bottoms with all levels of granulometry, and seagrass beds including Cymodocea nodosa and Posidonia oceanica meadows; the ecotones or interfaces between the three broad habitat categories were individualized as particular habitat types. The abiotic and biotic descriptors of the 14 habitat types individualized did not vary with time, except for a higher cover percentage and canopy height of macrophytes in the warm period, which increased the three-dimensional structure of some habitats. The taxonomic composition and density of juvenile fish assemblages were analyzed using both multivariate and univariate descriptors, after grouping the 57 fish species recorded into 41 well-individualized taxa. Juvenile fishes were recorded in all habitat types, with higher mean species richness and abundance during the warm than the cold period. The richest habitats in terms of both fish species richness and abundance were the natural rocky substrates and the interfaces between Posidonia beds and the other habitats. Although juvenile fish assemblage composition differed among habitat types and between periods, the most abundant fish species were Atherina sp., Sarpa salpa, Gobiidae, Symphodus spp., Pagellus spp. and several Diplodus species, which colonized 7 up to 14 different habitat types (depending on taxa) during their juvenile life. Most species settled in one or a few specific habitats but rapidly colonized adjacent habitats when growing. This study provided evidence of the role of all types of shallow coastal habitats as fish nurseries and their varying pattern of occupation in space and time by the different juvenile stages. It highlighted the importance of the mosaic of habitats and interfaces for the complete development of all juvenile life stages of fishes, and for the preservation of a high diversity of coastal fish assemblages and fisheries resources in the Mediterranean Sea. more...

Dispersal is a central process that affects population growth, gene flow, and ultimately species persistence. Here we investigate the extent to which gene flow occurs between fragmented populations of the deep-water brown algae Ericaria zosteroides (Turner) Greville (Sargassaceae, Fucales). These investigations were performed at different spatial scales from the bay of Marseille (western Provence) to Corsica. As dispersal of zygotes is shown to be limited over distances beyond a few meters, we used a multidisciplinary approach, based on Lagrangian modeling and population genomics to test the hypothesis that drifting of fertile parts of thallus (eggs on fertile branches), mediated by ocean currents, enable occasional gene flow between populations. Therefore we assessed the respective contribution of oceanographic connectivity, geographical isolation, and seawater temperatures to the genetic structure of this species. The genetic structure was assessed using 10,755 neutral SNPs and 12 outlier SNPs genotyped by dd-RAD sequencing in 261 individuals of E. zosteroides. We find that oceanographic connectivity is the best predictor of genetic structure, while differentiation in outlier SNPs can be explained by the depth of populations, as emphasized by the minimum seawater temperature predictor. However, further investigations will be necessary for clarifying how depth drives adaptive genetic differentiation in E. zosteroides. Our analyses revealed that local hydrodynamic conditions are correlated with the very high divergence of one population in the Bay of Marseille. Overall, the levels of gene flow mediated by drifting were certainly not sufficient to counteract differentiation by local genetic drift, but enough to allow colonization several kilometers away. This study stresses the need to consider secondary dispersal mechanisms of presumed low dispersal marine species to improve inference of population connectivity. more...

The direct and indirect impact of fish farms, shellfish aquaculture, and extensive forms of aquaculture such as seeding of juvenile sea urchins, on macrophytes (seaweeds and seagrasses), is reviewed in Mediterranean benthic ecosystems. Fish farms constitute a source of organic matter and nutrients (food and fecal pellets) that causes the extirpation of Posidonia oceanica seagrass meadows beneath and near to farm facilities. In addition to direct effects, the nitrogen enrichment of macrophytes tissues increases the grazing pressure by herbivorous fishes and sea urchins. In some cases, the impact can continue to increase several years after the cessation of farming activities. Natural restoration of extirpated seagrass meadows is generally unlikely at the human time scale. Shellfish aquaculture is the cause of the main flow of introduced macrophytes in the Mediterranean; the main vector is the importation of oyster spat from Japan and Korea. North-eastern Pacific seaweeds are now the dominant biotic component of some Mediterranean lagoons (e.g., Thau, Mar Piccolo, and Venice lagoons). In addition to direct effects, mussel aquaculture can constitute a source of larvae that flow with currents, the adults of which can overwhelm seaweed forests (e.g., Carpodesmia mediterranea). Shellfish aquaculture is also a source of fecal pellets, resulting in changes in bottom macrophytes, and a vector of diseases of metazoans, the extirpation of which may change the functioning of recipient macrophyte ecosystems. The edible sea urchin Paracentrotus lividus is sometimes erroneously considered as in decline due to over-harvesting. However, its abundance in the second half of the 20th century was probably a consequence of human impact (overfishing of its predatory fish, organic pollution. This man-induced proliferation resulted in the extirpation of seaweed forests (e.g., Carpodesmia spp., Treptacantha spp. – formerly Cystoseira spp. – Sargassum spp.; many species are endemic), which play a key role in Mediterranean coastal ecosystems. Therefore, the attempts to restore sea urchin abundance, via seeding of juveniles from hatcheries, has further artificialized the habitats rather than contributing to the restoration of natural ecosystems. Good practices guidelines are proposed aimed at minimizing the impact of aquaculture on macrophytes. more...

