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In recent years, the application of metagenomics techniques has advanced our understanding of plankton communities and their global distribution. Despite this progress, the relationship between the abundance distribution of diatom species and varying marine environmental conditions remains poorly understood. This study, leveraging data from the Tara Oceans expedition, tests the hypothesis that diatoms in sampled stations display a consistent species abundance distribution structure, as though they were sampled from a single ocean-wide metacommunity. Using a neutral sampling theory, we thus develop a framework to estimate the structure and diversity of diatom communities at each sampling station given the shape of the species abundance distribution of the metacommunity and the information of a reference station. Our analysis reveals a substantial temperature gradient in the discrepancies between predicted and observed biodiversity across the sampled stations. These findings challenge the hypothesis of a single neutral metacommunity, indicating that environmental differences substantially influence both the composition and structure of diatom communities.

Marine plankton play a crucial role in carbon storage, global climate, and ecosystem function. Planktonic ecosystems are embedded in patches of water that are continuously moving, stretching, and diluting. These processes drive inhomegeneities on a range of scales, with implications for the integrated ecosystem properties, but are hard to characterize. We present a theoretical framework that accounts for all these aspects; tracking the water patch hosting a drifting ecosystem along with its physical, environmental, and biochemical features. The theory resolves patch dilution and internal physical mixing as a function of oceanic strain and diffusion. Ecological dynamics are parameterized by an idealized nutrient and phytoplankton population and we specifically capture the time evolution of the biochemical spatial variances to represent within-patch heterogeneity. We find that, depending only on the physical processes to which the water patch is subjected, the plankton biomass response to a resource perturbation can vary in size up to six times. This work indicates that we must account for these processes when interpreting and modeling marine ecosystems and provides a framework with which to do so.

AbstractThe study of connectivity patterns in networks has brought novel insights across diverse fields ranging from neurosciences to epidemic spreading or climate. In this context, betweenness centrality has demonstrated to be a very effective measure to identify nodes that act as focus of congestion, or bottlenecks, in the network. However, there is not a way to define betweenness outside the network framework. By analytically linking dynamical systems and network theory, we provide a trajectory-based formulation of betweenness, called Lagrangian betweenness, as a function of Lyapunov exponents. This extends the concept of betweenness beyond the context of network theory relating hyperbolic points and heteroclinic connections in any dynamical system to the structural bottlenecks of the network associated with it. Using modeled and observational velocity fields, we show that such bottlenecks are present and surprisingly persistent in the oceanic circulation across different spatio-temporal scales and we illustrate the role of these areas in driving fluid transport over vast oceanic regions. Analyzing plankton abundance data from the Kuroshio region of the Pacific Ocean, we find significant spatial correlations between measures of diversity and betweenness, suggesting promise for ecological applications.

Marine organisms can disperse actively and passively. During early life stages, eggs and larvae of a plethora of taxa may be subject to transport over large distances by ocean currents, while adults often exhibit rather sedentary behaviors. Next generation stock assessment schemes broadly call for the need to include climate and oceanographic considerations, yet little progress has been made globally towards this direction. Red mullet Mullus barbatus is an important demersal resource in the Mediterranean with relatively long pelagic early-life stages suggesting a large dispersal potential. Here, we use two different putative stocks of red mullet in the Western Mediterranean Sea to test the effects of surface currents on the demography and inter- and intra-stock individual exchanges. The connectivity between spawning and nursery areas was investigated by simulating multi-annual lagrangian indices of larval retention, import, and self-recruitment. Our study demonstrates a certain degree of inter-stock connectivity along the main corridors of transport but also a substantial variability with some years when population sub-units appear to be more isolated. Regarding intra-stock metrics, we found that the main persistent spawning area of the Northern Spanish Coast stock may act as an export of individuals ending up in coastal nursery areas further south within or even outside the boundaries of the stock unit. By evaluating the passive dispersal and its impact on the spatial population structure and the intra-stock spatial dynamism of an exploited resource, our study provides insights to improve fisheries management with the incorporation of information on physical oceanography (from observing systems and operational models). Importantly, our methodology may readily be applied to other systems and/or be combined with additional methods (e.g., chemical, molecular or artificial markers) to facilitate the improvement of current management schemes and inform conservation stra

For more than a decade, high-throughput sequencing has transformed the study of marine planktonic communities and has highlighted the extent of protist diversity in these ecosystems. Nevertheless, little is known relative to their genomic diversity at the species-scale as well as their major speciation mechanisms. An increasing number of data obtained from global scale sampling campaigns is becoming publicly available, and we postulate that metagenomic data could contribute to deciphering the processes shaping protist genomic differentiation in the marine realm. As a proof of concept, we developed a FAIR pipeline and focused on the Mediterranean Sea to study three a priori abundant protist species: Bathycoccus prasinos, Pelagomonas calceolata and Phaeocystis cordata. We compared the genomic differentiation of each species in light of geographic, environmental and oceanographic distances. We highlighted that isolation-by-environment shapes the genomic differentiation of B. prasinos whereas P. cordata is impacted by geographic distance (i.e., isolation-by-distance). At present time, the use of metagenomics to accurately estimate the genomic differentiation of protists remains challenging since coverages are lower compared to traditional population surveys. However, our approach sheds light on ecological and evolutionary processes occurring within natural marine populations, and paves the way for future protist population metagenomic studies.

