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The Salento peninsula in southern Italy (Mediterranean Sea) is a strip of land between the Adriatic and the Ionian Seas, both characterized by local regimes of currents, different geological and physical backgrounds, and quite diversified fauna. In this area, specimens of the sea cucumber Holothuria tubulosa (Brünnich, 1768) (Echinodermata) were collected at four stations in the spring and autumn of 2020 to investigate a possible symbiotic association with the inquiline fish Carapus acus (Brünnich, 1768). Among the collected holothurians, five pearlfish specimens were found in the body cavity of four H. tubulosa collected at 10 m of depth, in autumn, at “Grotta Verde” in Marina di Andrano, Lecce (Ionian Sea). More than half of the sea cucumbers from the latter station hosted the symbiont, suggesting the presence of a shallow population of C. acus inhabiting this coastal area. Furthermore, morphometric analysis carried out on the collected fish helped to shed light on the population dynamics characterizing this neglected species. This is the first report of C. acus from Apulian waters, allowing us to unite previously disjoined areas and providing essential baseline knowledge for planning future in-depth analysis of this difficult-to-study fish in a geographical area that is strategic in terms of the conservation of Mediterranean biodiversity. Furthermore, the range of preferred host species is extended, as C. acus was previously known to prefer other sea cucumber species such as Parastichopus regalis (Cuvier, 1817) instead of H. tubulosa. Finally, the finding of C. acus in a single station and in only one season is not trivial and delivers baseline useful information for future conservation purposes.

Abstract Cold-water coral (CWC) reefs of the Angolan margin (SE Atlantic) are dominated by Desmophyllum pertusum and support a diverse community of associated fauna, despite hypoxic conditions. In this study, we use carbon and nitrogen stable isotope analyses (δ13C and δ15N) to decipher the trophic network of this relatively unknown CWC province. Although fresh phytodetritus is available to the reef, δ15N signatures indicate that CWCs (12.90 ± 1.00 ‰) sit two trophic levels above Suspended Particulate Organic Matter (SPOM) (4.23 ± 1.64 ‰) suggesting that CWCs are highly reliant on an intermediate food source, which may be zooplankton. Echinoderms and the polychaete Eunice norvegica occupy the same trophic guild, with high δ13C signatures (-14.00 ± 1.08 ‰) pointing to a predatory feeding behavior on CWCs and sponges, although detrital feeding on 13C enriched particles might also be important for this group. Sponges presented the highest δ15N values (20.20 ± 1.87 ‰), which could be due to the role of the sponge holobiont and bacterial food in driving intense nitrogen cycling processes in sponges’ tissue, helping to cope with the hypoxic conditions of the reef. Our study provides first insights to understand trophic interactions of CWC reefs under low-oxygen conditions.

Marine bioprospecting is a dynamic research field that explores the oceans and their biodiversity as noteworthy sources of new bioactive compounds. Anthozoans are marine animals belonging to the Cnidaria phylum characterized by highly specialized mechanosensory cells used both for defence against predators and prey capture. Here, high concentration of cnidocysts have been isolated from the Mediterranean zoanthid coral Parazoanthus axinellae (Schmidt, 1862) and their antimicrobial potential has been investigated. The cnidocyst extract exerted significant antibacterial activity against some human pathogens capable of developing resistance to conventional antibiotics such as Streptococcus agalactiae and Coccus sp., and against several Vibrio species, including some microbial strains for humans and farmed fish, such as Vibrio alginolyticus, Vibrio anguillarum, Vibrio fischeri, Vibrio harveyi, and Vibrio vulnificus. Results have been discussed in light of both the ecological aspects and biotechnological value of the cnidocyst extract in the nutritional, nutraceutical, and pharmaceutical fields.

Biological invasions, resulting from human activities, exert substantial impacts on ecosystems worldwide. This review focuses on marine invasive alien species (IAS) in Europe, examining the current state, proposing strategies to address the problem, and offering recommendations for enhanced management. Effective management of biological invasions relies on accessible, accurate data to inform decision-making. Information systems such as the European Alien Species Information Network (EASIN), Aquatic Non-Indigenous and Cryptogenic Species (AquaNIS), and World Register of Introduced Marine Species (WriMS) provide comprehensive databases on IAS, but their sustainability requires long-term maintenance, continuous updates, and support. Most countries lack specific monitoring programs for marine IAS, and standardization and improvement of monitoring methods are needed. Port monitoring plays a vital role in the early detection of new arrivals, and recent advancements in molecular techniques show promise for effective IAS monitoring. Risk screening tools are commonly employed to rank taxa based on their invasiveness potential in European regions, but variations in protocols can yield inconsistent results. European impact assessments highlight resource competition, novel habitat creation, and predation as primary mechanisms for negative impacts on biodiversity, while the creation of novel habitats represents a key mechanism for positive impacts. Preventing IAS introductions is critical, and measures such as ballast water treatment systems are implemented to reduce the likelihood of marine introductions. However, understanding introduction pathways remains uncertain for many IAS. Eradication and control efforts for marine IAS have limited success, emphasizing the need for enhanced biosecurity measures. Climate change, especially ocean warming, can intensify IAS impacts on native species and ecosystems. In climate change hotspots, some tropical aliens may, however, compensate for the loss of thermally sensitive natives with similar traits. Therefore, it is imperative to consider the interactions between climate change and IAS in developing effective management and conservation strategies. Enhancing IAS management in Europe entails i) securing adequate funding, ii) expanding the list of IAS of Union Concern to adequately cover marine invasions, iii) learning from countries with successful biosecurity practices, iv) sustaining information systems, v) improving monitoring and early warning systems with innovative technologies, vi) enhancing prediction models, vii) conducting integrated impact assessments and mapping cumulative IAS impacts, and vii) considering the potential benefits of IAS in ecosystem functioning and services.

