Positive feedback loop between introductions of non-native marine species and cultivation of oysters in Europe.

With globalization, agriculture and aquaculture activities are increasingly affected by diseases that are spread through movement of crops and stock. Such movements are also associated with the introduction of non-native species via hitchhiking individual organisms. The oyster industry, one of the most important forms of marine aquaculture, embodies these issues. In Europe disease outbreaks affecting cultivated populations of the naturalized oyster Crassostrea gigas caused a major disruption of production in the late 1960s and early 1970s. Mitigation procedures involved massive imports of stock from the species' native range in the northwestern Pacific from 1971 to 1977. We assessed the role stock imports played in the introduction of non-native marine species (including pathogens) from the northwestern Pacific to Europe through a methodological and critical appraisal of record data. The discovery rate of non-native species (a proxy for the introduction rate) from 1966 to 2012 suggests a continuous vector activity over the entire period. Disease outbreaks that have been affecting oyster production since 2008 may be a result of imports from the northwestern Pacific, and such imports are again being considered as an answer to the crisis. Although successful as a remedy in the short and medium terms, such translocations may bring new diseases that may trigger yet more imports (self-reinforcing or positive feedback loop) and lead to the introduction of more hitchhikers. Although there is a legal framework to prevent or reduce these introductions, existing procedures should be improved.

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.

Fronts, jumps and secondary introductions suggested as different invasion patterns in marine species, with an increase in spread rates over time.

Not all introduced (invasive) species in a region will spread from a single point of introduction. Long-distance dispersal or further introductions can obscure the pattern of spread, but the regional importance of such processes is difficult to gauge. These difficulties are further compounded when information on the multiple scale process of invasive species range expansion is reduced to one-dimensional estimates of spread (e.g. km yr(-1)). We therefore compared the results of two different metrics of range expansion: maximum linear rate of spread and accumulation of occupied grid squares (50 x 50 km) over time. An analysis of records for 54 species of introduced marine macrophytes in the Mediterranean and northeast Atlantic revealed cases where the invasion process was probably missed (e.g. Atlantic Bonnemaisonia hamifera) and suggested cases of secondary introductions or erratic jump dispersal (Dasysiphonia sp. and Womersleyella setacea). A majority of species analysed showed evidence for an accumulation of invaded sites without a clear invasion front. Estimates of spread rate are increasing for more recent introductions. The increase is greater than can be accounted for by temporally varying search effort and implies a historical increase in vector efficiency and/or a decreased resistance of native communities to invasion.

Biological invasion: Long term monitoring of the macroalgal flora of a major European harbor complex.

The marine macroflora of the ports of Le Havre and Antifer have been studied by citizen scientists since the late 1970s. In addition to analysis of the previous results, the field study was extended from 2010 to 2018. A total of 97 and 62 macroalgae were identified, respectively, including 14 NIS (non-indigenous species), the latter number being certainly an underestimate since microscopic species were not exhaustively sought and given the high number of cryptogenic species encountered. No new primary introduction of NIS for the NE Atlantic has been detected since the late 1970s. The origin of NIS and vector(s) of introduction were investigated. For all NIS, the donor region is the Indo-Pacific. Two likely vectors have been identified: maritime traffic for earlier introductions, and shellfish imports for more recent ones. The role of these ports and maritime traffic in the processes of introduction is discussed.

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.

The Caulerpa racemosa invasion: a critical review.

Caulerpa racemosa var. cylindracea is a marine Chlorophyta introduced into the Mediterranean Sea from south-western Australia. Since 1990, it has been invading the Mediterranean Sea and the Canary Islands, raising ecological problems. Although this invasion event can be considered as one of the most serious in the history of species introduced into the Mediterranean Sea, C. racemosa has not triggered as much attention as the famous "killer alga"Caulerpa taxifolia. The aim of the present study was: (i) to summarize the current state of knowledge with regard to the distribution, the various biological and ecological characteristics of the introduced C. racemosa and its impact on the Mediterranean coastal environment; (ii) to discuss the various hypotheses regarding the explanation for its rapid and successful spread; (iii) to investigate the disparity in the treatment of C. racemosa and Caulerpa taxifolia invasions; and (iv) to outline future research needs.

Macroalgal survival in ballast water tanks.

Despite a large amount of research into invasive species and their introductions, there have been no studies focused on macroalgal transport in ballast water. To address this, we collected replicate samples of ballast water from 12 ships in two Mediterranean harbours (Naples and Salerno). Filtered samples were kept in culture for a month at Mediterranean mean conditions (18 degrees C, 12:12h LD, 60micromol photons m(-2)s(-1)). Fifteen macroalgal taxa were cultured and differed according to the geographic origin of the ballast water. Most of the cultured algae were widely distributed species (e.g. Ulva spp. and Acinetospora-phase). However, Ulva ohnoi Hiraoka and Shimada, described from Japan, was hitherto unknown in the Mediterranean Sea. We show for the first time that ballast water can be an important vector for the transport of microscopic stages of macroalgae and that this can be a vector for the introduction of alien species.

An integrated method to evaluate and monitor the conservation state of coralligenous habitats: The INDEX-COR approach.

