Exploring the mesoscale connectivity of phytoplankton periodic assemblages' succession in northern Adriatic pelagic habitats.

An appropriate model for phytoplankton distribution patterns is critical for understanding biogeochemical cycles and trophic interactions in the oceans and seas. Because phytoplankton dynamics in coastal waters are more complex due to shallow depth and proximity to land, more accurate models applied to the correct spatial and temporal scales are needed. Our study investigates the role of the atmosphere and hydrosphere in pelagic habitat by modelling phytoplankton assemblages at two Long Term Ecological Research sites in the northern Adriatic Sea using niche-forming environmental variables (wind, temperature, salinity, river discharge, rain, and water column stratification). To study the synchronization between the phytoplankton community and these environmental variables at the two LTER sites, we applied current linear and nonlinear numerical methods for ecological modelling. The aim was to use periodic and/or non-periodic properties of the environmental variables to classify the phytoplankton assemblages at one LTER site (Gulf of Trieste - Slovenia) and then predict them at another LTER site 100 km away (Gulf of Venice - Italy). We found that periodicity played a role in the explanatory and predictive power of the environmental variables and that it was more important than non-periodic events in defining the common structure of the two pelagic habitats. The non-linear classification functions of the neural networks further increased the predictive power of these variables. We observed partial synchronization of communities at the mesoscale and differences between the original and predicted assemblages under similar environmental conditions. We conclude that mesoscale connectivity plays an important role in phytoplankton communities in the northern Adriatic. However, the loss of periodicity of niche-forming variables due to more frequent extreme meteorological and hydrological events could loosen these connections and affect the temporal succession of phytoplankton assemblages.

First basin scale spatial-temporal characterization of underwater sound in the Mediterranean Sea.

Anthropogenic underwater noise is an emergent pollutant. Despite several worldwide monitoring programs, only few data are available for the Mediterranean Sea, one of the global biodiversity hotspots. The results of the first continuous acoustic programme run at a transnational basin scale in the Mediterranean Sea are here presented. Recordings were done from March 2020 to June 2021, including the COVID-19 lockdown, at nine stations in the Northern Adriatic Sea. Spatial-temporal variations of the underwater sound are described, having one third octave band sound pressure levels (SPLs) from 10 Hz to 20 kHz as metrics. Higher and more variable SPLs, mainly related to vessel traffic, were found close to harbours, whereas Natura 2000 stations experienced lower SPLs. Lower values were recorded during the lockdown in five stations. Median yearly SPLs ranged between 64 and 95 as well as 70 and 100 dB re 1 µPa for 63 and 125 Hz bands, respectively. These values are comparable with those previously found in busy shallow EU basins but higher levels are expected during a business-as-usual period. This is a baseline assessment for a highly impacted and environmental valuable area, that needs to be managed in a new sustainable blue growth strategy.

First assessment of underwater sound levels in the Northern Adriatic Sea at the basin scale.

The protection of marine habitats from human-generated underwater noise is an emerging challenge. Baseline information on sound levels, however, is poorly available, especially in the Mediterranean Sea. To bridge this knowledge gap, the SOUNDSCAPE project ran a basin-scale, cross-national, long-term underwater monitoring in the Northern Adriatic Sea. A network of nine monitoring stations, characterized by different natural conditions and anthropogenic pressures, ensured acoustic data collection from March 2020 to June 2021, including the full lockdown period related to the COVID-19 pandemic. Calibrated stationary recorders featured with an omnidirectional Neptune Sonar D60 Hydrophone recorded continuously 24 h a day (48 kHz sampling rate, 16 bit resolution). Data were analysed to Sound Pressure Levels (SPLs) with a specially developed and validated processing app. Here, we release the dataset composed of 20 and 60 seconds averaged SPLs (one-third octave, base 10) output files and a Python script to postprocess them. This dataset represents a benchmark for scientists and policymakers addressing the risk of noise impacts on marine fauna in the Mediterranean Sea and worldwide.

An exceptionally high wave at the CNR-ISMAR oceanographic tower in the Northern Adriatic Sea.

On December 15, 2009, a very high wave crest was recorded by a local camera at the CNR-ISMAR oceanographic tower, 15 km offshore Venice in the Northern Adriatic Sea (Italy). The height of the estimated crest elevation appears well beyond the value (1,25·Hs) commonly used to identify a wave as freak. We document the wave event with a full description of the corresponding met-ocean conditions and related measurements, of which we provide a critical analysis.

The effects of COVID-19 induced lockdown measures on maritime settings of a coastal region.

