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Metabarcoding of environmental DNA has provided striking insights into marine microbial diversity. With this approach, we assessed the diversity of metazoan assemblages and their temporal variations at the Long-Term Ecological Research site MareChiara (LTER-MC) in the Gulf of Naples (Mediterranean Sea). The Illumina sequencing of the V4-18S rRNA fragment from 48 surface samples collected from 2011 to 2013 produced a total of 5 011 047 marine metazoan reads. The normalized dataset was generally dominated by copepods (60.3%), followed by annelids (34.7%) mostly represented by the invasive benthic polychaete Hydroides elegans. Non-copepod holoplankton was mainly represented by siphonophores, rotifers, and appendicularians, with occasional mass occurrences of jellyfish. The rest of meroplankton (mainly molluscs, annelids, and anthozoans) showed a high diversity, covering all 11 zooplankton phyla, from Porifera to Craniata. A high number of copepod genera were identified, with seasonal recurrence matching patterns observed in 30 years of studies in the Gulf of Naples. Despite limitations related to the molecular marker resolution and reference dataset, the study provided valuable insights into diversity and seasonal patterns of the whole metazoan assemblage, expanding the knowledge on rare or hardly identifiable taxa and confirming DNA metabarcoding as a powerful approach to be integrated at LTER sites.

Since 2007, the non-indigenous calanoid copepod Pseudodiaptomus marinus Sato, 1913 has been increasingly recorded in numerous European sites, spreading at an unexpectedly fast pace over a short time-span. This species presents specific biological and behavioural traits which make it of particular interest for ecological and applied research topics. On 29–30 January 2018, 29 scientists from nine European Countries established the EUROBUS (Towards a EURopean OBservatory of the nonindigenous calanoid copepod Pseudodiaptomus marinUS) Working Group (WG). This WG aimed at creating a European network of institutions and researchers working on the various aspects of the biology and ecology of P. marinus, with an open forum where sharing experience and know-how among WG participants. This brought to an updated distribution map of P. marinus in European waters, as well as to the identification of priority research lines and potential joint initiatives under the WGEUROBUS umbrella. This contribution, stemming from the experts participating at the WG, represents the manifesto of the current and future initiatives developed within WGEUROBUS. info:eu-repo/semantics/publishedVersion

Crowding has a major impact on the dynamics of many material and biological systems, inducing effects as diverse as glassy dynamics and swarming. While this issue has been deeply investigated for a variety of living organisms, more research remains to be done on the effect of crowding on the behaviour of copepods, the most abundant metazoans on Earth. To this aim, we experimentally investigate the swimming behaviour, used as a dynamic proxy of animal adaptations, of males and females of the calanoid copepod Centropages typicus at different densities of individuals (10, 50 and 100 ind. l −1 ) by performing three-dimensional single-organism tracking. We find that the C. typicus motion is surprisingly unaffected by crowding over the investigated density range. Indeed, the mean square displacements as a function of time always show a crossover from ballistic to Fickian regime, with poor variations of the diffusion constant on increasing the density. Close to the crossover, the displacement distributions display exponential tails with a nearly density-independent decay length. The trajectory fractal dimension, D 3D ≅ 1.5, and the recently proposed ‘ecological temperature’ also remain stable on increasing the individual density. This suggests that, at least over the range of animal densities used, crowding does not impact on the characteristics of C. typicus swimming motion, and that a homeostatic mechanism preserves the stability of its swimming performance.

The distribution and reproductive traits of copepods of the genus Clausocalanus were investigated during the Atlantic Meridional Transect cruise AMT-15, in September–October 2004 to estimate their ecological niches and secondary production in the epipelagic layer along a latitudinal cline (48°N–40°S). The distribution patterns of selected environmental parameters, i.e., temperature, salinity and chlorophyll a concentration, enabled eco-provinces to be identified as described by Longhurst (2006) . Clausocalanus represented on average 34% of total copepod abundance, with a large predominance of adult females and copepodites over males. Among the eleven Clausocalanus species found during the survey, eight species showed a wide distributional range, i.e.,C. paululus, C. pergens, C. furcatus, C. arcuicornis, C. jobei, C. parapergens, C. lividus, and C. mastigophorus, while C. ingens, C. brevipes, and C. laticeps were recorded only in the South Atlantic. The smallest C. furcatus, C. paululus, and C. pergens together accounted for 85% of total Clausocalanus adult abundance. The ecological niches were clearly separated among congeners of similar size and largely overlapped in congeners whose size differed. The small- and medium-sized species, which are egg-sac-spawners, had smaller clutch size and lower egg-production rate than the larger broadcaster congeners. Nevertheless, embryo viability was lower in broadcasters, which may explain their low abundance in terms of lower recruitment. A sex ratio largely skewed toward females in all Clausocalanus species and the observation of viable eggs in successive clutches from isolated females seem to indicate that re-mating is not necessary in this genus. Broadcast-spawners showed the highest weight-specific fecundity rates in the genus but similar secondary production to sac-spawners despite the fact that they never occurred at high abundance. In light of their abundant occurrence in oceanic waters and well-defined ecological niches, Clausocalanus species may be considered as good indicators of environmental conditions and monitored in relation to changes in ecosystem structure due to climate change.