A new method based on photographic sampling coupled with in situ observations was applied to 53 stations along the French Mediterranean coast, to assess the integrity of coralligenous reefs affected by different levels of anthropogenic pressure. The conservation state of the assemblages characterizing these habitats was then assessed by an index - the INDEX-COR - that integrates three metrics: (i) the sensitivity of the taxa to organic matter and sediment deposition, (ii) the observable taxonomic richness, and (iii) the structural complexity of the assemblages. The sensitivity of INDEX-COR was tested and showed good correlation with the Level of Pressure calculated for each station according to expert judgment and field observations.

An ecosystem-based approach to assess the status of Mediterranean algae-dominated shallow rocky reefs.

A conceptual model was constructed for the functioning the algae-dominated rocky reef ecosystem of the Mediterranean Sea. The Ecosystem-Based Quality Index (reef-EBQI) is based upon this model. This index meets the objectives of the EU Marine Strategy Framework Directive. It is based upon (i) the weighting of each compartment, according to its importance in the functioning of the ecosystem; (ii) biological parameters assessing the state of each compartment; (iii) the aggregation of these parameters, assessing the quality of the ecosystem functioning, for each site; (iv) and a Confidence Index measuring the reliability of the index, for each site. The reef-EBQI was used at 40 sites in the northwestern Mediterranean. It constitutes an efficient tool, because it is based upon a wide set of functional compartments, rather than upon just a few species; it is easy and inexpensive to implement, robust and not redundant with regard to already existing indices.

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.

First assessment of the Caulerpa racemosa (Caulerpales, Chlorophyta) invasion along the French Mediterranean coast.

The introduced green alga Caulerpa racemosa var. cylindracea has been rapidly spreading in the Mediterranean Sea since 1990. It was first observed in France in 1997 (Marseilles). In early 2004, the stretch of the French Mediterranean coastline and the surface area affected by the invasion were estimated at about 83 km and 4014 ha, respectively. The depth range of colonized areas was usually 10-35 m depth. Shallow (0-10 m) and deep (down to 40 m) dense meadows were rarely observed. In contrast to the dead matte of Posidonia oceanica, which constituted the most widely colonized substratum, dense P. oceanica meadows and fine sand with large ripple-marks were not invaded. Few rocky areas were colonized and coarse sand bottoms were usually colonized below 20 m depth. All the colonized areas were exposed to human activities and more than 40% were fishing areas. Mild climate, suitable substrata, presence of vectors of dispersal and absence of efficient biological control make the French Mediterranean coast particularly vulnerable to the further spread of the alga.

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.

Unexpected abundance and long-term relative stability of the brown alga Cystoseira amentacea, hitherto regarded as a threatened species, in the north-western Mediterranean Sea.

Cystoseira amentacea is a Mediterranean endemic alga thriving on very shallow rocky substrates. It has been considered as a threatened species, having experienced a steady decline and is therefore protected by international conventions. The historical distribution of the species has been assessed along the French Mediterranean coast, on the basis of 467 articles and herbarium vouchers. We have produced an accurate map of its current distribution and abundance along 1832 km of coastline, through in situ surveys. C. amentacea was observed along 1125 km of shoreline, including 33% of almost continuous or continuous belt. In most of its range, there is no evidence of loss, except in 4 areas of Provence, French Riviera and Corsica. A significant relation was found between the absence or low abundance of C. amentacea and the vicinity of ports and large sewage outfalls. The status of conservation of the species should therefore be reassessed.

TAXONOMIC REVISION AND GEOGRAPHIC DISTRIBUTION OF THE SUBGENUS SARGASSUM (FUCALES, PHAEOPHYCEAE) IN THE WESTERN AND CENTRAL PACIFIC ISLANDS BASED ON MORPHOLOGICAL AND MOLECULAR ANALYSES(1).

The species diversity of the subgenus Sargassum was reassessed for the southwestern Pacific with special focus on the Solomon Islands, Vanuatu, Fiji, and Wallis. Five taxa were recognized on the basis of morphological characters and corroborated by DNA analyses of the nuclear internal transcribed spacer 2 (ITS-2), chloroplastic partial rbcLS-operon, and mitochondrial cox3. Based on the study of diagnoses and type specimens, four taxa were identified to S. polyphyllum J. Agardh, S. polycystum C. Agardh, S. aquifolium (Turner) C. Agardh, and S. ilicifolium (Turner) C. Agardh, while one taxon remained unidentified. We present a key for identification that includes detailed descriptions of the species and illustrations of their morphological variability. In light of our findings, we propose to consider several new synonymies for S. aquifolium, S. ilicifolium, S. polycystum, and S. polyphyllum. We also include a review of Sargassum floras from Samoa, Tonga, and Nauru and discuss species distribution in the southwest and central Pacific. Finally, DNA phylogenies pointed to the polyphyly of section Acanthocarpicae and underlined the need for a new section in which to place S. polycystum, S. herporhizum Setch. et N. L. Gardner, and S. stolonifolium Phang et T. Yoshida. The new section Polycystae Mattio et Payri is described to fit species of the subgenus Sargassum with stolon-like branches.