The Mediterranean Sea is recognized as one of the most polluted areas by floating plastics. During the Tara Mediterranean expedition, an extensive sampling of plastic debris was conducted in seven ecoregions, from Gibraltar to Lebanon with the aim of providing reliable estimates of regional differences in floating plastic loads and plastic characteristics. The abundance, size, surface, circularity and mass of 75,030 pieces were analyzed and classified in a standardized multi-parameter database. Their average abundance was 2.60 × 105 items km−2 (2.25 × 103 to 8.50 × 106 km−2) resulting in an estimate of about 650 billion plastic particles floating on the surface of the Mediterranean. This corresponds to an average of 660 metric tons of plastic, at the lower end of literature estimates. High concentrations of plastic were observed in the northwestern coastal regions, north of the Tyrrhenian Sea, but also off the western and central Mediterranean basins. The Levantine basin south of Cyprus had the lowest concentrations. A Lagrangian Plastic Pollution Index (LPPI) predicting the concentration of plastic debris was validated using the spatial resolution of the data. The advanced state of plastic degradation detected in the analyses led to the conclusion that stranding/fragmentation/resuspension suggests it is a key process in the dynamics of floating plastic in Mediterranean surface waters. This is supported by the significant correlation between pollution sources and areas of high plastic concentration obtained by the LPPI.

The PRE-SWOT experiment was conducted onboard R/V García del Cid between 5 and 17 May 2018 in the southern region of the Balearic Islands (western Mediterranean Sea). PRE-SWOT aimed at anticipating the daily high-resolution 2D SSH fields that Surface Water & Ocean Topography (SWOT) will provide during the fast sampling phase after launch in selected areas of the global ocean. This experiment is a contribution to the preparatory SWOT cal/val activities and was coordinated with the PROTEUS-SWOT cruise (R/V Beautemps-Beauprè). The PRE-SWOT project (CTM2016-78607-P) is funded by the Spanish Research Agency and the European Regional Development Fund (AEI/FEDER, UE).

In the context of tracer transport in the ocean, we introduce a quantity, the crossroadness [1], which allows identifying the optimal disposition of a set of locations in order to monitor a given ocean surface region. The optimization is performed so that these sites observe the largest amount of water coming from the region and, at the same time, monitor waters coming from separate parts of the ocean. These are key criteria when deploying a marine observing network. Considering surface circulation, crossroadness measures at any location the extent of the ocean surface which transits in its neighborhood in a given time window. When the analysis is performed backward in time, this method allows us to identify the major sources which feed a target region. The method is first applied to a minimalistic model of a mesoscale eddy field, and then to realistic satellite-derived ocean currents in the Kerguelen area. In this region, we identify the optimal location of fixed stations capable of intercepting the trajectories of 43 surface drifters. We then illustrate the temporal persistence of the stations determined in this way. Finally, we identify possible hotspots of micro-nutrient enrichment for the recurrent spring phytoplanktonic bloom occurring there. Promising applications to other fields, such as larval connectivity or contaminant detection are discussed.

The study of connectivity patterns in networks has brought novel insights across diverse fields ranging from neurosciences to epidemic spreading or climate. In this context, betweenness centrality has demonstrated to be a very effective measure to identify nodes that act as focus of congestion, or bottlenecks, in the network. However, there is not a way to define betweenness outside the network framework. By analytically linking dynamical systems and network theory, we provide a trajectory-based formulation of betweenness, called Lagrangian betweenness, as a function of Lyapunov exponents. This extends the concept of betweenness beyond the context of network theory relating hyperbolic points and heteroclinic connections in any dynamical system to the structural bottlenecks of the network associated with it. Using modeled and observational velocity fields, we show that such bottlenecks are present and surprisingly persistent in the oceanic circulation across different spatio-temporal scales and we illustrate the role of these areas in driving fluid transport over vast oceanic regions. Analyzing plankton abundance data from the Kuroshio region of the Pacific Ocean, we find significant spatial correlations between measures of diversity and betweenness, suggesting promise for ecological applications.