The LOHAFEX iron fertilization experiment was conducted for 39 days in the closed core of a cyclonic mesoscale eddy located along the Antarctic Polar Front in the Atlantic sector of the Southern Ocean. Mixed layer (ML) waters were characterized by high nitrate (~ 20 μM), low dissolved iron (DFe ~ 0.2 nM) and low silicate concentrations (below 1 μM) restricting diatom growth. Upon initial fertilization, chlorophyll-a doubled during the first two weeks and stabilized thereafter, despite a second fertilization on day 21, due to an increase in grazing pressure. Biomass at the different trophic levels was mostly comprised of small autotrophic flagellates, the large copepod Calanus simillimus and the amphipod Themisto gaudichaudii. The downward flux of particulate material comprised mainly copepod fecal pellets that were remineralized in the upper 150 m of the water column with no significant deeper export. DFe concentrations in the upper 200 m were not significantly affected by the two fertilizations but after day 14 showed a greater variability (ranging from 0.3 to 1.3 nM) without a clear vertical pattern. Particulate iron concentrations (measured after 2 months at pH 1.4) decreased with time and showed a vertical pattern that indicated an important non-biogenic component at the bottom of the mixed layer. In order to assess the contribution of copepod grazing to iron cycling we used two different approaches: first, we measured for the first time in a field experiment copepod fecal pellet concentrations in the water column together with the iron content per pellet, and second, we devised a novel analytical scheme based on a two-step leaching protocol to estimate the contribution of copepod fecal pellets to particulate iron in the water column. Analysis of the iron content of isolated fecal pellets from C. simillimus showed that after the second fertilization, the iron content per fecal pellet was ~ 5 fold higher if the copepod had been captured in fertilized waters. We defined a new fraction termed leachable iron (pH 2.0) in 48 h (LFe48h) that, for the conditions during LOHAFEX, was shown to be an excellent proxy for the concentration of iron contained in copepod fecal pellets. We observed that, as a result of the second fertilization, iron accumulated in copepod fecal pellets and remained high at one third of the total iron stock in the upper 80 m. We hypothesize that our observations are due to a combination of two biological processes. First, phagotrophy of iron colloids freshly formed after the second fertilization by the predominant flagellate community resulted in higher Fe:C ratios per cell that, via grazing, lead to iron enrichment in copepod fecal pellets in fertilized waters. Second, copepod coprophagy could explain the rapid recycling of particulate iron in the upper 100–150 m, the accumulation of LFe48h in the upper 80 m after the second fertilization and provided the iron required for the maintenance of the LOHAFEX bloom for many weeks. Our results provide the first quantitative evidence of the major ecological relevance of copepods and their fecal products in the cycling of iron in silicate depleted areas of the Southern Ocean., This work was supported by the Government of Spain (MINECO CTM2008-01864-E/ANT and CTM2014-59244-C3-3-R). LML was supported by the Campus de Excelencia Internacional Program (Ministerio de Educación, Cultura y Deportes, España).

We explored the small-scale behavior of the calanoid copepod Temora stylifera in relation to the diatoms Chaetoceros socialis , Leptocylindrus aporus , Leptocylindrus danicus and Pseudo-nitzschia calliantha offered as monospecific diets at similar carbon concentrations. These four diatoms are characterized by distinct size, shape and colony forming ability and are important components of the autumnal bloom co-occurring with the seasonal peak of T. stylifera abundance in Mediterranean coastal waters. High-speed video recordings showed that T. stylifera acquired cells in a suspension feeding mode while creating feeding currents. Copepod behavior was quantified in terms of feeding, motion, and grooming activities. T. stylifera spent more time in hovering than cruising in presence of all diets. The solitary L. aporus and P. calliantha elicited longer feeding bouts, lower appendage beat frequency and shorter grooming events compared to the colonial L. danicus and C. socialis . Overall the present results indicate that the behavioral responses of T. stylifera to different diatom diets were species-specific. The observed behavioral plasticity may help T. stylifera to adjust rapidly to changes in the food environment and this can be advantageous in exploiting short-lived phytoplankton blooms.

The planktonic tunicates appendicularians and thaliaceans are highly efficient filter feeders on a wide range of prey size including bacteria and have shorter generation times than any other marine grazers. These traits allow some tunicate species to reach high population densities and ensure their success in a favourable environment. However, there are still few studies focusing on which genes and gene pathways are associated with responses of pelagic tunicates to environmental variability. Herein, we present the effect of food availability increase on tunicate community and gene expression at the Marquesas Islands (South-East Pacific Ocean). By using data from the Tara Oceans expedition, we show that changes in phytoplankton density and composition trigger the success of a dominant larvacean species (an undescribed appendicularian). Transcriptional signature to the autotroph bloom suggests key functions in specific physiological processes, i.e., energy metabolism, muscle contraction, membrane trafficking, and proteostasis. The relative abundance of reverse transcription-related Pfams was lower at bloom conditions, suggesting a link with adaptive genetic diversity in tunicates in natural ecosystems. Downstream of the bloom, pelagic tunicates were outcompeted by copepods. Our work represents the first metaomics study of the biological effects of phytoplankton bloom on a key zooplankton taxon.

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.