Abstract Understanding the impacts of microplastics on living organisms in aquatic habitats is one of the hottest research topics worldwide. Despite increased attention, investigating microplastics in underwater environments remains a problematic task, due to the ubiquitous occurrence of microplastic, its multiple modes of interactions with the biota, and to the diversity of the synthetic organic polymers composing microplastics in the field. Several studies on microplastics focused on marine invertebrates, but to date, the benthic sea slugs (Mollusca, Gastropoda, Heterobranchia) were not yet investigated. Sea slugs are known to live on the organisms on which they feed on or to snack while gliding over the sea floor, but also as users of exogenous molecules or materials not only for nutrition. Therefore, they may represent a potential biological model to explore new modes of transformation and/or management of plastic, so far considered to be a non-biodegradable polymer. In this study we analysed the stomachal content of Bursatella leachii, an aplysiid heterobranch living in the Mar Piccolo, a highly polluted coastal basin near Taranto, in the northern part of the Ionian Sea. Microplastics were found in the stomachs of all the six sampled specimens, and SEM/EDX analyses were carried out to characterize the plastic debris. The SEM images and EDX spectra gathered here should be regarded as a baseline reference database for future investigations on marine Heterobranchia and their interactions with microplastics.

This manuscript reports on four new jellyfish species in the Sea of Marmara (Turkey) (the scyphozoans Mawia benovici and Drymonema dalmatinum, hydrozoan Aequorea forskalea, and thaliaceanSalpa maxima) based on plankton collections made in the years 2019–2021. This is the first record of Mawia benovici in both Turkish coastal areas and the Sea of Marmara. The jellyfish that was previously re-corded as Drymonema sp. in the repetition Sea of Marmara was identified at the species level as D. dalmatinum. Furthermore, their possible introduction pathways are briefly discussed here

Knowledge of the reproductive strategy is a key prerequisite to predict population dynamics and potential invasiveness of both native and non-indigenous outbreak-forming species. In 2014 the Lessepsian upside-down jellyfish Cassiopea andromeda reached the harbor of Palermo (NW Sicily, Thyrrenian Sea), to date its established westernmost outpost in the Mediterranean Sea. To predict C. andromeda reproductive success in its novel habitat, gonad histology was carried out to record the number and size of mature and immature oocytes. Both male and female simultaneously presented gametes at all stages of development suggesting an asynchronous, yet apparently continuous, reproduction strategy. Indeed, oogenesis was observed throughout the year from pre-vitellogenic, vitellogenetic, and late-vitellogenetic to mature oocytes suggesting multiple reproductive events, as known in other Mediterranean Rhizostomeae. Oocytes were found from May to December, with two seasonal peaks of abundance (late spring = 392 and autumn = 272), suggesting imminent spawning events. Further, jellyfish size varied significantly throughout the year, with maximum diameter (up to 24 cm) in summer, and minimum diameter (6 cm) in winter. Small-sized jellyfish in winter belong to the new cohort, most probably arising from intense summer strobilation of polyps. Late spring fertilization, planula development, and metamorphosis, followed by polyp strobilation in the summer months, may explain the late appearance of a new jellyfish cohort, likely coincident with that recorded throughout winter.

A dual nutrition mode (i.e., mixotrophy) can be advantageous for alien species in a new environment. In Cearà (Brazil), the symbiotic jellyfish Cassiopea andromeda is rapidly spreading under diverse environmental conditions across natural and human-altered coastal habitats, such as mangroves and shrimp farms. Here we report on the trophic ecology of the alien upside-down jellyfish sampled in these two contrasting coastal habitats during the dry (July–October) and rainy (January–April) seasons, investigated by means of organic biomarkers (lipids, carbohydrates) and bulk tissue stable isotope (δ15N and δ13C) analyses. Total lipid content of jellyfish gonads was generally higher in shrimp farms, whereas no significant difference in carbohydrate concentration was found in jellyfish tissues from the two different habitats. Similarly, there were no significant differences in the δ15N values of jellyfish tissues from the two contrasting habitats, whereas the δ13C values were higher in jellyfish from shrimp farms. Overall, the higher carbon-enriched value in aquaculture ponds supports the hypothesis of differences of available food sources compared to the natural mangrove habitats, where food availability exhibits a stronger seasonality. In fact, aquaculture ponds are characterized by human-driven regular food supply, leading to more stable trophic conditions and to enhanced growth, lipid production, and gonadal output of C. andromeda jellyfish. This investigation may contribute to predicting how Cassiopea mixotrophy may contribute to explaining its differential success in different habitats.

Cassiopea andromeda (Forsskål, 1775), commonly found across the Indo-Pacific Ocean, the Red Sea, and now also in the warmest areas of the Mediterranean Sea, is a scyphozoan jellyfish that hosts autotrophic dinoflagellate symbionts (family Symbiodiniaceae). Besides supplying photosynthates to their host, these microalgae are known to produce bioactive compounds as long-chain unsaturated fatty acids, polyphenols, and pigments, including carotenoids, with antioxidant properties and other beneficial biological activities. By the present study, a fractionation method was applied on the hydroalcoholic extract from two main body parts (oral arms and umbrella) of the jellyfish holobiont to obtain an improved biochemical characterization of the obtained fractions from the two body parts. The composition of each fraction (i.e., proteins, phenols, fatty acids, and pigments) as well as the associated antioxidant activity were analyzed. The oral arms proved richer in zooxanthellae and pigments than the umbrella. The applied fractionation method was effective in separating pigments and fatty acids into a lipophilic fraction from proteins and pigment–protein complexes. Therefore, the C. andromeda–dinoflagellate holobiont might be considered as a promising natural source of multiple bioactive compounds produced through mixotrophic metabolism, which are of interest for a wide range of biotechnological applications.

