Operationalizing blue carbon principles in France: Methodological developments for Posidonia oceanica seagrass meadows and institutionalization.

Conservation of ecosystems is an important tool for climate change mitigation. Seagrasses, mangroves, saltmarshes and other marine ecosystems have particularly high capacities to sequester and store organic carbon (blue carbon), and are being impacted by human activities. Calls have been made to mainstream blue carbon into policies, including carbon markets. Building on the scientific literature and the French voluntary carbon standard, the 'Label Bas-Carbone', we develop the first method for the conservation of Posidonia oceanica seagrasses using carbon finance. This methodology assesses the emission reduction potential of projects that reduce physical impacts from boating and anchoring. We show how this methodology was institutionalized thanks to a tiered approach on key parameters including carbon stocks, degradation rates, and decomposition rates. We discuss future needs regarding (i) how to strengthen the robustness of the method, and (ii) the expansion of the method to restoration of seagrasses and to other blue carbon ecosystems.

Impact of sea level rise on the Mediterranean Lithophyllum byssoides rims.

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.

The Station Marine d'Endoume, Marseille: 150 years of natural history.

When marine natural sciences began to be the concern of most European scientists, in the middle of the 19th century, Marseille, in southern France, was no exception. The creation, ca. 150 years ago, of the first Zoology Laboratory of the Faculty of Sciences of Marseille took place in 1868. Under the leadership of Antoine-Fortuné Marion, it soon led to the creation of the Station Marine d'Endoume (SME) in 1889. Marion's pioneering work survived both world wars and was then taken to another dimension by Jean-Marie Pérès, head of the marine station from 1948 to 1983. This institution is still alive to date. We here inventoried all the taxa described by SME scientists (1870 to 2021) and arranged them in a public database. Three main periods of activity at the SME are described, as well as the focus made through time to different groups of taxa, selected ecosystems, or biogeographic areas. Through many examples, it was possible to document how these naturalistic, taxonomic descriptions contributed to a broader scientific knowledge within this period. Finally, we discussed trends in taxonomic and naturalistic research, based on the SME experience.

Dominance of Scorpaena maderensis among scorpaenids of littoral rocky reefs in Corsica (NW Mediterranean): Further evidence of Mediterranean Sea warming.

The population of Scorpaenidae was investigated in the Nature Reserve of Scàndula (Corsica, France, NW Mediterranean). While absent from Corsica a few decades ago, the Madeira rockfish Scorpaena maderensis is now the most abundant scorpaenid species in shallow rocky reef areas, far outnumbering Scorpaena notata and Scorpaena porcus. Considered as a subtropical species of Atlantic origin, the northward progression of the thermophilic S. maderensis and its dominance in Corsican waters is further evidence of the ongoing warming of the Mediterranean Sea.

Contribution of Posidonia oceanica meadows in the context of climate change mitigation in the Mediterranean Sea.

Coastal marine vegetation has been recently highlighted for its highly efficient carbon storage capacity. Among the sixty-four species of seagrass, Posidonia oceanica, a Mediterranean endemic species, appears to be the most effective in carbon fixation and storage. Based on new data from the study of one of the largest P. oceanica meadows in the Mediterranean Sea (100 km of coastline, 20 425 ha), and a synthesis of available data from the whole of the Mediterranean basin, the aim of this work is to evaluate the amount of carbon fixed each year by P. oceanica and sequestered in the matte, in relation with the mitigation of the impact of climate change (carbon sink). The mean total carbon fixation (blades, sheaths and rhizomes) per year varies between 33.5 and 426.6 g C.m-2 and the mean carbon sequestration (long-term sink in the matte), corresponding to the sheath and rhizome tissues, varies between 7.7 and 84.4 g C.m-2, with a clear decreasing trend according to depth because of the meadow density decrease. The synthesis of a hundred measurements made throughout the Mediterranean Sea and at depths between 0.5 and 32.0 m provides a basis for estimating the average annual carbon fixation and sequestration rate throughout the Mediterranean basin. The fixation of the blades is estimated at 1 024 t C.ha-1.yr-1, that of the sheaths at 220 t C ha-1.yr-1 and that of the rhizomes at 58 t C ha-1.yr-1; i.e. a total fixation rate of 1 302 t C ha-1.yr-1 and sequestration rate (dead sheaths and rhizomes) of 278 t C ha-1.yr-1. This annual carbon fixation represents only 0.61% on average of CO2 emissions/releases for all Mediterranean countries but in the large Mediterranean islands this fixation is on average 3.1% and can reach almost 14.4% for Corsica. Moreover, the major advantage of the P. oceanica meadow lies in its capacity to store carbon from annual carbon sequestration for centuries to millennia and can be compared to several terrestrial ecosystems considered to be efficient in carbon storage (peatlands).

The ups and downs of a canopy-forming seaweed over a span of more than one century.