We compare the long-term and seasonal patterns of abundance and phenology of the cyclopoid copepod Oithona similis at the L4 site (1988–2013) in the North Atlantic and at the LTER-MC site (1984–2013) in the Mediterranean Sea to investigate whether high temperature limits the occurrence of this species with latitudinal cline. The two sites are well suited to testing this hypothesis as they are characterized by similar chlorophyll a concentration (Chl a) but different temperature [sea surface temperature (SST)]. The abundance of O. similis at L4 was ∼10 times higher than at LTER-MC. Moreover, this species had several peaks of abundance during the year at L4 but a single peak in spring at LTER-MC. The main mode of temporal variability in abundance was seasonal at both sites. The abundance of O. similis was negatively correlated with SST only at LTER-MC, whereas it was positively correlated with Chl a at both sites. Oithona similis had a temperature optimum between 15 and 20°C reaching maximum abundance at ∼16.5°C at LTER-MC, but showed no Chl a optimum at either site. We conclude that the abundance of O. similis increases with prey availability up to 16.5°C and that temperature >20°C represents the main limiting factor for population persistence.

While the effects of industrial contamination in coastal areas may persist for years in benthos communities, plankton should not show permanent impairments because of their high spatial dynamics, fast turnover times and pronounced seasonality. To test this hypothesis, in 2019 we conducted five surveys in the Bay of Pozzuoli (Gulf of Naples, Mediterranean Sea), in front of a dismissed steel factory and in the adjacent inshore coastal waters. High seasonal variability was observed for bacteria, phytoplankton and mesozooplankton, whereas plankton spatial gradients were relatively smooth during each survey. Plankton biomass and diversity did not reveal any effects of past industrial activities not even at the innermost stations of the Bay, which however showed some signals of present anthropogenic pressure. Hydrodynamic and morphological features likely play a prominent role in maintaining a relatively good status of the plankton of the Bay, which hints at the relevance of coastal circulation and meteorological dynamics to revitalize areas impacted by human activities.

Microplastics (MP) have gained enormous attention in the last years as they are among the major contaminants of soil and oceans, able to insert themselves in the food chain and cause severe damage to plants life, fauna and human health. Many techniques have been proposed to analyze the abundance, concentration and presence of such material in water, but data are still incomplete. Here, we propose digital holography (DH) as a non-invasive technique to search for and reconstruct the shape of different types of MP. We prove DH is a suitable tool for identifying MP also in the presence of other objects in water, such as diatoms.

Suspensions of small planktonic copepods represent a special category in the realm of active matter, as their size falls within the range of colloids, while their motion is so complex that it cannot be rationalized according to basic self-propelled particle models. Indeed, the wide range of individual variability and swimming patterns resemble the behaviour of much larger animals. By analysing hundreds of three dimensional trajectories of the planktonic copepod Clausocalanus furcatus we investigate the possibility of detecting how the motion of this species is affected by different external conditions, such as the presence of food and the effect of gravity. While this goal is hardly achievable by direct inspection of single organism trajectories, we show that this is possible by focussing on simple average metrics commonly used to characterize colloidal suspensions, such as the mean square displacement and the dynamic correlation functions. We find that the presence of food leads to the onset of a clear localization that separates a short-time ballistic from a long-time diffusive regime. Such a benchmark reflects the tendency of C. furcatus to remain temporally feeding in a limited space and disappears when food is absent. Localization is clearly evident in the horizontal plane, but is negligible in the vertical direction, due to the effect of gravity. Our results suggest that simple average descriptors may provide concise and useful information on the swimming properties of planktonic copepods, even though single organism behaviours are strongly heterogeneous.

Spread of alien species (AS) is a serious threat to marine habitats and analysis of principal descriptors of their occurrence is pivotal to set reliable conservation strategies. 2. In order to assess the susceptibility of marine habitats to biological invasions, a dataset was gathered of the occurrence of 3899 species from 29 phyla, taken from 93 marine sites located along the Italian coast in the period 2000-2012.

Mortality affects the dynamics of zooplankton populations with important effects on trophic interactions and biogeochemical fluxes in marine environments, but is still one of the processes least investigated in the field. In the present study, the non-predatory mortality in copepod assemblages and species was investigated by applying the neutral red staining method to identify and quantify copepod carcasses throughout an annual cycle in a Mediterranean coastal site (station LTER-MC in the inner Gulf of Naples). Carcasses accounted on average for 10.3% (±9.7%) of total copepod abundance and were most abundant in spring, summer and autumn. Carcasses were represented predominantly by copepodites (78.9 ± 22.0%) and occurred more frequently and abundantly in calanoids than in other copepod orders, with interspecific differences in their abundance and temporal patterns. Using carcass abundances from field data and decomposition times from laboratory observations, we estimated non-predatory mortality rates of key calanoids that are common and abundant in Mediterranean coastal waters. Non-predatory mortality rates averaged 0.13 day−1 in Paracalanus parvus, 0.07 day−1 in Clausocalanus spp., 0.06 day−1 in Temora stylifera and 0.04 day−1 in Acartia clausi. Non-predatory mortality rates in these populations were not correlated with temperature, salinity or chlorophyll a.

Anchovies represent the largest world’s marine fish catches and the current threats on their populations impose a sustainable exploitment based on sound scientific information. In the European anchovy (Engraulis encrasicolus), the existence of several populations has been proposed but a global view is missing. Using a multidisciplinary approach, here we assessed the divergence among different ecotypes and its possible causes. SNPs have revealed two functionally distinct ecotypes overlapping in the Central Mediterranean, with one ecotype confined near the river estuaries. The same SNPs outliers also segregated two distinct populations in the near Atlantic, despite their large spatial distance. In addition, while most studies suggested that adaptation to low salinity is key to divergence, here we show that the offshore ecotype has higher environmental tolerance and an opportunistic feeding behaviour, as assessed by the study of environmental conditions, anchovy diet and trophic levels, and passive egg dispersal. These results provide insights into the anchovy evolutionary history, stressing the importance of behaviour in shaping ecotypes.

