Cross-basin and cross-taxa patterns of marine community tropicalization and deborealization in warming European seas.

Ocean warming and acidification, decreases in dissolved oxygen concentrations, and changes in primary production are causing an unprecedented global redistribution of marine life. The identification of underlying ecological processes underpinning marine species turnover, particularly the prevalence of increases of warm-water species or declines of cold-water species, has been recently debated in the context of ocean warming. Here, we track changes in the mean thermal affinity of marine communities across European seas by calculating the Community Temperature Index for 65 biodiversity time series collected over four decades and containing 1,817 species from different communities (zooplankton, coastal benthos, pelagic and demersal invertebrates and fish). We show that most communities and sites have clearly responded to ongoing ocean warming via abundance increases of warm-water species (tropicalization, 54%) and decreases of cold-water species (deborealization, 18%). Tropicalization dominated Atlantic sites compared to semi-enclosed basins such as the Mediterranean and Baltic Seas, probably due to physical barrier constraints to connectivity and species colonization. Semi-enclosed basins appeared to be particularly vulnerable to ocean warming, experiencing the fastest rates of warming and biodiversity loss through deborealization.

Seawater physics and chemistry along the Med-SHIP transects in the Mediterranean Sea in 2016.

The Mediterranean Sea has been sampled irregularly by research vessels in the past, mostly by national expeditions in regional waters. To monitor the hydrographic, biogeochemical and circulation changes in the Mediterranean Sea, a systematic repeat oceanographic survey programme called Med-SHIP was recommended by the Mediterranean Science Commission (CIESM) in 2011, as part of the Global Ocean Ship-based Hydrographic Investigations Program (GO-SHIP). Med-SHIP consists of zonal and meridional surveys with different frequencies, where comprehensive physical and biogeochemical properties are measured with the highest international standards. The first zonal survey was done in 2011 and repeated in 2018. In addition, a network of meridional (and other key) hydrographic sections were designed: the first cycle of these sections was completed in 2016, with three cruises funded by the EU project EUROFLEETS2. This paper presents the physical and chemical data of the meridional and key transects in the Western and Eastern Mediterranean Sea collected during those cruises.

Ecosystem-level responses to multiple stressors using a time-dynamic food-web model: The case of a re-oligotrophicated coastal embayment (Saronikos Gulf, E Mediterranean).

Multiple stressors may combine in unexpected ways to alter the structure of ecological systems, however, our current ability to evaluate their ecological impact is limited due to the lack of information concerning historic trophic interactions and ecosystem dynamics. Saronikos Gulf is a heavily exploited embayment in the E Mediterranean that has undergone significant ecological alterations during the last 20 years including a shift from long-standing eutrophic to oligotrophic conditions in the mid-2000's. Here we used a historical Ecopath food-web model of Saronikos Gulf (1998-2000) and fitted the time-dynamic module Ecosim to biomass and catch time series for the period 2001-2020. We then projected the model forward in time from 2021 to 2050 under 8 scenarios to simulate ecosystem responses to the individual and combined effect of sea surface temperature increase, primary productivity shifts and fishing effort release. Incorporating trophic interactions, climate warming, fishing and primary production improved model fit, depicting that both fishing and the environment have historically influenced ecosystem dynamics. Retrospective simulations of the model captured historical biomass and catch trends of commercially important stocks and reproduced successfully the marked recovery of marine resources 10 years after re-oligotrophication. In future scenarios increasing temperature had a detrimental impact on most functional groups, increasing and decreasing productivity had a positive and negative effect on all respectively, while fishing reductions principally benefited top predators. Combined stressors produced synergistic or antagonistic effects depending on the direction and magnitude of change of each stressor in isolation while their overall impact seemed to be strongly mediated via food-web interactions. Such holistic approaches advance of our mechanistic understanding of ecosystems enabling us to develop more effective management strategies in the face of a rapidly changing marine environment.

Metal and metallothionein concentrations in mesozooplankton from an oligotrophic offshore area in the eastern Mediterranean Sea (Cretan Passage/Levantine Sea).