Canopy-forming seaweeds constitute marine forests that deliver ecosystem services. The worldwide range shift, sharp decline or loss of many of these forests, caused by the cumulative impact of increasing human pressure and climate change, have been widely documented. Contrasting examples, reflecting higher than expected resilience, have been more rarely reported. Here, we took the opportunity of having at our disposal a two-century suite of documents (herbarium vouchers, articles) and a ~120-year observation period, dealing with a long-lived brown seaweed, Cystoseira mediterranea, along a well-explored Mediterranean coastline in the Gulf of Lions, to depict the fate of its populations. In addition, we provided baselines for future surveys, with a high degree of accuracy. The northernmost population, scattered on rare suitable substrates, gradually declined and has been extinct since the 1980s. The length of shore occupied by the southern population showed a long-term decline trend, with two sharp minima followed by partial recovery. The causes of the decline differ between sites and periods: coastal development, pollution, competition with mussels, heatwaves and exceptional storms. Overall, the Gulf of Lions populations reflects long-lasting resilience, higher than expected, and a health status that is better than that reported for many other canopy-forming seaweeds.

Response of rocky shore communities to anthropogenic pressures in Albania (Mediterranean Sea): Ecological status assessment through the CARLIT method.

The lower mid-littoral and shallow subtidal communities were studied in the district of Vlora (Albania), three years after the establishment of a Marine Protected Area, with particular attention to the long-lived species. The bioconstructions built in the mid-littoral zone by the calcified rhodobiont Lithophyllum byssoides were in poor condition and sometimes even dead. In contrast, the brown alga Cystoseira amentacea constituted lush stands. For assessing the ecological status of the studied area, the CARLIT method, based upon macroalgal communities, was applied. The observed range of ecological status was wide ('high' through 'bad') and was overall among the lowest assessed to date in the Mediterranean Sea. The occurrence of extensive sea-urchin barren-grounds, though not taken into consideration by the CARLIT index, confirmed the poor condition of large sectors of the study area. Overall, the CARLIT index is well correlated with anthropogenic pressures, as assessed by the LUSI index.

Distribution and abundance of Ostreopsis spp. and associated species (Dinophyceae) in the northwestern Mediterranean: the region and the macroalgal substrate matter.

The range of Ostreopsis (Dinophyceae), a taxon harmful to both human health and ecosystems, has spread from a tropical and subtropical range of distribution to temperate areas, such as the Mediterranean Sea. This study has evidenced widespread summer occurrence in the northwestern Mediterranean, from French Catalonia to the French Riviera and Corsica. Ostreopsis spp. are usually associated with two other dinophycean taxa, Prorocentrum lima and Coolia spp. No obvious correlation (indicative of competition and/or facilitation) between the abundance of the three taxa was evidenced. In addition to local variability, we observed regional variability, with low abundance and local absence in French Catalonia and Languedoc, which contrasted with overall abundance and blooms in Provence, the French Riviera and Corsica, especially in late summer. Possible causes for this regional variability are discussed. Furthermore, the three taxa can grow on a variety of macroalgal substrates: 34 taxa belonging to the fleshy, bushy, flat and erect morpho-functional groups. Some macroalgal species were correlated with either high or low abundance of the studied dinophycean taxa and could therefore enhance or hinder their blooming.

Towards a framework for assessment and management of cumulative human impacts on marine food webs.

Effective ecosystem-based management requires understanding ecosystem responses to multiple human threats, rather than focusing on single threats. To understand ecosystem responses to anthropogenic threats holistically, it is necessary to know how threats affect different components within ecosystems and ultimately alter ecosystem functioning. We used a case study of a Mediterranean seagrass (Posidonia oceanica) food web and expert knowledge elicitation in an application of the initial steps of a framework for assessment of cumulative human impacts on food webs. We produced a conceptual seagrass food web model, determined the main trophic relationships, identified the main threats to the food web components, and assessed the components' vulnerability to those threats. Some threats had high (e.g., coastal infrastructure) or low impacts (e.g., agricultural runoff) on all food web components, whereas others (e.g., introduced carnivores) had very different impacts on each component. Partitioning the ecosystem into its components enabled us to identify threats previously overlooked and to reevaluate the importance of threats commonly perceived as major. By incorporating this understanding of system vulnerability with data on changes in the state of each threat (e.g., decreasing domestic pollution and increasing fishing) into a food web model, managers may be better able to estimate and predict cumulative human impacts on ecosystems and to prioritize conservation actions.