International audience; Copepods belonging to the Oncaeidae family are commonly and abundantly found in marine zooplankton. In the Mediterranean Sea, forty-seven oncaeid species occur, of which eleven in the Gulf of Naples. In this Gulf, several Oncaea species were morphologically analysed and described at the end of the XIX century by W. Giesbrecht. In the same area, oncaeids are being investigated over seasonal and inter-annual scales at the long-term coastal station LTER-MC. In the present work, we identified six oncaeid species using the nuclear ribosomal internal transcribed spacers (ITS rDNA) and the mitochondrial cytochrome c oxidase subunit I (mtCOI). Phylogenetic analyses based on these two genomic regions validated the sisterhood of the genera Triconia and the Oncaea sensu stricto. ITS1 and ITS2 phylogenies produced incongruent results about the position of Oncaea curta, calling for further investigations on this species. We also characterised the ITS2 region by secondary structure predictions and found that all the sequences analysed presented the distinct eukaryotic hallmarks. A Compensatory Base Change search corroborated the close relationship between O. venusta and O. curta and between O. media and O. venusta already identified by ITS phylogenies. The present results, which stem from the integration of molecular and morphological taxonomy, represent an encouraging step towards an improved knowledge of copepod biodiversity: The two complementary approaches, when applied to long-term copepod monitoring, will also help to better understanding their genetic variations and ecological niches of co-occurring species.

The internal organization of plankton communities plays a key role in biogeochemical cycles and in the functioning of aquatic ecosystems. In this study, the structure of a marine plankton community (including both unicellular and multicellular organisms) was inferred by applying an ecological network approach to species abundances observed weekly at the long-term ecological research station MareChiara (LTER-MC) in the Gulf of Naples (Tyrrhenian Sea, Mediterranean Sea) in the summers of 2002–2009. Two distinct conditions, characterized by different combination of salinity and chlorophyll values, alternated at the site: one influenced by coastal waters, herein named ‘green’, and the other reflecting more offshore conditions, named ‘blue’. The green and blue ‘phases’ showed different keystone biological elements: namely, large diatoms and small-sized flagellates, respectively. Several correlations amongst species belonging to different trophic groups were found in both phases (connectance ~0.30). In the green phase, several links between phytoplankton and mesozooplankton and within the latter were detected, suggesting matter flow from microbes up to carnivorous zooplankton. A microbial-loop-like sub-web, including mixo- and heterotrophic dinoflagellates and ciliates, was present in the green phase, but it was relatively more important in the blue phase. The latter observation suggests a more intense cycling of matter at the microbial trophic level in the blue phase. These results show that different modes of ecological organization can emerge from relatively small changes in the composition of aquatic communities coping with environmental variability. This highlights a significant plasticity in the internal structure of plankton webs, which should be taken into account in predictions of the potential effects of climatic oscillations on aquatic ecosystems and biogeochemical cycles therein.

Mazzocchi, M. G. et al., The cruises conducted in the spring and autumn of 2008 in the frame of the European project SESAME represented the first coordinated surveys that allowed acquiring a quasi-synoptic picture of epipelagic mesozooplankton in most regions of the Mediterranean Sea. Seasonal differences were recorded in biomass, total abundance, and community composition and structure. In both seasons, it did not appear a clear west-east decreasing gradient in total standing stock, but rather regional discontinuities. However, west or east preferences were observed in the distribution of some zooplanktonic groups and copepod species. An artificial neural network analysis (SOM) identified, in both seasons, a clear mesozooplankton regionalization, which resembled the autotrophic regimes based on color remote sensing data. The correspondence between the distribution of zooplankton communities and the trophic regimes appeared more precise in spring, when the increased concentration of chlorophyll a makes the Mediterranean Sea a more heterogeneous environment, but it was still visible in the more uniform oligotrophic autumn conditions. Three distinct types of mesozooplankton communities seem to flourish in the investigated regions: the first type is the most widespread and thrives in the >non-blooming> areas, the second type occurs in the >intermittently-blooming> areas, and the third type is a characteristic of areas with recurrent and intense phytoplankton blooms. Overall, the well defined regionalization of mesozooplankton that appears from our results corroborates the view of the Mediterranean Sea as a mosaic environment, as previously emerged from the analyses of different biological compartments. © 2013 Elsevier B.V., This is a contribution of the European IP SESAME (FP6), Work Package-2 on “Coordinated acquisition of new multidisciplinary data in the Southern European Seas”.

