Deficiencies in monitoring practices of marine protected areas in southern European seas.

Worldwide, states are gazetting new Marine Protected Areas (MPAs) to meet the international commitment of protecting 30% of the seas by 2030. Yet, protection benefits only come into effect when an MPA is implemented with activated regulations and actively managed through continuous monitoring and adaptive management. To assess if actively managed MPAs are the rule or the exception, we used the Mediterranean and Black Seas as a case study, and retrieved information on monitoring activities for 878 designated MPAs in ten European Union (EU) countries. We searched for scientific and grey literature that provides information on the following aspects of MPA assessment and monitoring: ecological (e.g., biomass of commercially exploited fish), social (e.g., perceptions of fishers in an MPA), economic (e.g., revenue of fishers) and governance (e.g., type of governance scheme). We also queried MPA authorities on their past and current monitoring activities using a web-based survey through which we collected 123 responses. Combining the literature review and survey results, we found that approximately 16% of the MPA designations (N = 878) have baseline and/or monitoring studies. Most monitoring programs evaluated MPAs based solely on biological/ecological variables and fewer included social, economic and/or governance variables, failing to capture and assess the social-ecological dimension of marine conservation. To increase the capacity of MPAs to design and implement effective social-ecological monitoring programs, we recommend strategies revolving around three pillars: funding, collaboration, and technology. Following the actionable recommendations presented herein, MPA authorities and EU Member States could improve the low level of MPA monitoring to more effectively reach the 30% protection target delivering benefits for biodiversity conservation.

Biochemical Characterization of Cassiopea andromeda (Forsskål, 1775), Another Red Sea Jellyfish in the Western Mediterranean Sea.

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.

Co-inertia multivariate approach for the evaluation of anthropogenic impact on two commercial fish along Tyrrhenian coasts.

Aliphatic hydrocarbon levels were determined by the GC/MS technique in fish livers of Engraulis encrasicolus (Ee) and Trachurus trachurus (Tt), collected from a particular area of the Mediterranean Sea, called GSA 10, which is located exactly in Tyrrhenian Sea between Campania coast and North Sicily coast. The aim was to evaluate their potential use as specific bioindicators towards this class of contaminants. Both Tt and Ee are considered to be pollution monitoring bioindicators, due to their dominance in marine communities and economic fishing interest. Ee showed a higher tendency to bioaccumulate TAHs, due to the lower quantity of fatty acids in liver tissues with respect to Tt. The area under study has been characterised a) chemically with the acquisition of temperature, oxygen and salinity profiles along the water column, and b) ecologically with the determination of amino acid contents in fish eyes, in order to gain information on the adaptation to environmental changes. Moreover, specific activities of two hydrolytic enzymes, such as alkaline phosphatase and peroxidase in fish epidermal mucus, together with lactate in blood plasma and cortisol levels, have been investigated for the first time, in order to obtain insights into the effects of hydrocarbons on animal welfare. A multiple co-inertia analysis was also applied to chemical and environmental parameters, in order to explore any possible correlation between different variables. The multivariate approach showed a clear spatial distribution between environmental and chemical variables in Ee, whilst there was an absence of a spatial trend in Tt. Moreover, the chemometric analysis showed a very high correlation between amino acid profiles and environmental variables for both species, confirming the possibility of being used as ecological welfare indices for short-term environmental variations.

Pelagia benovici sp. nov. (Cnidaria, Scyphozoa): a new jellyfish in the Mediterranean Sea.

A bloom of an unknown semaestome jellyfish species was recorded in the North Adriatic Sea from September 2013 to early 2014. Morphological analysis of several specimens showed distinct differences from other known semaestome species in the Mediterranean Sea and unquestionably identified them as belonging to a new pelagiid species within genus Pelagia. The new species is morphologically distinct from P. noctiluca, currently the only recognized valid species in the genus, and from other doubtful Pelagia species recorded from other areas of the world. Molecular analyses of mitochondrial cytochrome c oxidase subunit I (COI) and nuclear 28S ribosomal DNA genes corroborate its specific distinction from P. noctiluca and other pelagiid taxa, supporting the monophyly of Pelagiidae. Thus, we describe Pelagia benovici sp. nov. Piraino, Aglieri, Scorrano & Boero.

Mediterranean fish communities are struggling to adapt to global warming. Evidence from the western coast of Italy.