The terrestrial and marine Port-Cros National (PCNP) was established in 1963; it was then made up only of the Archipelago of Port-Cros. Since 2012, it has been extended to include a vast land and sea area, including not only islands but also part of the mainland, the new PCNP (N-PCNP); the marine core area and the adjacent marine area cover approximately 120,000 ha and extend over 63 km as the crow flies, from east to west. Taxon richness is just one descriptor of biodiversity among others (e.g., functional and ecosystem diversity), and is far from being the most reliable one; however, it deserves to be taken into consideration, provided that certain prerequisites are met, because it constitutes a convenient measure of, e.g., the research effort and the diversity of habitats. The number of reported macrophyte taxa amounts to 502: 73 green algae, 316 red algae, 104 brown algae and 9 magnoliophyta and other taxa. Two new combinations are proposed: Ericaria brachycarpa var. claudiae and Gongolaria montagnei var. compressa. This gamma species diversity is far from being exceptionally high, but rather is within the norm for the Mediterranean, if we take into account the size of the area considered. The number of reported taxa per site is highly heterogeneous throughout the N-PCNP area; it is, as expected, correlated with the number of studies per site. The research effort peaked in the 1970–1980s, and then irregularly declined, which may seem surprising in this era of biodiversity launched at the 1992 Rio Summit. The exceptionally extensive database available, covering more than a century, provides the basis for a critical analysis of the concept of biodiversity, as proclaimed by the general public and the ‘greens’, which can be naive or biased, and of the concept of ‘heritage value species’, which the authors of this article consider to be a ‘toxic concept’, as opposed to ‘ordinary biodiversity’, which enables ecosystem functioning. However, this database, straddling both areas highly impacted by humans (coastal development, tourist resorts) and areas that are effectively protected, does not highlight obvious changes over time. more...

The Port-Cros National Park (PCNP), established in 1963, was one of the earliest terrestrial and marine parks in the Mediterranean Sea. From 2012, it engaged in a major redefinition and extension of its territory (N-PCNP—New Port-Cros National Park). This case is particularly interesting insofar as the protected area has been competently and efficiently managed since its creation, and protection and management measures have been strictly implemented: in the Mediterranean, the PCNP has often been considered as a benchmark. Here, we critically analyse almost 60 years of the management of the biodiversity and the human uses, with their share of successes and failures, certain operations which are today regarded as errors, and a doctrine today of a priori non-interventionism, in contrast to the doctrine in vogue in the early years. Of particular interest is the change in outlook with regard to actions favouring flagship species, such as building a tower for bats, setting up artificial nests for seabirds, and constructing an artificial reef at sea. The question of the natural arrival of the wild boar, a native species, and the hostility of the public and some species-centred scientists, is particularly instructive. We analyse these changes in the light of the ongoing trends in concepts in ecology and nature conservation, and the shift from a species-centred to an ecosystem-centred approach. It is worth emphasizing that a critical review of almost 60 years of management is a very rare exercise in a national park anywhere in the world. more...

The present study reports on observations carried out in the Tropical North Atlantic in summer and autumn 2017, documenting Sargassum aggregations using both ship-deck observations and satellite sensor observations at three resolutions (MSI-10 m, OLCI-300 m, VIIRS-750 m and MODIS-1 km). Both datasets reported that in summer, Sargassum aggregations were mainly observed off Brazil and near the Caribbean Islands, while they accumulated near the African coast in autumn. Based on in situ observations, we propose a five-class typology allowing standardisation of the description of in situ Sargassum raft shapes and sizes. The most commonly observed Sargassum raft type was windrows, but large rafts composed of a quasi-circular patch hundreds of meters wide were also observed. Satellite imagery showed that these rafts formed larger Sargassum aggregations over a wide range of scales, with smaller aggregations (of tens of m2 area) nested within larger ones (of hundreds of km2). Match-ups between different satellite sensors and in situ observations were limited for this dataset, mainly because of high cloud cover during the periods of observation. Nevertheless, comparisons between the two datasets showed that satellite sensors successfully detected Sargassum abundance and aggregation patterns consistent with in situ observations. MODIS and VIIRS sensors were better suited to describing the Sargassum aggregation distribution and dynamics at Atlantic scale, while the new sensors, OLCI and MSI, proved their ability to detect Sargassum aggregations and to describe their (sub-) mesoscale nested structure. The high variability in raft shape, size, thickness, depth and biomass density observed in situ means that caution is called for when using satellite maps of Sargassum distribution and biomass estimation. Improvements would require additional in situ and airborne observations or very high-resolution satellite imagery. more...