Dispersal plays a key role to connect populations and, if limited, is one of the main processes to maintain and generate regional biodiversity. According to neutral theories of molecular evolution and biodiversity, dispersal limitation of propagules and population stochasticity are integral to shaping both genetic and community structure. We conducted a parallel analysis of biological connectivity at genetic and community levels in marine groups with different dispersal traits. We compiled large data sets of population genetic structure (98 benthic macroinvertebrate and 35 planktonic species) and biogeographic data (2193 benthic macroinvertebrate and 734 planktonic species). We estimated dispersal distances from population genetic data (i.e., FST vs. geographic distance) and from β-diversity at the community level. Dispersal distances ranked the biological groups in the same order at both genetic and community levels, as predicted by organism dispersal ability and seascape connectivity: macrozoobenthic species without dispersing larvae, followed by macrozoobenthic species with dispersing larvae and plankton (phyto- and zooplankton). This ranking order is associated with constraints to the movement of macrozoobenthos within the seabed compared with the pelagic habitat. We showed that dispersal limitation similarly determines the connectivity degree of communities and populations, supporting the predictions of neutral theories in marine biodiversity patterns., RADIALES (IEO)

Plankton is a hugely diverse community including both unicellular and multicellular organisms, whose individual dimensions span over seven orders of magnitude. Plankton is a fundamental part of biogeochemical cycles and food-webs in aquatic systems. While knowledge has progressively accumulated at the level of single species and single trophic processes, the overwhelming biological diversity of plankton interactions is insufficiently known and a coherent and unifying trophic framework is virtually lacking. We performed an extensive review of the plankton literature to provide a compilation of data suitable for implementing food-web models including plankton trophic processes at high taxonomic resolution. We identified the components of the plankton community at the Long Term Ecological Research Station MareChiara in the Gulf of Naples. These components represented the sixty-three nodes of a plankton food-web. To each node we attributed biomass and vital rates, i.e. production, consumption, assimilation rates and ratio between autotrophy and heterotrophy in mixotrophic protists. Biomasses and rates values were defined for two opposite system’s conditions; relatively eutrophic and oligotrophic states. We finally identified 817 possible trophic links within the web and provided each of them with a relative weight, in order to define a diet-matrix, valid for both trophic states, which included all consumers, fromn anoflagellates to carnivorous plankton. Vital rates for plankton resulted, as expected, very wide; this strongly contrasts with the narrow ranges considered in plankton system models implemented so far. Moreover, the amount and variety of trophic links highlighted by our review is largely excluded by state-of-the-art biogeochemical and food-web models for aquatic systems. Plankton models could potentially benefit from the integration of the trophic diversity outlined in this paper: first, by using more realistic rates; second, by better defining trophic roles of consumers in the planktonic web. We suggest that most trophic habits present in planktonic organisms must be contemplated in new generation plankton models.

1. Spread of alien species (AS) is a serious threat to marine habitats and analysis of principal descriptors of their occurrence is pivotal to set reliable conservation strategies. 2. In order to assess the susceptibility of marine habitats to biological invasions, a dataset was gathered of the occurrence of 3899 species from 29 phyla, taken from 93 marine sites located along the Italian coast in the period 2000–2012. 3. In total, 61 AS belonging to 11 phyla have been recorded. Invertebrates were the most represented (63%). Alien species were found in all the habitats examined (EUNIS, level 2), although they showed highest abundance in benthic habitats. Most of the AS were associated with a single EUNIS habitat, while some of them were present in more than one habitat. Trans-habitat occurrence suggests the potential invasiveness of AS. 4. According to statistical analysis, AS recorded could have been more numerous, since some of the marine habitats seemed to be still unsaturated. The model that best describes the spread of AS takes account of both native species richness (Rn) and EUNIS habitat type as explanatory variables. The number of observed AS was directly related to Rn and it was highest in rocky circalittoral and infralittoral habitats. 5. The results of this macro-ecological study focus on the importance of performing large-scale studies, since adopting ecosystem approaches to marine invasion management seems especially fruitful. 6. The results, moreover, highlight the importance of AS monitoring of different habitats, from those subjected to anthropogenic pressure, historically considered to be hubs of introduction of AS, to the most biologically rich and diverse marine habitats. Indeed, it is necessary to set monitoring strategies to detect the introduction, the distribution and persistence of AS over time. These recommendations are especially significant in the light of the strategic plans currently under formulation in Mediterranean countries with regard to AS monitoring

We analyzed the copepod assemblages over two decades (1984–2006) in a coastal ongoing time-series at Station MC in the inner Gulf of Naples (Tyrrhenian Sea, Western Mediterranean), which is part of the International network of Long-Term Ecological Research (LTER). The seasonal and interannual time courses of species abundance and composition were related to depth integrated temperature, salinity, and chlorophyll a , which provide essential information on the local environmental dynamics. Our aims were to characterize the main modes of copepod variability and to highlight possible changes occurred in the period in relation to the local environmental dynamics. Despite the great variability at seasonal and interannual scales, our site did not show evidence of discontinuities or trends in water column properties as compared to other Mediterranean sites for the same period, which we interpret as resulting from the position of Station MC that is exposed to the influence of a complex climate forcing. Abrupt changes did not appear for most of the key representative species (e.g., Acartia clausi , Centropages typicus , Paracalanus parvus , Temora stylifera , and juveniles of Clausocalanus spp./ P. parvus ) beyond the high interannual variability in the abundance patterns. A few indications suggest that our station might have acquired less coastal characters (e.g., decreasing chlorophyll a concentrations), but the signals from the copepod assemblages appeared only in rare species. A significant increase was observed in the occurrence of some typical offshore calanoids (e.g., Neocalanus gracilis , Scolecithricella spp.), while a few species typical of confined areas disappeared (e.g., Acartia margalefi , Paracartia latisetosa ). STATICO analysis showed a significant resilience in the seasonal cycle of the copepod assemblages at Station MC, even when there was high variability in the environmental parameters. While the changes recorded in the least abundant species may be indicative of long-term variations, we interpret the overall persistence of the basic community as resulting from the flexibility of coastal species to adapt to variation in environmental conditions.