The spread of coronavirus (COVID-19) caused an unprecedented implementation of lockdown measures across world's nations. Veneto Region, located in North-Eastern Adriatic Sea was one of the first maritime regions in Italy and Europe subjected to progressive lockdown restrictions. We systematically analyse the effects of national lockdown policies on maritime settings of the region using Automated Identification System (AIS) data from fishing vessels, passenger ships, tanker and cargo vessels collected through the Aqua Alta Oceanographic Tower (AAOT). We derive consequences on vessel activities during the March-April 2020 lockdown, by using a data-driven, comparative spatio-temporal analysis of vessel trajectories. Results show that compared to the same period of 2017, vessel activity were reduced by 69% during the lockdown, fishing activities reduced by 84% and passenger traffic by 78%. We register a restart of fishing activity in the third week of April 2020. We suggest that the presented conceptual and spatial assessment protocol can guide future research on environmental and socio-economic effects of COVID-19 on marine realms and contribute to further interdisciplinary research with other marine scientific fields.

Dinoflagellate resting cysts from surface sediments of the Adriatic Ports: Distribution and potential spreading patterns.

The ability of microalgae to preserve viable in coastal sediments as resting forms provides a reservoir of biodiversity and a useful tool to determine species spreadings. This study represents the first port baseline survey on dinoflagellate cysts, investigated in nine Adriatic ports during a cross border project. 40 dinoflagellate taxa were detected. The assemblages resulted in all ports dominated by Lingulodinium polyedra and Alexandrium minutum/affine/tamutum group. General separation to the western and eastern side of the Adriatic regarding cysts assemblage composition, partially abundance, was observed. Six taxa were detected as non-indigenous species for the Adriatic. Two taxa are included in the list of harmful aquatic organisms, indicating the potential threat of ballast waters in the Adriatic. Potential spreading of taxa by general circulation and ballast waters, intra- and extra-Adriatic was investigated. The entering in to force of the ballast waters management regulations should enhance prospects to minimize future harmful impacts.

Phytoplankton diversity in Adriatic ports: Lessons from the port baseline survey for the management of harmful algal species.

An inventory of phytoplankton diversity in 12 Adriatic ports was performed with the port baseline survey. Particular emphasis was put on the detection of harmful aquatic organisms and pathogens (HAOP) because of their negative impact on ecosystem, human health, and the economy. Phytoplanktonic HAOP are identified as species, either native or non-indigenous (NIS), which can trigger harmful algal blooms (HAB). A list of 691 taxa was prepared, and among them 52 were classified as HAB and five as NIS. Records of toxigenic NIS (Pseudo-nitzschia multistriata, Ostreopsis species including O. cf. ovata) indicate that the intrusion of non-native invasive phytoplankton species has already occurred in some Adriatic ports. The seasonal occurrence and abundance of HAOP offers a solid baseline for a monitoring design in ports in order to prevent ballast water uptake and possible expansion of HAOP outside their native region.

Strategy of port baseline surveys (PBS) in the Adriatic Sea.

Port baseline surveys (PBS) provide species inventories in and around ports, with a focus on non-indigenous species that may have been introduced by vessels, primarily via ballast water. PBS are an essential tool to support effective management strategies for non-indigenous as well as native harmful aquatic organisms and pathogens (HAOP). This paper describes the methodology of PBS that were conducted in 12 Adriatic ports. The PBS employed existing protocols that were adapted to meet the characteristics of the Adriatic sites. Their results are reported in several papers included in this special issue, each of which is devoted to a specific community. An overview of existing surveys protocols - which provide valuable support to decision-making and to design effective monitoring of non-indigenous species - is also supplied.

Habitat Heterogeneity and Connectivity: Effects on the Planktonic Protist Community Structure at Two Adjacent Coastal Sites (the Lagoon and the Gulf of Venice, Northern Adriatic Sea, Italy) Revealed by Metabarcoding.