The study of connectivity patterns in networks has brought novel insights across diverse fields ranging from neurosciences to epidemic spreading or climate. In this context, betweenness centrality has demonstrated to be a very effective measure to identify nodes that act as focus of congestion, or bottlenecks, in the network. However, there is not a way to define betweenness outside the network framework. Here we introduce the “Lagrangian betweenness”, an analogous quantity which relies only on the information provided by trajectories sampled across a generic dynamical system in the form of Finite Time Lyapunov Exponents, a widely used metric in Dynamical Systems Theory and Lagrangian oceanography. Our theoretical framework reveals a link between regions of high betweenness and the hyperbolic behavior of trajectories in the system. For example, it identifies bottlenecks in fluid flows where particles are first brought together and then widely dispersed. This has many potential applications including marine ecology and pollutant dispersal. We first test our definition of betweenness in an idealized double-gyre flow system. We then apply it in the characterization of transport by real geophysical flows in the semi-enclosed Adriatic Sea and the Kerguelen region of the highly turbulent Antarctic Circumpolar Current. In both cases, patterns of Lagrangian betweenness identify hidden bottlenecks of tracer transport that are surprisingly persistent across different spatio-temporal scales. In the marine context, high Lagrangian betweenness regions represent the optimal compromise between the heterogeneity of water origins and destinations, suggesting that they may be associated with relevant diversity reservoirs and hot-spots in marine ecosystems. Our new metric could also provide a novel approach useful for the management of environmental resources, informing strategies for marine spatial planning, and for designing observational networks to control pollutants or early-warn

The study of connectivity patterns in networks has brought novel insights across diverse fields ranging from neurosciences to epidemic spreading or climate. In this context, betweenness centrality has demonstrated to be a very effective measure to identify nodes that act as focus of congestion, or bottlenecks, in the network. However, there is not a way to define betweenness outside the network framework. By analytically linking dynamical systems and network theory, we provide a trajectory-based formulation of betweenness, called Lagrangian betweenness, as a function of Lyapunov exponents. This extends the concept of betweenness beyond the context of network theory relating hyperbolic points and heteroclinic connections in any dynamical system to the structural bottlenecks of the network associated with it. Using modeled and observational velocity fields, we show that such bottlenecks are present and surprisingly persistent in the oceanic circulation across different spatio-temporal scales and we illustrate the role of these areas in driving fluid transport over vast oceanic regions. Analyzing plankton abundance data from the Kuroshio region of the Pacific Ocean, we find significant spatial correlations between measures of diversity and betweenness, suggesting promise for ecological applications.

Connectivity is a fundamental structural feature of a network that determines the outcome of any dynamics that happens on top of it. However, an analytical approach to obtain connection probabilities between nodes associated with to paths of different lengths is still missing. Here, we derive exact expressions for random-walk connectivity probabilities across any range of numbers of steps in a generic temporal, directed, and weighted network. This allows characterizing explicit connectivity realized by causal paths as well as implicit connectivity related to motifs of three nodes and two links called here pitchforks. We directly link such probabilities to the processes of tagging and sampling any quantity exchanged across the network, hence providing a natural framework to assess transport dynamics. Finally, we apply our theoretical framework to study ocean transport features in the Mediterranean Sea. We find that relevant transport structures, such as fluid barriers and corridors, can generate contrasting and counterintuitive connectivity patterns bringing novel insights into how ocean currents drive seascape connectivity.

After the launch of the Surface Water and Ocean Topography (SWOT) satellite planned for 2022, the region around the Balearic Islands (western Mediterranean Sea) will be the target of several in situ sampling campaigns aimed at validating the first available tranche of SWOT data. In preparation for this validation, the PRE-SWOT cruise in 2018 was conceived to explore the three-dimensional (3D) circulation at scales of 20 km that SWOT aims to resolve, included in the fine-scale range (1–100 km) as defined by the altimetric community. These scales and associated variability are not captured by contemporary nadir altimeters. Temperature and salinity observations reveal a front that separates local Atlantic Water in the northeast from recent Atlantic Water in the southeast, and extends from the surface to ~150 m depth with maximum geostrophic velocities of the order of 0.20 m s−1 and a geostrophic Rossby number that ranges between −0.24 and 0.32. This front is associated with a 3D vertical velocity field characterized by an upwelling cell surrounded by two downwelling cells, one to the east and the other to the west. The upwelling cell is located near an area with high nitrate concentrations, possibly indicating a recent inflow of nutrients. Meanwhile, subduction of chlorophyll-a in the western downwelling cell is detected in glider observations. The comparison of the altimetric geostrophic velocity with the CTD-derived geostrophic velocity, the ADCP horizontal velocity, and drifter trajectories, shows that the present-day resolution of altimetric products precludes the representation of the currents that drive the drifter displacement. The Lagrangian analysis based on these velocities demonstrates that the study region has frontogenetic dynamics not detected by altimetry. Our results suggest that the horizontal component of the flow is mainly geostrophic down to scales of 20 km in the study region and during the period analyzed, and should therefore be resolvable by SWO