Coastal zooplankton have been investigated since 1984 at a Long Term Ecological Research station MC (LTER-MC) in the inner Gulf of Naples (Tyrrhenian Sea, Western Mediterranean). The sampling site, located between the littoral and the open sea systems, has very active hydrography that affects plankton communities. The present work was aimed at establishing whether, in such a dynamic and variable environment, species associations and homogeneous periods could be identified as characteristic and stable features of the mesozooplankton over the period 1984–2006. Hierarchical clustering was applied to assess species associations based on a matrix of similarities between species (R-mode), and homogeneous periods based on a matrix of similarities between observations (Q-mode). The Indicator Value index [IndVal, Dufrene and Legendre (1997) Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr., 67, 345–366] was calculated to identify species characterizing each period. Five taxonomic groups with well-defined composition and abundance were identified as robust associations that likely reflect different modes of community functioning. The temporal course of these associations was largely shaped by strong seasonal forcing comprising both physical and biological (e.g. trophic) signals. These associations persisted over the long term, thus indicating some stable characters in the Naples zooplankton time-series, providing evidence of resilience in communities in highly variable coastal conditions.

44 pages, 19 figures, 7 tables, We present an overview of the plankton studies conducted during the last 25 years in the epipelagic offshore waters of the Mediterranean Sea. This quasi-enclosed sea is characterized by a rich and complex physical dynamics with distinctive traits, especially in regard to the thermohaline circulation. Recent investigations have basically confirmed the long-recognised oligotrophic nature of this sea, which increases along both the west-east and the north-south directions. Nutrient availability is low, especially for phosphorous (N:P up to 60), though this limitation may be buffered by inputs from highly populated coasts and from the atmosphere. Phytoplankton biomass, as chl a, generally displays low values (less than 0.2 μg chl a l−1) over large areas, with a modest late winter increase. A large bloom (up to 3 μg l−1) is observed throughout the late winter and spring exclusively in the NW area. Relatively high biomass values are recorded in fronts and cyclonic gyres. A deep chlorophyll maximum is a permanent feature for the whole basin, except during the late winter mixing. It is found at increasingly greater depths ranging from 30 m in the Alboran Sea to 120 m in the easternmost Levantine basin. Primary production reveals a west-east decreasing trend and ranges between 59 and 150 g C m−2 y−1 (in situ measurements). Overall, the basin is largely dominated by small autotrophs, microheterotrophs and egg-carrying copepod species. The microorganisms (phytoplankton, viruses, bacteria, flagellates and ciliates) and zooplankton components reveal a considerable diversity and variability over spatial and temporal scales, although the latter is poorly studied. Examples are the wide diversity of dinoflagellates and coccolithophores, the multifarious role of diatoms or picoeukaryotes, and the distinct seasonal or spatial patterns of the species-rich copepod genera or families which dominate the basin. Major dissimilarities between western and eastern basins have been highlighted in species composition of phytoplankton and mesozooplankton, but also in the heterotrophic microbial components and in their relationships. Superimposed to these longitudinal differences, a pronounced biological heterogeneity is also observed in areas hosting deep convection, fronts, cyclonic and anti-cyclonic gyres or eddies. In such areas, the intermittent nutrient enrichment promotes a switching between a small-sized microbial community and diatom-dominated populations. A classical food web readily substitutes the microbial food web in these cases. These switches, likely occurring within a continuum of trophic pathways, may greatly increase the flux towards higher trophic levels, in spite of the apparent heterotrophy. Basically, the microbial system seems to be both bottom-up and top-down controlled. A >multivorous web> is shown by the great variety of feeding modes and preferences and by the significant and simultaneous grazing impact on phytoplankton and ciliates by mesozooplankton, This paper was supported by a grant provided by the European Network of Excellence EUR-OCEANS contract number 511106-2. We thank the authors who directly provided their data. Additional funds were provided to D. Vaqué by the MICROVIS project (CTM 2007-62140) (Spanish Ministry of Science) and to M. Ribera d’Alcalá by the EU IP SESAME (contract number 036949)

Prey capture is a fundamental process for the success of copepods in food-diluted environments. This process is influenced by several factors, including swimming and predatory habits. This work is aimed at characterising the kinematic and fractal properties of the swimming trajectories and reconstructing the predatory horizon of the small calanoid copepod Clausocalanus furcatus. Results indicate that the motion of C. furcatus resembles a random process, mainly evolving in one direction, whereas its predatory horizon is confined to a small region frontal to the anterior end of the copepod. These outcomes are discussed in terms of specific adaptation taking into account the natural conditions experienced by C. furcatus in its environment.