Climate change has significant impacts on marine ecosystems, resulting in disruptions in biological interactions, shifts in community composition, and changes in the physiology of fish and other marine organisms. In this study conducted in the central Mediterranean Sea, the mean temperature of the catch (MTC) was employed as an indicator to investigate the climatological factors influencing the fish community. The MTC, which utilizes species-preferred temperatures, was calculated using bottom temperature (BT) data weighted against scientific catches. The estimated MTC increasing rates were 0.01 °C year-1 for the entire community, 0.017 °C year-1 for the shelf break, and 0.004 °C year-1 for the continental slope assemblage. We found that MTC is increasing at a lower rate compared to BT, suggesting a progressive under-adaptation of the fish community that seems not fully able to keep up with the ongoing pace of warming. The study identified sea surface temperature and bottom temperature as key drivers of changes in fish community composition. Notably, the fish community composition exhibited drastic changes over the studied period, and we suggest that the MTC can be a useful index to monitor such changes within the context of the EU's climate change adaptation strategy.

Reproductive cycle and gonadal output of the Lessepsian jellyfish Cassiopea andromeda in NW Sicily (Central Mediterranean Sea).

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.

Short-term exposure to concurrent biotic and abiotic stressors may impair farmed molluscs performance.

Global warming, through increasing temperatures, may facilitate the spread and proliferation of outbreak-forming species which may find favourable substrate conditions on artificial aquaculture structures. The presence of stinging organisms (cnidarian hydroids) in the facilities fouling community are a source of pollution that can cause critical problems when in-situ underwater cleaning processes are performed. Multiple stressor experiments were carried out to investigate the cumulative effect on farmed mussels' functional traits when exposed to realistic stressful conditions, including presence of harmful cnidarian cells and environmental conditions of increasing temperature and short-term hypoxia. Exposure to combined stressors significantly altered mussels' performance, causing metabolic depression and low filtering activity, potentially delaying, or inhibiting their recovery ability and ultimately jeopardizing organisms' fitness. Further research on the stressors properties and occurrence is needed to obtain more realistic responses from organisms to minimize climate change impacts and increase ecosystem and marine economic activities resilience to multiple stressors.

Snapshot of the Distribution and Biology of Alien Jellyfish Cassiopea andromeda (Forsskål, 1775) in a Mediterranean Touristic Harbour.

Harbors are hotspots for the introduction of alien species, and, usually, investigations on their host populations help fill the knowledge gap in their pathways of invasion and in their impacts on marine biodiversity and ecosystems. In 2014, the upside-down alien jellyfish Cassiopea andromeda invaded a Mediterranean touristic harbor ("Cala"), and its abundance has since increased over time. In the present study, the distribution and trophic behavior of C. andromeda in Cala were investigated for the years 2017-2018 through visual sampling, and GIS-based statistical and stable isotope analyses. Since Cala is a hard-to-reach area (with many anchor cables and boats), Megabenthos Underwater Video was used to count the number and estimate the size of jellyfishes. The variations in size throughout the study period suggest that the population of C. andromeda is quite established in Cala at depths lower than 7.5 m. The ranges of the environmental parameters recorded (temperature, salinity, and transparency) were consistent with the ideal conditions for maintaining a Cassiopea population, but they did not seem to influence aggregation. Additionally, the carbon and nitrogen isotopic signatures studied highlight the mixotrophic behavior of this species. These preliminary results confirm the capacity of C. andromeda to live and reproduce in heavily anthropized areas.

Small-scale fisheries catch more threatened elasmobranchs inside partially protected areas than in unprotected areas.

Elasmobranchs are heavily impacted by fishing. Catch statistics are grossly underestimated due to missing data from various fishery sectors such as small-scale fisheries. Marine Protected Areas are proposed as a tool to protect elasmobranchs and counter their ongoing depletion. We assess elasmobranchs caught in 1,256 fishing operations with fixed nets carried out in partially protected areas within Marine Protected Areas and unprotected areas beyond Marine Protected Areas borders at 11 locations in 6 Mediterranean countries. Twenty-four elasmobranch species were recorded, more than one-third belonging to the IUCN threatened categories (Vulnerable, Endangered, or Critically Endangered). Catches per unit of effort of threatened and data deficient species were higher (with more immature individuals being caught) in partially protected areas than in unprotected areas. Our study suggests that despite partially protected areas having the potential to deliver ecological benefits for threatened elasmobranchs, poor small-scale fisheries management inside Marine Protected Areas could hinder them from achieving this important conservation objective.

Importance of the Lunar Cycle on Mesopelagic Foraging by Atlantic Bluefin Tuna in the Upwelling Area of the Strait of Messina (Central Mediterranean Sea).

The influence of the lunar cycle on bluefin tuna foraging in the upwelling area of the Strait of Messina was investigated by exploring trophic interaction with mesopelagic fish and cephalopod prey. To focus on how the lunar cycle could affect availability of mesopelagic prey for this predator, we tested potential differences in the diet related to each lunar phase. Moreover, we considered two potential impacts of the lunar cycle: the lunar irradiance and the strength of currents. Overall, 2672 prey items were mesopelagic fish and cephalopods, representing 60.7% of overall diet by number. The main mesopelagic fish prey items were lanternfishes and dragonfishes, while Onychoteuthis banksii was the most important cephalopod prey. In summary, the Strait of Messina has highly specific hydrodynamic and biological features which strongly depend on upwelling currents, which in turn are influenced by the lunar cycle (new and full moon with strong currents, quarters with fewer currents). Upwelling causes water mixing, bringing to the surface a large amount of mesopelagic fauna which become more readily available to tuna. Lunar irradiance contributes to the variation of prey composition, increasing the success of visual predation on mesopelagic resources at high light in the water column.