Spatial variability of Cd, Cu, Cr, Ni, Zn, Fe, Mn, Pb and metallothionein (MT) concentrations were determined in mesozooplankton samples along the west-east axis of the Cretan Passage in the western Levantine Sea (Eastern Mediterranean). Metal and MT values from the present study are proposed as background levels, due to the lack of substantial anthropogenic activities in the area, where only maritime traffic and atmospheric deposition could be potential sources. Higher concentrations, of both metals and MTs, were recorded mainly at the western stations indicating higher metal bioavailability than in the eastern part. An inverse relationship of the metal zooplankton levels with zooplankton biomass, abundance and vital rates (production, respiration and ingestion rates), as well as salinity, was evident. We discuss the hypothesis that physical and biological characteristics of the marine environment, affecting growth dynamics of phytoplankton and zooplankton communities, may also act on metal uptake in oligotrophic marine systems.

The Influence of the Toxic Dinoflagellate Alexandrium minutum, Grown under Different N:P Ratios, on the Marine Copepod Acartia tonsa.

HABs pose a threat to coastal ecosystems, the economic sector and human health, and are expanding globally. However, their influence on copepods, a major connector between primary producers and upper trophic levels, remains essentially unknown. Microalgal toxins can eventually control copepod survival and reproduction by deterring grazing and hence reducing food availability. We present several 24-h experiments in which the globally distributed marine copepod, Acartia tonsa, was exposed to different concentrations of the toxic dinoflagellate, Alexandrium minutum, grown under three N:P ratios (4:1, 16:1 and 80:1), with the simultaneous presence of non-toxic food (the dinoflagellate Prorocentrum micans). The different N:P ratios did not affect the toxicity of A. minutum, probably due to the low toxicity of the tested strain. Production of eggs and pellets as well as ingested carbon appeared to be affected by food toxicity. Toxicity levels in A. minutum also had an effect on hatching success and on the toxin excreted in pellets. Overall, A. minutum toxicity affected the reproduction, toxin excretion and, to an extent, the feeding behavior of A. tonsa. This work indicates that even short-term exposure to toxic A. minutum can impact the vital functions of A. tonsa and might ultimately pose serious threats to copepod recruitment and survival. Still, further investigation is required for identifying and understanding, in particular, the long-term effects of harmful microalgae on marine copepods.

Metal and metallothionein levels in zooplankton in relation to environmental exposure: spatial and temporal variability (Saronikos Gulf, Greece).

Metal and metallothionein (MT) in mixed zooplankton were investigated as means of monitoring metal availability regarding environmental exposure. Spatial and temporal variability of Cd, Cu, Ni, Zn, Fe, Mn and Pb in zooplankton and seawater were studied in Saronikos Gulf (Aegean Sea, Eastern Mediterranean), once every second month during an annual cycle (2011-2012). Particulate organic carbon and chlorophyll α were also measured in seawater samples. Median zooplankton metal concentrations were 0.65, 32.4, 7.1, 864, 1420, 40.2 and 26.8 µg g-1 dw for Cd, Cu, Ni, Zn, Fe, Mn and Pb, respectively, and 109 µg g-1 ww for MTs. Metal levels in zooplankton and MTs were higher at sites influenced by human-derived pressures. Additionally, metal concentrations in pelagic fish flesh from the Greek MED-POL data base were used for bioconcentration and biomagnification factors calculation. Bioconcentration from water to zooplankton was higher than metal transfer from either seston to zooplankton or zooplankton to fish.

A light-induced shortcut in the planktonic microbial loop.

Mixotrophs combine photosynthesis with phagotrophy to cover their demands in energy and essential nutrients. This gives them a competitive advantage under oligotropihc conditions, where nutrients and bacteria concentrations are low. As the advantage for the mixotroph depends on light, the competition between mixo- and heterotrophic bacterivores should be regulated by light. To test this hypothesis, we incubated natural plankton from the ultra-oligotrophic Eastern Mediterranean in a set of mesocosms maintained at 4 light levels spanning a 10-fold light gradient. Picoplankton (heterotrophic bacteria (HB), pico-sized cyanobacteria, and small-sized flagellates) showed the fastest and most marked response to light, with pronounced predator-prey cycles, in the high-light treatments. Albeit cell specific activity of heterotrophic bacteria was constant across the light gradient, bacterial abundances exhibited an inverse relationship with light. This pattern was explained by light-induced top-down control of HB by bacterivorous phototrophic eukaryotes (PE), which was evidenced by a significant inverse relationship between HB net growth rate and PE abundances. Our results show that light mediates the impact of mixotrophic bacterivores. As mixo- and heterotrophs differ in the way they remineralize nutrients, these results have far-reaching implications for how nutrient cycling is affected by light.

Dispersal similarly shapes both population genetics and community patterns in the marine realm.

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 ß-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.