The Lagoon of Venice (LoV) and the Gulf of Venice (GoV), two adjacent coastal Long Term Ecological Research (LTER) sites in the northern Adriatic Sea, represent a transitional/marine coupled ecosystem under the influence of regional and local factors. In this study, these sites were sampled on four dates from April 2016 to February 2017 for environmental DNA and relevant abiotic variables, aiming to assess the relative importance of habitat heterogeneity and connectivity in structuring the protist community. High Throughput Sequencing of V4-18S rRNA gene from 56 samples collected at seven stations produced ca 6 million reads, grouped into 7,336 Operational Taxonomic Units (OTUs) at 97% similarity, which were affiliated to protists belonging to 34 taxonomic groups. The whole community was dominated by Bacillariophyta, especially in spring-summer in the LoV, and by Dinophyta, mainly in the GoV. Ciliophora, Syndiniales, and Cryptophyceae were the next more abundant groups. The community structure varied across the seasons and was different in the two ecosystems, which shared 96% of the reads but showed a high proportion of OTUs distributed preferentially in one of the two sites (specialists) and a different partitioning of trophic categories. GoV specialists were mainly Dinophyceae (>56%), followed by Syndiniales and Bacillariophyta, while the LoV specialists were distributed among several groups, including Bacillariophyta, Syndiniales, Ciliophora, Cryptophyceae, and Trebouxiophyceae. The main abiotic drivers of the differences between protist communities were salinity and temperature, which however explained a minor part of the variance (17%), pointing at a higher relevance of biotic factors and inter-taxa relationships. This was more evident in the LoV, where the network analysis highlighted a higher number of OTUs' connections than in the GoV. Overall, the metabarcoding approach allowed to depict the composition of the whole protist community in the lagoon and adjacent coastal waters with high resolution, revealing many taxa so far not reported in the area. In addition, despite no clear barrier to dispersal processes, differences in the relative abundance and temporal variability of local protist communities indicate that environmental heterogeneity, in these adjacent and connected ecosystems, can be strong enough to allow for ecological segregation.

Plankton dynamics across the freshwater, transitional and marine research sites of the LTER-Italy Network. Patterns, fluctuations, drivers.

A first synoptic and trans-domain overview of plankton dynamics was conducted across the aquatic sites belonging to the Italian Long-Term Ecological Research Network (LTER-Italy). Based on published studies, checked and complemented with unpublished information, we investigated phytoplankton and zooplankton annual dynamics and long-term changes across domains: from the large subalpine lakes to mountain lakes and artificial lakes, from lagoons to marine coastal ecosystems. This study permitted identifying common and unique environmental drivers and ecological functional processes controlling seasonal and long-term temporal course. The most relevant patterns of plankton seasonal succession were revealed, showing that the driving factors were nutrient availability, stratification regime, and freshwater inflow. Phytoplankton and mesozooplankton displayed a wide interannual variability at most sites. Unidirectional or linear long-term trends were rarely detected but all sites were impacted across the years by at least one, but in many case several major stressor(s): nutrient inputs, meteo-climatic variability at the local and regional scale, and direct human activities at specific sites. Different climatic and anthropic forcings frequently co-occurred, whereby the responses of plankton communities were the result of this environmental complexity. Overall, the LTER investigations are providing an unparalleled framework of knowledge to evaluate changes in the aquatic pelagic systems and management options.

Seasonal rather than spatial variability drives planktonic and benthic bacterial diversity in a microtidal lagoon and the adjacent open sea.

Coastal lagoons are highly productive ecosystems, which are experiencing a variety of human disturbances at increasing frequency. Bacteria are key ecological players within lagoons, yet little is known about the magnitude, patterns and drivers of diversity in these transitional environments. We carried out a seasonal study in the Venice Lagoon (Italy) and the adjacent sea, to simultaneously explore diversity patterns in different domains (pelagic, benthic) and their spatio-temporal variability, and test the role of environmental gradients in structuring assemblages. Community composition differed between lagoon and open sea, and between domains. The dominant phyla varied temporally, with varying trends for the two domains, suggesting different environmental constraints on the assemblages. The percentage of freshwater taxa within the lagoon increased during higher river run-off, pointing at the lagoon as a dynamic mosaic of microbial taxa that generate the metacommunity across the whole hydrological continuum. Seasonality was more important than spatial variability in shaping assemblages. Network analyses indicated more interactions between several genera and environmental variables in the open sea than the lagoon. Our study provides evidences for a temporally dynamic nature of bacterial assemblages in lagoons and suggests that an interplay of seasonally influenced environmental drivers shape assemblages in these vulnerable ecosystems.

New oxylipins produced at the end of a diatom bloom and their effects on copepod reproductive success and gene expression levels.

Diatoms are dominant photosynthetic organisms in the world's oceans and are considered essential in the transfer of energy to higher trophic levels. However, these unicellular organisms produce secondary metabolites deriving from the oxidation of fatty acids, collectively termed oxylipins, with negative effects on predators, such as copepods, that feed on them (e.g. reduction in survival, egg production and hatching success) and, indirectly, on higher trophic levels. Here, a multidisciplinary study (oxylipin measurements, copepod fitness, gene expression analyses, chlorophyll distribution, phytoplankton composition, physico-chemical characteristics) was carried out at the end of the spring diatom bloom in April 2011 in the Northern Adriatic Sea (Mediterranean Sea) in order to deeply investigate copepod-diatom interactions, chemical communication and response pathways. The results show that the transect with the lowest phytoplankton abundance had the lowest copepod egg production and hatching success, but the highest oxylipin concentrations. In addition, copepods in both the analyzed transects showed increased expression levels of key stress-related genes (e.g. heat-shock proteins, catalase, glutathione S-transferase, aldehyde dehydrogenase) compared to control laboratory conditions where copepods were fed with the dinoflagellate Prorocentrum minimum which does not produce any oxylipins. New oxylipins that have never been reported before for microalgae are described for the first time, giving new insights into the complex nature of plant-animal signaling and communication pathways at sea. This is also the first study providing insights on the copepod response during a diatom bloom at the molecular level.