The buffer effect of canopy-forming algae on vermetid reefs' functioning: A multiple stressor case study.

Biodiversity plays a key role for our planet by buffering ongoing and future changes in environmental conditions. We tested if canopy-forming algae enhancing biodiversity (CEB) in a Mediterranean intertidal reef ecological community could alleviate the effect of stressors (heat waves and pollution from sewage) on community metabolic rates (as expressed by oxygen consumption) used as a proxy of community functioning. CEB exerted a buffering effect related to the properties of stressor: physical-pulsing (heat wave) and chronic-trophic (sewage). After a simulated heat wave, CEB was effective in buffering the impacts of detrimental temperatures on the functioning of the community. In reefs exposed to chronic sewage effluents, benefits derived from CEB were less evident, which is likely due to the stressor's contextual action. The results support the hypothesis that ecological responses depend on stressor typology acting at local level and provide insights for improving management measures to mitigate anthropogenic disturbance.

Integrating functional traits into correlative species distribution models to investigate the vulnerability of marine human activities to climate change.

Climate change and particularly warming are significantly impacting marine ecosystems and the services they provided. Temperature, as the main factor driving all biological processes, may influence ectotherms metabolism, thermal tolerance limits and distribution species patterns. The joining action of climate change and local stressors (including the increasing human marine use) may facilitate the spread of non-indigenous and native outbreak forming species, leading to associated economic consequences for marine coastal economies. Marine aquaculture is one among the most economic anthropogenic activities threatened by multiple stressors and in turn, by increasing hard artificial substrates at sea would facilitate the expansion of these problematic organisms and face negative consequences regarding facilities management and farmed organisms' welfare. Species Distribution Models (SDMs) are considered powerful tools for forecasting the future occurrences and distributions of problematic species used to preventively aware stakeholders. In the current study, we propose the use of combined correlative SDMs and mechanistic models, based on individual thermal performance curve models calculated through non-linear least squares regression and Bayesian statistics (functional-SDM), as an ecological relevant tool to increase our ability to investigate the potential indirect effect of climate change on the distributions of harmful species for human activities at sea, taking aquaculture as a food productive example and the benthic cnidarian Pennaria disticha (one of the most pernicious fouling species in aquaculture) as model species. Our combined approach was able to improve the prediction ability of both mechanistic and correlative models to get more ecologically informed "whole" niche of the studied species. Incorporating the mechanistic links between the organisms' functional traits and their environments into SDMs through the use of a Bayesian functional-SDM approach would be a useful and reliable tool in early warning ecological systems, risk assessment and management actions focused on important economic activities and natural ecosystems conservation.

Seasonal variability of diet and trophic level of the gelatinous predator Pelagia noctiluca (Scyphozoa).

Jellyfish populations apparently have increased in some places around the world and human problems with them also have increased. However, effects of jellyfish outbreaks in the ecosystems remain poorly understood and little or no information is available on their dietary preferences - in relation to the seasonal shifts of prey abundance - and on the potential variability of their impact on marine food webs. The mauve stinger Pelagia noctiluca (Forsskål, 1775) is by far the most common outbreak-forming scyphozoan jellyfish in the Western Mediterranean. By use of a combination of stomach contents, stable isotope (SI) and fatty acid (FA) analyses, we tested the hypothesis that changes in the seasonal dietary sources of P. noctiluca parallel changes in the FA and SI composition. Stomach content and biomarker analyses suggested that P. noctiluca is not a selective predator, cyclically shifting between carnivory and omnivory depending on the seasonality of accessible prey. The combination of SI and FA analyses highlighted the importance of microzooplankton as prey. Specific FA biomarkers showed that the diet of P. noctiluca changed seasonally depending on the availability of living plankton or suspended detritus. This study also revealed significant biochemical differences between jellyfish somatic and gonadal tissues, with total fatty acid concentration in the gonads up to ten times higher than in the somatic tissues.

First evidence of inbreeding, relatedness and chaotic genetic patchiness in the holoplanktonic jellyfish Pelagia noctiluca (Scyphozoa, Cnidaria).

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.

Jellyfish as prey: frequency of predation and selective foraging of Boops boops (Vertebrata, Actinopterygii) on the mauve stinger Pelagia noctiluca (Cnidaria, Scyphozoa).

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.