Hacia un Marco de Trabajo para la Evaluación y el Manejo de los Impactos Humanos Acumulativos sobre las Redes Alimenticias Marinas Resumen El manejo efectivo con base en los ecosistemas requiere entender la respuesta de los ecosistemas a múltiples amenazas humanas en lugar de enfocarse en amenazas individuales. Para entender holísticamente la respuesta de los ecosistemas a las múltiples amenazas antropogénicas es necesario saber cómo estas amenazas afectan a los diferentes componentes dentro de los ecosistemas y cómo alteran finalmente el funcionamiento de los ecosistemas. Usamos el estudio de caso de la red alimenticia del pasto marino del Mediterráneo (Posidonia oceanica) y la obtención de conocimiento de expertos en una aplicación de los pasos iniciales de un método para la evaluación de los impactos humanos acumulativos sobre las redes alimenticias. Produjimos un modelo de red alimenticia de pastos marinos, determinamos las principales relaciones tróficas, identificamos a las principales amenazas para los componentes de la red y evaluamos la vulnerabilidad de los componentes a esas amenazas. Algunas amenazas tuvieron impactos altos (p. ej.: infraestructura costera) o bajos (p. ej.: escorrentía agrícola) sobre todos los componentes de la red, mientras que otros (p. ej.: carnívoros introducidos) tuvieron impactos muy diferentes sobre cada componente. Partir al ecosistema en sus componentes nos permitió identificar amenazas no vistas previamente y reevaluar la importancia de las amenazas percibidas comúnmente como mayores. Al incorporar este entendimiento de la vulnerabilidad del sistema con datos sobre los cambios en el estado de cada amenaza (p. ej.: disminución de la contaminación doméstica e incremento de la pesca) al modelo de red alimenticia, los manejadores pueden ser capaces de estimar y predecir de mejor manera los impactos humanos acumulativos sobre los ecosistemas y priorizar las acciones de conservación.

Competition between the invasive macrophyte Caulerpa taxifolia and the seagrass Posidonia oceanica: contrasting strategies.

BACKGROUND: Plant defense strategy is usually a result of trade-offs between growth and differentiation (i.e. Optimal Defense Theory--ODT, Growth Differentiation Balance hypothesis--GDB, Plant Apparency Theory--PAT). Interaction between the introduced green alga Caulerpa taxifolia and the endemic seagrass Posidonia oceanica in the Mediterranean Sea offers the opportunity to investigate the plausibility of these theories. We have accordingly investigated defense metabolite content and growth year-round, on the basis of an interaction gradient. RESULTS: When in competition with P. oceanica, C. taxifolia exhibits increased frond length and decreased Caulerpenyne--CYN content (major terpene compound). In contrast, the length of P. oceanica leaves decreases when in competition with C. taxifolia. However, the turnover is faster, resulting in a reduction of leaf longevity and an increase on the number of leaves produced per year. The primary production is therefore enhanced by the presence of C. taxifolia. While the overall concentration of phenolic compounds does not decline, there is an increase in some phenolic compounds (including ferulic acid and a methyl 12-acetoxyricinoleate) and the density of tannin cells. CONCLUSION: Interference between these two species determines the reaction of both, confirming that they compete for space and/or resources. C. taxifolia invests in growth rather than in chemical defense, more or less matching the assumptions of the ODT and/or PAT theories. In contrast, P. oceanica apparently invests in defense rather than growth, as predicted by the GDB hypothesis. However, on the basis of closer scrutiny of our results, the possibility that P. oceanica is successful in finding a compromise between more growth and more defense cannot be ruled out.

Does coverage matter at mesoscale within a Posidonia oceanica seagrass meadow?

Despite the fact that coverage is one of the most widely used descriptors for seagrass meadows, the spatial structure of coverage at mesoscale has not often been taken into account. The present work investigates the structure of P. oceanica coverage at mesoscale and its possible relationship with several factors (depth, type of substrate, relative level within the meadow, type of shoot density and level of anthropic pressure). Five classes of coverage structure are delineated within P. oceanica meadows and statistical analysis of the data did not provide evidence of a link with the factors taken into consideration. This result could be explained by the prevailing role of endogenic processes in the structuring of the P. oceanica meadow.

Unexpected response of the seagrass Posidonia oceanica to a warm-water episode in the north western Mediterranean Sea.

The response of Posidonia oceanica (Linnaeus) Delile to the warm-water episode of summer 1999 was studied by means of the technique of lepidochronology. Study sites include three sites affected by the mass mortality event of benthic invertebrates and one not affected. The results showed a significant decline in some parameters (number of leaves and/or rhizome growth) for the three sites affected by the mass mortality event for the year following the warm-water episode (1999-2000). A similar decline was not observed for the unaffected site. The fact that high temperatures could have a negative impact on deep Posidonia oceanica near its cold limit of distribution is an unexpected result.

Biological pollution in the Mediterranean Sea: invasive versus introduced macrophytes.

The authors have listed 85 species of macrophytes that have probably been introduced to the Mediterranean. Among them, nine species can be considered as invasive, i.e., playing a conspicuous role in the recipient ecosystems, taking the place of keystone species and/or being economically harmful: Acrothamnion preissii, Asparagopsis armata, Lophocladia lallemandii, Womersleyella setacea (Rhodophyta), Sargassum muticum, Stypopodium schimperi (Fucophyceae), Caulerpa racemosa, Caulerpa taxifolia and Halophila stipulacea (Plantae). These data fit well the Williamson and Fitter's "tens rule", which states that, on average, 1 out of 10 introduced species becomes invasive. Though some features (e.g. life traits, geographical origin) can increase the likelihood of a successful invasion, the success of invaders is far from being predictable. Since the beginning of the 20th century, the number of introduced species to the Mediterranean has nearly doubled every 20 years. Should these kinetics continue, and according to the tens rule, it can be expected that 5-10 newly introduced macrophytes shall become invasive in the next 20 years.