Crise, A. ... et. al.-- 12 pages, 1 figure, 5 tables, PERSEUS project aims to identify the most relevant pressures exerted on the ecosystems of the Southern European Seas (SES), highlighting knowledge and data gaps that endanger the achievement of SES Good Environmental Status (GES) as mandated by the Marine Strategy Framework Directive (MSFD). A complementary approach has been adopted, by a meta-analysis of existing literature on pressure/impact/knowledge gaps summarized in tables related to the MSFD descriptors, discriminating open waters from coastal areas. A comparative assessment of the Initial Assessments (IAs) for five SES countries has been also independently performed. The comparison between meta-analysis results and IAs shows similarities for coastal areas only. Major knowledge gaps have been detected for the biodiversity, marine food web, marine litter and underwater noise descriptors. The meta-analysis also allowed the identification of additional research themes targeting research topics that are requested to the achievement of GES. © 2015 The Authors, This work has been partially funded by the EC FP7 PERSEUS Project (Grant. Agr. 287600)

The Mediterranean Sea is one of the areas most affected by alien invasions, which are continuously increasing also due to intense human activities and environmental changes that favor the introduction of species previously unable to colonize the basin. This is the case of the copepods of the genus Pseudodiaptomus, first described in the Indian Ocean and considered one of the most resistant to unfavorable conditions but never recorded in the Mediterranean until 2011 though present in adjacent seas. Pseudodiaptomus marinus, in particular, is common in shallow marine-brackish waters and is one of the species often found in ballast waters and in aquaculture plants. Native of Japan, it has started spreading since 1950s and its populations have established in several harbours, eutrophic inlets and lagoons along the coasts of the Pacific and Indian Oceans. In the last few years, P. marinus has been increasingly reported in European Seas (Mediterranean Sea and North Sea). In this paper, we review the invasion history of this species with a special emphasis on its records in the Mediterranean Sea, and its occurrence and establishment in Sicilian waters. We also compare the biological traits and population dynamics of P. marinus with those of other representative of the genus and discuss about the possible mechanisms of introduction in new environments. Aim of our work is to understand the reasons of successful invasion of P. marinus and the environmental and biological factors that may lead to its further biogeographic expansion.

Mating behaviour is an emerging topic in the study of copepod ecology, having important consequences at both individual and population levels. Mate-finding has been shown to depend upon the perception of cues released by the conspecific, as well as on the swimming behaviour of both the male and the female. In order to understand the differences in the strategies adopted in the pre-copulatory phase by males and females, we analysed the swimming tracks of the freshwater calanoid Leptodiaptomus ashlandi. The three-dimensional trajectories described by males and females have been analysed in terms of their three-dimensional fractal dimension D3D, a parameter which can synthetically but appropriately characterize the degree of contortion of a three-dimensional path. D3D values for L. ashlandi males during the pre-detection phase are higher than the corresponding ones for females, indicating that the male explores more extended volumes of fluid, thus increasing the probability of encountering a female.

We integrated field and laboratory data with modeling to determine the extent to which the temporal patterns in population abundance of a copepod species as observed at sea may be explained by differences in production and mortality rates due to diet. A Lagrangian individual-based model utilizing birth and mortality rates whose values and variance were derived from the effects of dietary composition was implemented to simulate the growth of the multi-staged population of Temora stylifera. The four diets considered were represented by unialgal cultures of the dinoflagellate Prorocentrum minimum or the diatom Thalassiosira rotula, a mixture of the two species, and natural particle assemblages

In the future, if marine science is to achieve any progress in addressing biological diver- sity of ocean plankton, then it needs to sponsor development of new technology. One requirement is the development of high-resolution sensors for imaging field-collected and in situ specimens in a non-invasive manner. The rapid automatic categorisation of species must be accompanied by the creation of very large distributed databases in the form of high-resolution 3D rotatable images of species, which could become the standard reference source for automatic identification. These 3D images will serve as classification standards for field applications, and (in adjusted optical quality) as training templates for image analysis systems based on statistical and other pattern-matching processes. This paper sets out the basic argument for such developments and proposes a long-term solution to achieve these aims.

Three-dimensional swimming, in 20 trajectories of Daphnia pulex, was characterized in terms of its kinematic properties. Results show that the random component was stronger than the deterministic component at time scales >1-2 s. Such random movements may have evolved to outwit potential predators and prey.

Copepods of the genus Clausocalanus Giesbrecht, 1888 are among the most abundant calanoids in the Mediterranean Sea, both in coastal and offshore regions. The vertical distribution of C. arcuicornis, C. furcatus, C. jobei, C. lividus, C. mastigophorus, C. parapergens, C. paululus, and C. pergens, which co-occur in the upper 200 m in the Gulf of Naples (Tyrrhenian Sea), was investigated during an annual sampling cycle conducted at an offshore station in 2002. The quantitative data on distribution of each species were analysed in relation to the environmental parameters. The patterns that we observed in the seasonal cycles and vertical distribution provided insights on the ecological niches of the eight Clausocalanus species.

The population structure (NI to adults) of the copepods Centropages typicus and Temora stylifera was investigated in parallel at two sites in the eutrophic and oligotrophic waters of the Gulf of Naples (Tyrrhenian Sea) with the aim of identifying the environmental factors that modulate the occurrence of these calanoids that are abundant in the neritic waters of the Mediterranean Sea. Monthly zooplankton sampling was conducted from May 1999 to June 2000 in 5–7 discrete depth layers in the upper 200 m with a 70-μm mesh net. Temperature, salinity, and fluorescence profiles were recorded by CTD casts. The remarkable differences observed in the temporal and spatial patterns of the two populations throughout their developmental cycles seem to result more from different responses to environmental factors than from different life-history traits. C. typicus was abundant in early to mid-summer and was distributed in the whole water column; T. stylifera followed in late summer–autumn and occurred preferentially in the upper 30 m. Temperature was the main regulating factor for T. stylifera. The availability of autotrophic food appeared relatively more important for C. typicus, although unequivocal signals were not observed, suggesting that more complex trophic interactions likely affect the dynamics of the two populations.