Temperature is a fundamental variable for all biological processes. It influences the metabolism and tolerance limits of all living organisms, affecting species phenology and distribution patterns. It also facilitates the spread of non-indigenous species and the proliferation and expansion of native outbreak-forming species. Pennaria disticha is a colonial benthic cnidarian reported to be invasive in different Indian and Pacific coastal areas, as well as a harmful member of fouling communities found in Mediterranean marine aquaculture farms. Using the most basal functional trait (i.e., thermal tolerance), we explored the potential of P. disticha to colonize different habitats across the Mediterranean Sea in future warming scenarios. Respiration rate was measured as a proxy of P. disticha metabolism under 12 different experimental temperatures. The obtained thermal tolerance dataset was used to create a thermal performance curve (TPC). We then scaled modeled curve to occurrence probability to map species potential metabolic habitat suitability and phenological shifts within the Mediterranean Sea when subjected to different warming scenarios. Prediction maps for future climatic conditions showed a potential temporal and spatial expansion of P. disticha in the Western and Central Mediterranean. The present data increases our understanding of the ecological performance and potential distribution of an invasive and outbreak-forming species. This information will contribute to the development of early warning systems and to the design and implementation of risk assessment and management plans.

Medusae of Turritopsis dohrnii undergo reverse development in response to physical damage, adverse environmental conditions, or aging. Senescent, weakened or damaged medusae transform into a cluster of poorly differentiated cells (known as the cyst stage), which metamorphose back into a preceding life cycle stage, the polyp. During the metamorphosis, cell transdifferentiation occurs. The cyst represents the intermediate stage between a reverting medusa and a healthy polyp, during which cell transdifferentiation and tissue reorganization take place. Here we characterize and compare the transcriptomes of the polyp and newborn medusa stages of T. dohrnii with that of the cyst, to identify biological networks potentially involved in the reverse development and transdifferentiation processes. The polyp, medusa and cyst of T. dohrnii were sequenced through Illumina RNA-sequencing and assembled using a de novo approach, resulting in 92,569, 74,639 and 86,373 contigs, respectively. The transcriptomes were annotated and comparative analyses among the stages identified biological networks that were significantly over-and under-expressed in the cyst as compared to the polyp and medusa stages. Biological processes that occur at the cyst stage such as telomerase activity, regulation of transposable elements and DNA repair systems, and suppression of cell signaling pathways, mitotic cell division and cellular differentiation and development may be involved in T. dohrnii’s reverse development and transdifferentiation. Our results are the first attempt to understand T. dohrnii’s life-cycle reversal at the genetic level, and indicate possible avenues of future research on developmental strategies, cell transdifferentiation, and aging using T. dohrnii as a non-traditional in vivo system.

The extant graptolite Rhabdopleura recondita has been so far recorded only as inhabiting a bryozoan skeleton. Its larval settlement and metamorphosis are possible in the absence of a bryozoan zoarium, whereas further colony development may require that the larva is hidden inside the bryozoan host. This dependence may constrain the development of R. recondita tube and tubaria compared to the other Rhabdopleura species that develop without a host. We report here on larval settlement and metamorphosis in the absence/presence of a bryozoan host skeleton. We also make the first attempt to test the phenotypical response of R. recondita tubes and tubaria under variable hydrodynamic regimes in laboratory conditions. After 40 days, no significant variation was detected in the number or length of the newly formed tubes. These findings suggest that R. recondita eventually resides in a narrow velocity range and that tube and tubarium development is largely invariable.

The moon jellyfish Aurelia coerulea (Scyphozoa) is one of the most common and largest jellyfish inhabiting coastal lagoons, confined bays, and marinas of temperate and subtropical coastal waters. The annual population dynamics of A. coerulea along with some bacterial parameters (bacterial size and biomass, total coliforms, faecal coliforms, intestinal enterococci, culturable Vibrio spp., and culturable bacteria at 37 °C), sea surface temperature (SST), salinity, and an array of nutrients (ammonia, nitrites, nitrates, phosphates, silicates, total nitrogen, and total phosphorus) were assessed in the Varano lagoon (Adriatic Sea) that is subject to anthropogenic pollution. Statistical analyses revealed that jellyfish outbreaks and their consequent biomass deposition significantly correlated to seawater temperature, total nitrogen, phosphates, and ammonia concentrations while negative correlations appeared with nitrite and nitrate concentrations. In addition, bacterial biomass and Vibrio abundance correlated with each other and temperature, jellyfish density, and total nitrogen. These findings suggest that environmental changes could trigger the occurrence of jellyfish bursts in the lagoon which, in turn, may act as one of the central drivers of processes regulating some bacterial components. The positive relationship between jellyfish flush-and-crash dynamics and SST suggests that ongoing global warming will seemingly increase jellyfish outbreaks.

Biodiversity is a broad concept that encompasses the diversity of nature, from the genetic to the habitat scale, and ensures the proper functioning of ecosystems. The Mediterranean Sea, one of the world’s most biodiverse marine basins, faces major threats, such as overexploitation of resources, pollution and climate change. Here we provide the first multi-taxa inventory of marine organisms and coastal terrestrial flora recorded in southeastern Salento (Ionian Sea, Italy), realized during the project “Biodiversity MARE Tricase”, which provided the first baseline of species living in the area. Sampling was carried out by SCUBA and free diving, fishing gears, and citizen science from 0 to 70 m. Overall, 697 taxa were found between March 2016 and October 2017, 94% of which were identified to the species level. Of these, 19 taxa represented new records for the Ionian Sea (36 additional new records had been reported in previous publications on specific groups, namely Porifera and Mollusca Heterobranchia), and two findings represented the easternmost records in the Mediterranean Sea (Helicosalpa virgula and Lampea pancerina). For eight other taxa, our findings represented the only locality in the Ionian Sea, besides the Straits of Messina. In addition to the species list, phenological events (e.g., blooms, presence of reproductive traits and behaviour) were also reported, with a focus on gelatinous plankton. Our results reveal that even for a relatively well-known area, current biodiversity knowledge may still be limited, and targeted investigations are needed to fill the gaps. Further research is needed to understand the distribution and temporal trends of Mediterranean biodiversity and to provide baseline data to identify ongoing and future changes.