Phytoplankton cell lysis associated with polyunsaturated aldehyde release in the Northern Adriatic Sea.

Diatoms are able to react to biotic and abiotic stress, such as competition, predation and unfavorable growth conditions, by producing bioactive compounds including polyunsaturated aldehydes (PUAs). PUAs have been shown to act against grazers and either enhance or inhibit the growth of different phytoplankton and bacteria both in culture and in the field. Presence of nanomolar concentrations of dissolved PUAs in seawater has been reported in the North Adriatic Sea (Mediterranean), suggesting that these compounds are released in seawater following diatom cell lysis. However, the origin of the PUAs and their effects on natural phytoplankton assemblages remain unclear. Here we present data from four oceanographic cruises that took place during diatom blooms in the northern Adriatic Sea where concentrations of particulate and dissolved PUAs were monitored along with phytoplankton cell lysis. Cell lysis was positively correlated with both concentrations of particulate and dissolved PUAs (R = 0.69 and R = 0.77, respectively), supporting the hypothesis that these compounds are released by cell lysis. However, the highest concentration of dissolved PUAs (2.53 nM) was measured when cell lysis was high (0.24 d(-1)) but no known PUA-producing diatoms were detected, suggesting either that other organisms can produce PUAs or that PUA-producing enzymes retain activity extracellularly after diatom cells have lysed. Although in situ concentrations of dissolved PUAs were one to three orders of magnitude lower than those typically used in laboratory culture experiments, we argue that concentrations produced in the field could induce similar effects to those observed in culture and therefore may help shape plankton community composition and function in the oceans.

Integrated evaluation of environmental parameters influencing Vibrio occurrence in the coastal Northern Adriatic Sea (Italy) facing the Venetian lagoon.

In the marine environment, the persistence and abundance of Vibrio are related to a number of environmental parameters. The influence of the different environmental variables in determining the Vibrio occurrence could be different in the specific geographic areas around the world. Moreover, oceanographic parameters are generally interdependent and should not be considered separately when their influence on bacterial presence and concentration is tested. In this study, an integrated approach was used to identify key parameters determining the abundance of Vibrio spp in marine samples from the Venetian Lagoon in Italy, which is an important area for fish farming and tourism. Multivariate techniques have been adopted to analyze the dataset: using PCA, it was shown that a relatively high proportion of the total variance in this area was mainly due to two independent variables, namely salinity and temperature. Using cluster analysis, it was possible to categorize different groups with homogeneous features as regards space ("stations") and time ("seasons") distribution, as well as to quantify the values of environmental variables and the Vibrio abundances in each category. Furthermore, integrating key environmental factors and bacterial concentration values, it was possible to identify levels of salinity and sea surface temperature which were optimal for Vibrio concentration in water, plankton, and sediment samples. The identification of key environmental variables conditioning Vibrio occurrence should facilitate ocean monitoring, making it possible to predict unexpected variations in marine microflora which determine possible public health risks in coastal areas.

Quantification of dissolved and particulate polyunsaturated aldehydes in the Adriatic sea.

Polyunsaturated aldehydes (PUA) are supposed to play critical roles in chemically-mediated plankton interactions. Laboratory studies suggest that they act as mediators of chemical defense and chemical communication. PUA are oxylipins containing an α,ß,γ,δ-unsaturated aldehyde structure element and are mainly found in diatoms. We present here a detailed surface mapping of PUA during a spring bloom of the diatom Skeletonema marinoi in the Adriatic Sea. We monitored dissolved PUA, as well as particulate PUA, which are produced by phytoplankton after cell disintegration. Our survey revealed a patchy distribution of PUA and shows that at most stations S. marinoi is the major contributor to the overall PUA. Our data also suggest that lysis of a diatom bloom can contribute significantly to the dissolved PUA concentrations and that other producers, which are smaller in cell size compared to diatoms, have to be taken into account as well if the total PUA content of marine samples is considered. The analyses of samples collected in deeper water suggests that diatom contribution to PUA decreases with depth, while smaller-sized unidentified organisms take place as dominant contributors to the PUA concentrations.