The objective of this research was to quantify the abundance and distribution of nanoplanktonic particles available to frequently occurring juvenile and adult planktonic copepods in the epipelagial of the open subtropical and tropical ocean. We chose locations off Puerto Rico and off the Bahamas to achieve our purpose. The hydrography for all stations revealed a thin upper mixed layer, a thermocline with varying temperature decreases, salinities approaching and surpassing 37‰, and chlorophyll concentrations increasing with depth from 0.05 to 0.25 μg l -1 . Nanoplankton was grouped into five size ranges from 2-4 to 10-20 μm equivalent spherical diameter (ESD), and four cell types (photosynthetic nanoflagellate, Pnano; heterotrophic nanoflagellate, Hnano; photosynthetic dinflagellate, Pdino; heterotrophic dinoflagellate, Hdino). The 10-20 μm range had the highest total cell volume, and the 8-10 μm range the lowest values; for cell type, volumes of Pnano and Hdino were highest, and Hnano lowest. Total particle concentrations covering all depths and stations ranged from 3.4 to 17.5 μg C 1 -1 . At a large percentage of sampled depths total cell volumes of heterotrophs came close to those of autotrophs. Total autotroph volume was higher in the smaller size ranges than in the larger ones, while the opposite was found for heterotrophs. Calculations for ingestion rates of females of the oceanic calanoid Paracalanus aculeatus at environmental food levels and size distributions revealed that the 10-20 μm ESD particle size fraction contributed between 46 and 82% of the total ingested nanoplanktonic cell C. At nearly all the observed abundances in our study this copepod would obtain sufficient amounts for basal metabolic needs, and at the majority of stations and depths have additional energy assimilated to cover enhanced metabolic demands and limited reproduction.

The goal of our project is to increase our knowledge about the co-existence of the numerous small copepod species in the subtropical and tropical open oceans by combining experimental and field studies. In this paper we present the vertical distribution of the copepod assemblages collected in the upper 100 m of the water column during a diel cycle at an oceanic site off Bermuda in summer 2000. Our quantitative analysis ranged from the gross features of the community to order, genus, species and sex, ending with the population structure for one representative calanoid (Mecynocera clausii). Total abundance of copepods sampled with both 63 and 200 μm meshes increased from near the surface to 100 m depth. Of the four orders, the Calanoida and Cyclopoida increased slightly with depth, whereas the Poecilostomatoida hardly occurred in the upper 30 m and then increased by more than one order of magnitude towards 100 m depth. Harpacticoida showed even abundance through the vertical column during the night. Some abundant genera/species displayed hardly any or small differences in abundance over the upper 100 m (e.g. Calocalanus, Oithona); that distribution also showed that genera with entirely different motion and feeding behaviour can occupy the same vertical environment. A second group appeared to avoid the upper mixed layer and upper thermocline almost completely (e.g. Oncaea, Mecynocera clausii, Clausocalanus pergens, Eucalanus hyalinus, Haloptilus longicornis). A third limited group seemed to prefer the upper mixed layer (e.g. Cliusocalanus furcatus). Some genera were characterized by a paucity of males whereas others had as many males as females. Our results showed that quantifying the distribution and abundance of genera, species and their copepodid stages seems to be the preferable approach, leading towards an understanding of the functioning of the epipelagial copepod community. We hypothesize that a combination of variables, including feeding behaviour, temperature preference and also predation, could determine the ranges of vertical distribution, which would be flexible because of the effect of each of the variables. It is reasoned that predation was of minor impact in these oligotrophic waters because of the very low abundance of carnivorous species.

Several dinoflagellate species form nonmotile, thick-walled resting cysts in their life cycle. Cysts can be ingested by planktonic and benthic organisms, but there is scarce information concerning their survival after the passage through the digestive apparatus of the grazers. We tested the germination capability of cysts produced by two neritic dinoflagellates, Scrippsiella trochoidea (F. Stein) A.R. Loeblich and Scrippsiella ramonii Montresor, after their ingestion by four copepod species. Experiments have been carried out with four species: Acartia clausi Giesbrecht, 1889; Centropages typicus Kroyer, 1849; Temora stylifera Dana, 1849; and Clausocalanus lividus Frost and Fleminger, 1968. Copepods were fed either with motile cells or cysts, and feeding and clearance rates were estimated for A. clausi , C. lividus and T. stylifera . Grazing rates on both dinoflagellates was much higher for vegetative cells than for cysts. Resting cysts were isolated from the faecal pellets and incubated to test their germination capability. S. trochoidea cysts eaten by C. typicus and T. stylifera showed a high germination rate, while cysts of the same species were not viable after the passage through the gut of A. clausi and C. lividus . In contrast, S. ramonii cysts were never able to germinate after being ingested by copepods. The observed variation in viability among the two cyst types and the different survival rates observed for S. trochoidea cysts might be related to differences in cyst morphology and to differences in the digestive process among the tested copepod species.

Individual adult females of the cyclopoid copepod Oithona plumifera were continuously observed for up to 60 min to evaluate their motion and feeding behaviour. These females move only about five times per minute, each time with a brief relocation jump. They sink rather slowly while mainly in a horizontal position, which enhances the probability of encountering suitable food particles. They perceive food particles hydrodynamically via the setae on their first antennae, their prosome appendages and their abdomen. They appear to be able to obtain sufficient food even at tropical open ocean food abundances. A comparison with the calanoid Clausocalanus furcatus reveals notable differences in behaviour between these two species, which dominate the copepod assemblages in oligotrophic environments.