Ecological profiling of non-native species is essential to predict their dispersal and invasiveness potential across different areas of the world. Cassiopea is a monophyletic taxonomic group of scyphozoan mixotrophic jellyfish including C. andromeda, a recent colonizer of sheltered, shallow-water habitats of the Mediterranean Sea, such as harbors and other light-limited, eutrophic coastal habitats. To assess the ecophysiological plasticity of Cassiopea jellyfish and their potential to spread across the Mare Nostrum by secondary introductions, we investigated rapid photosynthetic responses of jellyfish to irradiance transitions-from reduced to increased irradiance conditions (as paradigm of transition from harbors to coastal, meso/oligotrophic habitats). Laboratory incubation experiments were carried out to compare oxygen fluxes and photobiological variables in Cassiopea sp. immature specimens pre-acclimated to low irradiance (PAR = 200 μmol photons m-2 s-1) and specimens rapidly exposed to higher irradiance levels (PAR = 500 μmol photons m-2 s-1). Comparable photosynthetic potential and high photosynthetic rates were measured at both irradiance values, as also shown by the rapid light curves. No significant differences were observed in terms of symbiont abundance between control and treated specimens. However, jellyfish kept at the low irradiance showed a higher content in chlorophyll a and c (0.76±0.51SD vs 0.46±0.13SD mg g-1 AFDW) and a higher Ci (amount of chlorophyll per cell) compared to jellyfish exposed to higher irradiance levels. The ratio between gross photosynthesis and respiration (P:R) was >1, indicating a significant input from the autotrophic metabolism. Cassiopea sp. specimens showed high photosynthetic performances, at both low and high irradiance, demonstrating high potential to adapt to sudden changes in light exposure. Such photosynthetic plasticity, combined with Cassiopea eurythermal tolerance and mixotrophic behavior, jointly suggest the upside-down jellyfish as a potentially successful invader in the scenario of a warming Mediterranean Sea.

Marine invertebrates represent a vast, untapped source of bioactive compounds. Cnidarians are represented by nearly 10,000 species that contain a complex mixture of venoms, collagen, and other bioactive compounds, including enzymes, oligosaccharides, fatty acids, and lipophilic molecules. Due to their high abundance in coastal waters, several jellyfish taxa may be regarded as candidate targets for the discovery of novel lead molecules and biomaterials and as a potential source of food/feed ingredients. The moon jellyfish Aurelia coerulea is one of the most common jellyfish worldwide and is particularly abundant in sheltered coastal lagoons and marinas of the Mediterranean Sea, where it first appeared—as an alien species—in the last century, when Pacific oyster cultivation began. In the present study, the antioxidant and lysozyme antibacterial activities associated with extracts from different medusa compartments—namely the umbrella, oral arms, and secreted mucus—were investigated. Extracts from the oral arms of A. coerulea displayed significant antioxidant activity. Similarly, lysozyme-like activity was the highest in extracts from oral arms. These findings suggest that A. coerulea outbreaks may be used in the search for novel cytolytic and cytotoxic products against marine bacteria. The geographically wide occurrence and the seasonally high abundance of A. coerulea populations in coastal waters envisage and stimulate the search for biotechnological applications of jellyfish biomasses in the pharmaceutical, nutritional, and nutraceutical sectors.

Increasing frequency of native jellyfish proliferations and massive appearance of non-indigenous jellyfish species recently concur to impact Mediterranean coastal ecosystems and human activities at sea. Nonetheless, jellyfish biomass may represent an exploitable novel resource to coastal communities, with reference to its potential use in the pharmaceutical, nutritional, and nutraceutical Blue Growth sectors. The zooxanthellate jellyfish Cassiopea andromeda, Forsskål, 1775 (Cnidaria, Rhizostomeae) entered the Levant Sea through the Suez Canal and spread towards the Western Mediterranean to reach Malta, Tunisia, and recently also the Italian coasts. Here we report on the biochemical characterization and antioxidant activity of C. andromeda specimens with a discussion on their relative biological activities. The biochemical characterization of the aqueous (PBS) and hydroalcoholic (80% ethanol) soluble components of C. andromeda were performed for whole jellyfish, as well as separately for umbrella and oral arms. The insoluble components were hydrolyzed by sequential enzymatic digestion with pepsin and collagenase. The composition and antioxidant activity of the insoluble and enzymatically digestible fractions were not affected by the pre-extraction types, resulting into collagen- and non-collagen-derived peptides with antioxidant activity. Both soluble compounds and hydrolyzed fractions were characterized for the content of proteins, phenolic compounds, and lipids. The presence of compounds coming from the endosymbiont zooxanthellae was also detected. The notable yield and the considerable antioxidant activity detected make this species worthy of further study for its potential biotechnological sustainable exploitation.

Science is addressing global societal challenges, and due to limitations in research financing, scientists are turning to the public at large to jointly tackle specific environmental issues. Citizens are therefore increasingly involved in monitoring programs, appointed as citizen scientists with potential to delivering key data at near to no cost to address environmental challenges, therein fostering scientific knowledge and advising policy- and decision-makers. One of the first and most successful examples of marine citizen science in the Mediterranean is represented by the integrative and collaborative implementation of several jellyfish-spotting campaigns in Italy, Spain, Malta, and Tunisia starting in 2009. Altogether, in terms of time coverage, geographic extent, and number of citizen records, these represent the most effective marine citizen science campaigns thus far implemented in the Mediterranean Sea. Here, we analyzed a collective database merging records over the above four countries, featuring more than 100,000 records containing almost 25,000 observations of jellyfish specimens collected over a period of 3 to 7 years (from 2009 to 2015) by citizen scientists participating in any of the national citizen science programs included in this analysis. Such a wide citizen science exercise demonstrates a valuable and cost-effective tool to understanding ecological drivers of jellyfish proliferation over the Western and Central Mediterranean basins, as well as a powerful contribution to developing tailored adaptation and management strategies; mitigating jellyfish impacts on human activities in coastal zones; and supporting implementation of marine spatial planning, Blue Growth, and conservation strategies.