Habitat connectivity is central to several key evolutionary and ecological processes, having implications for the spatial structuring of marine populations. For benthic species that have no or little mobility as adults, connectivity is evaluated by analyzing the dispersal of propagules across the seascape. We investigate fine-scale connectivity of coralligenous habitats in Marseille Bay (located in the north-western Mediterranean) sea using high-resolution cartography and a particle-tracking model ran over a range of parameters derived from the biological traits of potential target species. We present annual and seasonal means of various connectivity diagnostics measuring the retention and exchange of propagules among coralligenous patches. A synthetic view is obtained by clustering individual patches into “coralligenous provinces” (ensemble of patches tightly connected by ocean currents). When discussing our results against historical observations, we highlight some genetic breaks, sharp community changes and ecological clusters that fit well our simulated connectivity patterns. Consistent findings include reduced biodiversity along Côte Bleue, high biodiversity at Planier Island and the presence of various dispersal barriers, which evolve with dispersal durations and provide the backbone of habitat connectivity. Our results help to apprehend and test hypothesis on marine population structures, providing useful information for ecologists and conservationists.

Understanding how climate change will affect oceanic fluid transport is crucial for environmental applications and human activities. However, a synoptic characterization of the influence of climate change on mesoscale stirring and transport in the surface ocean is missing. To bridge this gap, we exploit a high‐resolution, fully coupled climate model of the Mediterranean basin using a Network Theory approach. We project significant increases of horizontal stirring and kinetic energies in the next century, likely due to increments of available potential energy. The future evolution of basin‐scale transport patterns hints at a rearrangement of the main hydrodynamic provinces, defined as regions of the surface ocean that are well mixed internally but with minimal cross‐flow across their boundaries. This results in increased heterogeneity of province sizes and stronger mixing in their interiors. Our approach can be readily applied to other oceanic regions, providing information for the present and future marine spatial planning.

Quantifying the mechanisms of tracer dispersion in the ocean remains a central question in oceanography, for problems ranging from nutrient delivery to phytoplankton, to the early detection of contaminants. Until now, most of the analysis has been based on Lagrangian concepts of transport, often focusing on the identification of features that minimize fluid exchange among regions, or more recently, on network tools which focus instead on connectivity and transport pathways. Neither of these approaches, however, allows us to rank the geographical sites of major water passage, and at the same time, to select them so that they monitor waters coming from separate parts of the ocean. These are instead key criteria when deploying an observing network. Here, we address this issue by estimating at any point the extent of the ocean surface which transits through it in a given time window. With such information we are able to rank the sites with major fluxes that intercept waters originating from different regions. We show that this allows us to optimize an observing network, where a set of sampling sites can be chosen for monitoring the largest flux of water dispersing out of a given region. When the analysis is performed backward in time, this method allows us to identify the major sources which feed a target region. The method is first applied to a minimalistic model of a mesoscale eddy field, and then to realistic satellite-derived ocean currents in the Kerguelen area. In this region, we identify the optimal location of fixed stations capable of intercepting the trajectories of 43 surface drifters, along with statistics on the temporal persistence of the stations determined in this way. We then identify possible hotspots of micro-nutrient enrichment for the recurrent spring phytoplanktonic bloom occurring here. Promising applications to other fields, such as larval connectivity, marine spatial planning or contaminant detection, are then discussed.

Marine resources stewardships are progressively becoming more receptive to an effective incorporation of both ecosystem and environmental complexities into the analytical frameworks of fisheries assessment. Understanding and predicting marine fish production for spatially and demographically complex populations in changing environmental conditions is however still a difficult task. Indeed, fisheries assessment is mostly based on deterministic models that lack realistic parameterizations of the intricate biological and physical processes shaping recruitment, a cornerstone in population dynamics. We use here a large metapopulation of a harvested fish, the European hake (Merluccius merluccius), managed across transnational boundaries in the northwestern Mediterranean, to model fish recruitment dynamics in terms of physics‐dependent drivers related to dispersal and survival. The connectivity among nearby subpopulations is evaluated by simulating multi‐annual Lagrangian indices of larval retention, imports, and self‐recruitment. Along with a proxy of the regional hydroclimate influencing early life stages survival, we then statistically determine the relative contribution of dispersal and hydroclimate for recruitment across contiguous management units. We show that inter‐annual variability of recruitment is well reproduced by hydroclimatic influences and synthetic connectivity estimates. Self‐recruitment (i.e., the ratio of retained locally produced larvae to the total number of incoming larvae) is the most powerful metric as it integrates the roles of retained local recruits and immigrants from surrounding subpopulations and is able to capture circulation patterns affecting recruitment at the scale of management units. We also reveal that the climatic impact on recruitment is spatially structured at regional scale due to contrasting biophysical processes not related to dispersal. Self‐recruitment calculated for each management unit explains between 19% and 32.9% of the