Jellyfish as food represent a millennial tradition in Asia. Recently, jellyfish have also been proposed as a valuable source of protein in Western countries. To identify health risks associated with the potential human consumption of jellyfish as food, trace element accumulation was assessed in the gonads and umbrella tissues of the Mediterranean Rhizostoma pulmo (Macri, 1778), sampled over a period of 16 months along the shallow coastal waters a short distance from the city of Taranto, an area affected by metallurgic and oil refinery sources of pollution. Higher tissue concentrations of trace elements were usually detected in gonads than in umbrella tissue. In particular, significant differences in the toxic metalloid As, and in the metals Mn, Mo, and Zn, were observed among different tissues. The concentrations of vanadium were slightly higher in umbrella tissues than in gonads. No positive correlation was observed between element concentration and jellyfish size, suggesting the lack of bioaccumulation processes. Moreover, toxic element concentrations in R. pulmo were found below the threshold levels for human consumption allowed by Australian, USA, and EU Food Regulations. These results corroborate the hypothesis that R. pulmo is a safe, potentially novel food source, even when jellyfish are harvested from coastal areas affected by anthropogenic impacts.

Jellyfish blooms are frequent and widespread in coastal areas worldwide, often associated with significant ecological and socio-economic consequences. Recent studies have also suggested cnidarian jellyfish may act as vectors of bacterial pathogens. The scyphomedusa Rhizostoma pulmo is an outbreak-forming jellyfish widely occurring across the Mediterranean basin. Using combination of culture-based approaches and a high-throughput amplicon sequencing (HTS), and based on available knowledge on a warm-affinity jellyfish-associated microbiome, we compared the microbial community associated with R. pulmo adult jellyfish in the Gulf of Taranto (Ionian Sea) between summer (July 2016) and winter (February 2017) sampling periods. The jellyfish-associated microbiota was investigated in three distinct compartments, namely umbrella, oral arms, and the mucus secretion. Actinobacteria, Bacteroidetes, Chlamydiae, Cyanobacteria, Deinococcus-Thermus, Firmicutes, Fusobacteria, Planctomycetes, Proteobacteria, Rhodothermaeota, Spirochaetes, Tenericutes, and Thaumarchaeota were the phyla isolated from all the three R. pulmo compartments in the sampling times. In particular, the main genera Mycoplasma and Spiroplasma, belonging to the class Mollicutes (phylum Tenericutes), have been identified in all the three jellyfish compartments. The taxonomic microbial data were coupled with metabolic profiles resulting from the utilization of 31 different carbon sources by the BIOLOG Eco-Plate system. Microorganisms associated with mucus are characterized by great diversity. The counts of culturable heterotrophic bacteria and potential metabolic activities are also remarkable. Results are discussed in terms of R. pulmo ecology, the potential health hazard for marine and human life as well as the potential biotechnological applications related to the associated microbiome.

The Mediterranean basin is one of the regions heavily affected by jellyfish bloom phenomena, mainly due to the presence of scyphozoans, such as Rhizostoma pulmo. The jellyfish have few natural predators, and their bodies represent an organic-rich substrate that can support rapid bacterial growth with great impact on the structure of marine food webs. In Asiatic countries, jellyfish are widely studied for their health benefits, but their nutritional and nutraceutical values still remain poorly characterized. In this study, the differences in the 1H NMR spectroscopy metabolic profiles of R. pulmo female gonads and body fractions (including umbrella and oral arms), in different sampling periods, were studied. For each body compartment both lipid and aqueous extracts were characterized and their 1H NMR metabolic profiles subjected to multivariate analysis. From a statistical analysis of the extracts, a higher contents of ω-3 polyunsaturated fatty acids (PUFAs), amino acid and osmolytes (homarine, betaine, taurine) with important roles in marine invertebrates were observed in female gonads, whereas umbrella and oral arms showed similar metabolic profiles. These results support a sustainable exploitation of the jellyfish for the extraction of bioactive compounds useful in nutraceutical, nutricosmetics, and functional food fields.

Aquaculture is increasing rapidly to meet global seafood demand. Some hydroid populations have been linked to mortality and health issues in finfish and shellfish, but their dynamics in and around aquaculture farms remain understudied. In the present work, two experiments, each with 36 panels, tested colonization (factors: depth, season of immersion) and succession (factors: depth, submersion duration) over one year. Hydroid surface cover was estimated for each species, and data were analyzed with multivariate techniques. The assemblage of hydrozoans was species-poor, although species richness, frequency and abundance increased with time, paralleling the overall increase in structural complexity of fouling assemblages. Submersion duration and season of immersion were particularly important in determining the species composition of the assemblages in the succession and colonization experiments, respectively. Production of water-borne propagules, including medusae, from the hydroids was observed from locally abundant colonies, among them the well-known fouling species Obelia dichotoma, potentially representing a nuisance for cultured fish through contact-driven envenomations and gill disorders. The results illustrate the potential importance of fouling hydroids and their medusae to the health of organisms in the aquaculture industry.

[This corrects the article DOI: 10.1371/journal.pone.0195352.].

The mucus produced by many marine organisms is a complex mixture of proteins and polysaccharides forming a weak watery gel. It is essential for vital processes including locomotion, navigation, structural support, heterotrophic feeding and defence against a multitude of environmental stresses, predators, parasites, and pathogens. In the present study we focused on mucus produced by a benthic cnidarian, the sea anemone Actinia equina (Linnaeus, 1758) for preventing burial by excess sedimentation and for protection. We investigated some of the physico-chemical properties of this matrix such as viscosity, osmolarity, electrical conductivity, protein, carbohydrate, and total lipid contents. Some biological activities such as hemolytic, cytotoxic, and antibacterial lysozyme-like activities were also studied. The A. equina mucus is mainly composed by water (96.2% ± 0.3%), whereas its dry weight is made of 24.2% ± 1.3% proteins and 7.8% ± 0.2% carbohydrates, with the smallest and largest components referable to lipids (0.9%) and inorganic matter (67.1%). The A. equina mucus matrix exhibited hemolytic activity on rabbit erythrocytes, cytotoxic activity against the tumor cell line K562 (human erythromyeloblastoid leukemia) and antibacterial lysozyme-like activity. The findings from this study improve the available information on the mucus composition in invertebrates and have implications for future investigations related to exploitation of A. equina and other sea anemones’ mucus as a source of bioactive compounds of high pharmaceutical and biotechnological interest.

Jellyfish are recorded with increasing frequency and magnitude in many coastal areas and several species display biological features comparable to the most popular Asiatic edible jellyfish. The biochemical and antioxidant properties of wild gelatinous biomasses, in terms of nutritional and nutraceutical values, are still largely unexplored. In this paper, three of the most abundant and commonly recorded jellyfish species (Aurelia sp.1, Cotylorhiza tuberculata and Rhizostoma pulmo) in the Mediterranean Sea were subject to investigation. A sequential enzymatic hydrolysis of jellyfish proteins was set up by pepsin and collagenase treatments of jellyfish samples after aqueous or hydroalcoholic protein extraction. The content and composition of proteins, amino acids, phenolics, and fatty acids of the three species were recorded and compared. Protein content (mainly represented by collagen) up to 40% of jellyfish dry weight were found in two of the three jellyfish species (C. tuberculata and R. pulmo), whereas the presence of ω-3 and ω-6 polyunsaturated fatty acids (PUFAs) was significantly higher in the zooxanthellate jellyfish C. tuberculata only. Remarkable antioxidant ability was also recorded from both proteinaceous and non proteinaceous extracts and the hydrolyzed protein fractions in all the three species. The abundance of collagen, peptides and other bioactive molecules make these Mediterranean gelatinous biomasses a largely untapped source of natural compounds of nutraceutical, cosmeceutical and pharmacological interest.

In recent years, caged finfish mariculture across European seas suffered production losses by severe fish mortality, following episodic outbreaks of invertebrate cnidarian stingers. Due to their stinging cells and injectable venoms, medusozoan jellyfish, or drifting propagules of polyp colonies at high density may impair caged fish health through toxic effects on vulnerable tissues of gills and skin, and related secondary bacterial infections. Gill disorders in European sea bass (Dicentrarchus labrax) fish farms along the Spanish Mediterranean coast are commonly reported, but regular monitoring of the frequency of cnidarian outbreaks and their potential impacts on caged fish is still poorly enforced. In this study, two sea bass mariculture farms in Southern Spain (Málaga; Almería) were monitored biweekly for zooplankton, phytoplankton and fish gills condition, over 13 or 30 months for the Málaga and Almería facilities, respectively, within the period 2012–2014. Significant, direct correlations were found among low water temperature, recorded fish mortalities, and high abundances of planktonic cnidarians, particularly of the hydrozoan siphonophores Muggiaea atlantica and M. kochii, and the larval stage of Ectopleura larynx, a common member of cage biofouling communities. A significant relationship between cnidarian densities and the quantitative scoring of gill pathology was also observed. In addition, high densities of long-bristled planktonic diatoms (Chaetoceros spp.) coincided with a major fish mortality event (April 2012, Almería farm). Standardised monitoring of plankton dynamics and composition may help in promoting response capacities of Mediterranean mariculture managers to fish health challenges (such as stinging plankton blooms) by (a) improving diagnostic tools and preventative countermeasures and (b) supporting the development of science-based spatial planning and sustainable growth of coastal mariculture.

Polyps of two moon jellyfish species, Aurelia coerulea and A. relicta, from two Adriatic Sea coastal habitats were incubated under multiple combinations of temperature (14, 21°C), salinity (24, 37 ppt) and food regime (9.3, 18.6, 27.9 μg C ind-1 week-1) to comparatively assess how these factors may influence major asexual reproduction processes in the two species. Both species exhibited a shared pattern of budding mode (Directly Budded Polyps: DBP; Stolonal Budded Polyps: SBP), with DBP favoured under low food supply (9.3 μg C ind -1 week-1) and low temperature (14°C), and SBP dominant under high temperature (21°C). However, A. coerulea showed an overall higher productivity than A. relicta, in terms of budding and podocyst production rates. Further, A. coerulea exhibited a wide physiological plasticity across different temperatures and salinities as typical adaptation to ecological features of transitional coastal habitats. This may support the hypothesis that the invasion of A. coerulea across coastal habitats worldwide has been driven by shellfish aquaculture, with scyphistoma polyps and resting stages commonly found on bivalve shells. On the contrary, A. relicta appears to be strongly stenovalent, with cold, marine environmental optimal preferences (salinity 37 ppt, T ranging 14-19°C), corroborating the hypothesis of endemicity within the highly peculiar habitat of the Mljet lake. By exposing A. relicta polyps to slightly higher temperature (21°C), a previously unknown developmental mode was observed, by the sessile polyp regressing into a dispersive, temporarily unattached and tentacle-less, non-feeding stage. This may allow A. relicta polyps to escape climatic anomalies associated to warming of surface layers and deepening of isotherms, by moving into deeper, colder layers. Overall, investigations on species-specific eco-physiological and ontogenetic potentials of polyp stages may contribute to clarify the biogeographic distribution of jellyfish and the phylogenetic relationships among evolutionary related sister clades.

Biological fouling organisms on fish cages represent a major issue and costly factor in marine finfish aquaculture. Cnidarians have been identified as one of the most problematical groups, contributing significantly to the occlusion and structural stress of the cage nets, but also dramatically affecting farmed species health in aquaculture facilities worldwide. Recently, significant relationships were established in different Spanish aquaculture facilities between hydrozoans and juvenile fish affected by gill injuries and mortality episodes. Community composition, growth rate and reproductive potential of biofouling were monitored on fish cages over two seasonal periods of fry cages farming, located in southern Spain (SW Alboran Sea), with a special focus on cnidarians. Biomass and community composition of biofouling changed with time and between studied periods, with a marked seasonality in colonization periods and taxonomic composition, particularly for the colonial hydrozoans. The hydroids Ectopleura larynx and Pennaria disticha were found at the highest densities. P. disticha was responsible for major biomass contribution to total hydroid biomass with the fastest growth rates. In addition, actinulae larvae of E. larynx were identified in zooplankton samples at high densities especially during periods of fry introduction in sea cages (when fish are highly vulnerable). These results corroborate evidence of the detrimental influence of fouling cnidarians in Mediterranean finfish aquaculture due to a direct harmful impact on fish health. Investigations on population dynamics, reproductive biology and envenomation potential of fouling hydrozoans should be regarded as key component of best monitoring practices to ensure good farmed fish welfare, maximization of aquaculture production and overall marine spatial planning.

The jellyfish Rhizostoma pulmo, Macrì 1778 (Cnidaria, Rhizostomae) undergoes recurrent outbreaks in the Mediterranean coastal waters, with large biomass populations representing a nuisance or damage for marine and maritime activities. A preliminary overview of the antioxidant activity (AA) of R. pulmo proteinaceous compounds is provided here based on the extraction and characterization of both soluble and insoluble membrane-fractioned proteins, the latter digested by sequential enzymatic hydrolyses with pepsin and collagenases. All jellyfish proteins showed significant AA, with low molecular weight (MW) proteins correlated with greater antioxidant activity. In particular, collagenase-hydrolysed collagen resulted in peptides with MW lower than 3 kDa, ranging 3⁻10 kDa or 10⁻30 kDa, with AA inversely proportional to MW. No cytotoxic effect was detected on cultured human keratinocytes (HEKa) in a range of protein concentration 0.05⁻20 μg/mL for all tested protein fractions except for soluble proteins higher than 30 kDa, likely containing the jellyfish venom compounds. Furthermore, hydrolyzed jellyfish collagen peptides showed a significantly higher AA and provided a greater protective effect against oxidative stress in HEKa than the hydrolyzed collagen peptides from vertebrates. Due to a high reproductive potential, jellyfish may represent a potential socioeconomic opportunity as a source of natural bioactive compounds, with far-reaching beneficial implications. Eventually, improvements in processing technology will promote the use of untapped marine biomasses in nutraceutical, cosmeceutical, and pharmaceutical fields, turning marine management problems into a more positive perspective.

Jellyfish outbreaks in marine coastal areas represent an emergent problem worldwide, with negative consequences on human activities and ecosystem functioning. However, potential positive effects of jellyfish biomass proliferation may be envisaged as a natural source of bioactive compounds of pharmaceutical interest. We investigated the biochemical composition of mature female gonads and lysozyme antibacterial activity of oocytes in the Mediterranean barrel jellyfish Rhizostoma pulmo. Chemical characterization was performed by means of multinuclear and multidimensional NMR spectroscopy. The ovaries of R. pulmo were mainly composed of water (93.7 ± 1.9% of wet weight), with organic matter (OM) and dry weight made respectively of proteins (761.76 ± 25.11 µg mg−1 and 45.7 ± 1.5%), lipids (192.17 ± 10.56 µg mg−1 and 9.6 ± 0.6%), and carbohydrates (59.66 ± 2.72 µg mg−1 and 3.7 ± 0.3%). The aqueous extract of R. pulmo gonads contained free amino acids, organic acids, and derivatives; the lipid extract was composed of triglycerides (TG), polyunsaturated fatty acids (PUFAs), diunsaturated fatty acids (DUFAs), monounsaturated fatty acids (MUFAs), saturated fatty acids (SFAs), and minor components such as sterols and phospholipids. The R. pulmo oocyte lysate exhibited an antibacterial lysozyme-like activity (mean diameter of lysis of 9.33 ± 0.32 mm corresponding to 1.21 mg/mL of hen egg-white lysozyme). The occurrence of defense molecules is a crucial mechanism to grant healthy development of mature eggs and fertilized embryos (and the reproductive success of the species) by preventing marine bacterial overgrowth. As a corollary, these results call for future investigations for an exploitation of R. pulmo biomasses as a resource of bioactive metabolites of biotechnological importance including pharmaceuticals and nutrition.

On a global scale, jellyfish populations in coastal marine ecosystems exhibit increasing trends of abundance. High-density outbreaks may directly or indirectly affect human economical and recreational activities, as well as public health. As the interest in biology of marine jellyfish grows, a number of jellyfish metabolites with healthy potential, such as anticancer or antioxidant activities, is increasingly reported. In this study, the Mediterranean “fried egg jellyfish” Cotylorhiza tuberculata (Macri, 1778) has been targeted in the search forputative valuable bioactive compounds. A medusa extract was obtained, fractionated, characterized by HPLC, GC-MS and SDS-PAGE and assayed for its biological activity on breast cancer cells (MCF-7) and human epidermal keratinocytes (HEKa). The composition of the jellyfish extract included photosynthetic pigments, valuable ω-3 and ω-6 fatty acids, and polypeptides derived either from jellyfish tissues and their algal symbionts. Extract fractions showed antioxidant activity and the ability to affect cell viability and intercellular communication mediated by gap junctions (GJIC) differentially in MCF-7and HEKa cells. A significantly higher cytotoxicity and GJIC enhancement in MCF-7 compared to HEKa cells was recorded. A putative action mechanism for the anticancer bioactivity through the modulation of GJIC has been hypothesized and its nutraceutical and pharmaceutical potential was discussed.

Jellyfish are of particular concern for marine finfish aquaculture. In recent years repeated mass mortality episodes of farmed fish were caused by blooms of gelatinous cnidarian stingers, as a consequence of a wide range of hemolytic, cytotoxic, and neurotoxic properties of associated cnidocytes venoms. The mauve stinger jellyfish Pelagia noctiluca (Scyphozoa) has been identified as direct causative agent for several documented fish mortality events both in Northern Europe and the Mediterranean Sea aquaculture farms. We investigated the effects of P. noctiluca envenomations on the gilthead sea bream Sparus aurata by in vivo laboratory assays. Fish were incubated for 8 hours with jellyfish at 3 different densities in 300 l experimental tanks. Gill disorders were assessed by histological analyses and histopathological scoring of samples collected at time intervals from 3 hours to 4 weeks after initial exposure. Fish gills showed different extent and severity of gill lesions according to jellyfish density and incubation time, and long after the removal of jellyfish from tanks. Jellyfish envenomation elicits local and systemic inflammation reactions, histopathology and gill cell toxicity, with severe impacts on fish health. Altogether, these results shows P. noctiluca swarms may represent a high risk for Mediterranean finfish aquaculture farms, generating significant gill damage after only a few hours of contact with farmed S. aurata. Due to the growth of the aquaculture sector and the increased frequency of jellyfish blooms in the coastal waters, negative interactions between stinging jellyfish and farmed fish are likely to increase with the potential for significant economic losses.

Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the future.

In recent years, jellyfish blooms have attracted considerable scientific interest for their potential impacts on human activities and ecosystem functioning, with much attention paid to jellyfish as predators and to gelatinous biomass as a carbon sink. Other than qualitative data and observations, few studies have quantified direct predation of fish on jellyfish to clarify whether they may represent a seasonally abundant food source. Here we estimate predation frequency by the commercially valuable Mediterranean bogue, Boops boops on the mauve stinger jellyfish, Pelagia noctiluca, in the Strait of Messina (NE Sicily). A total of 1054 jellyfish were sampled throughout one year to quantify predation by B. boops from bite marks on partially eaten jellyfish and energy density of the jellyfish. Predation by B. boops in summer was almost twice that in winter, and they selectively fed according to medusa gender and body part. Calorimetric analysis and biochemical composition showed that female jellyfish gonads had significantly higher energy content than male gonads due to more lipids and that gonads had six-fold higher energy content than the somatic tissues due to higher lipid and protein concentrations. Energetically, jellyfish gonads represent a highly rewarding food source, largely available to B. boops throughout spring and summer. During the remainder of the year, when gonads were not very evident, fish predation switched towards less-selective foraging on the somatic gelatinous biomass. P. noctiluca, the most abundant jellyfish species in the Mediterranean Sea and a key planktonic predator, may represent not only a nuisance for human leisure activities and a source of mortality for fish eggs and larvae, but also an important resource for fish species of commercial value, such as B. boops.

Genetic drift and non-random mating seldom influence species with large breeding populations and high dispersal potential, characterized by unstructured gene pool and panmixia at a scale lower than the minimum dispersal range of individuals. In the present study, a set of nine microsatellite markers was developed and used to investigate the spatio-temporal genetic patterns of the holoplanktonic jellyfish Pelagia noctiluca (Scyphozoa) in the Southern Tyrrhenian Sea. Homozygote excess was detected at eight loci, and individuals exhibited intra-population relatedness higher than expected by chance in at least three samples. This result was supported by the presence of siblings in at least 5 out 8 samples, 4 of which contained full-sib in addition to half-sib dyads. Having tested and ruled out alternative explanations as null alleles, our results suggest the influence of reproductive and behavioural features in shaping the genetic structure of P. noctiluca, as outcomes of population genetics analyses pointed out. Indeed, the genetic differentiation among populations was globally small but highlighted: a) a spatial genetic patchiness uncorrelated with distance between sampling locations, and b) a significant genetic heterogeneity between samples collected in the same locations in different years. Therefore, despite its extreme dispersal potential, P. noctiluca does not maintain a single homogenous population, but rather these jellyfish appear to have intra-bloom localized recruitment and/or individual cohesiveness, whereby siblings more likely swarm together as a single group and remain close after spawning events. These findings provide the first evidence of family structures and consequent genetic patchiness in a species with highly dispersive potential throughout its whole life cycle, contributing to understanding the patterns of dispersal and connectivity in marine environments.

Between March-May 2013 a massive Salpa maxima bloom was recorded by a citizen science study along the Ionian and Adriatic coast of the Salento peninsula (Italy). Citizen records were substantiated with field inspections along the coast and during an oceanographic campaign in the Otranto Channel. Salps clogged nets, impairing fishing activities along the coast. Swimmers were scared by the gelatinous appearance of the salps, and thought they were jellyfish. At the end of the bloom the dead bodies of the colonies, that were up to 6-7 m long, were accumulated along the coast and stirred by the waves, forming foams along dozens of kilometers of coast. The bloom also occurred at the Tremiti Islands, north of the Gargano Peninsula. The possible impacts of such events on the functioning of pelagic systems are discussed.

Gelatinous zooplankton outbreaks have increased globally owing to a number of human-mediated factors, including food web alterations and species introductions. The invasive ctenophore Mnemiopsis leidyi entered the Black Sea in the early 1980s. The invasion was followed by the Azov, Caspian, Baltic and North Seas, and, most recently, the Mediterranean Sea. Previous studies identified two distinct invasion pathways of M. leidyi from its native range in the western Atlantic Ocean to Eurasia. However, the source of newly established populations in the Mediterranean Sea remains unclear. Here we build upon our previous study and investigate sequence variation in both mitochondrial (Cytochrome c Oxidase subunit I) and nuclear (Internal Transcribed Spacer) markers in M. leidyi, encompassing five native and 11 introduced populations, including four from the Mediterranean Sea. Extant genetic diversity in Mediterranean populations (n = 8, N a = 10) preclude the occurrence of a severe genetic bottleneck or founder effects in the initial colonizing population. Our mitochondrial and nuclear marker surveys revealed two possible pathways of introduction into Mediterranean Sea. In total, 17 haplotypes and 18 alleles were recovered from all surveyed populations. Haplotype and allelic diversity of Mediterranean populations were comparable to populations from which they were likely drawn. The distribution of genetic diversity and pattern of genetic differentiation suggest initial colonization of the Mediterranean from the Black-Azov Seas (pairwise F ST = 0.001-0.028). However, some haplotypes and alleles from the Mediterranean Sea were not detected from the well-sampled Black Sea, although they were found in Gulf of Mexico populations that were also genetically similar to those in the Mediterranean Sea (pairwise F ST = 0.010-0.032), raising the possibility of multiple invasion sources. Multiple introductions from a combination of Black Sea and native region sources could be facilitated by intense local and transcontinental shipping activity, respectively.