Role of light and photophysiological properties on phytoplankton succession during the spring bloom in the north-western Mediterranean Sea

This study aimed to determine the role of light on the succession of the phytoplankton community during the spring bloom in the northwestern Mediterranean Sea. To this end, three successive Lagrangian experiments were carried out between March and April 2003. The three experiments correspond to distinct phases of the bloom development (pre-bloom, bloom peak and post-bloom, respectively) and therefore to different trophic conditions. Phytoplankton (sampled on a daily scale) was grouped in size-based classes (pico and nano+micro) each of them were characterised in terms of chemotaxonomic composition, primary production and photophysiological properties. The phytoplankton community evolved with time changing in both size-class dominance and specie/group dominance within each size class. The bloom peak was characterised by highly dynamic condition (i.e. vertical mixing) and by the dominance of both small (pico) and large (nano and micro) diatoms, as a result of their capacity to photoacclimate to changing light regimes (‘physiological plasticity’). Concluding, we suggest that the physiological adaptation to light is the main factor driving the succession of the phytoplankton community during the first phases of the bloom (until the onset of thermal stratification) in the western Mediterranean Sea.

Basin scale distribution of zooplankton in the Ligurian Sea (north-western Mediterranean) in late autumn

Mesozooplankton biomass and abundance were evaluated in epipelagic waters at 59 stations covering the Italian sector of the Ligurian Sea (north-western Mediterranean) in December 1990. This region is characterised by a cyclonic circulation which encloses a central divergence zone and is associated with a main thermohaline front offshore the western Ligurian coast. At the end of autumn, mesozooplankton biomass (range: 0.80–4.24 mg DW m−3) and the abundance (range: 83.8–932 ind. m−3) were lower in the divergence zone. On the contrary, in the Ligurian frontal zone at the periphery of the divergence and on the eastern continental shelf the greatest values of biomass and abundance were recorded. Copepods and appendicularians dominated the mesozooplankton community, the main taxa being the copepods Clausocalanus spp. (46% of total zooplankton) and Oithona spp. (15%) and the appendicularian Fritillaria spp. (12%). Three hydrological sub-regions, i.e. the divergence, the eastern continental shelf and the periphery of the divergence, were characterised by different zooplankton communities and characteristic species. Environmental differences between the three zones were mainly related to changes in bottom topography, sea surface temperatures and quantity of particulate organic matter. Vertical mesozooplankton abundance and taxa distribution from the surface to 1,900 m depth were also examined in one station. The results showed that the bulk of the community was concentrated in the upper 200 m, small copepods being dominant particularly in the upper 50 m. The copepod community was more diversified in sub-superficial waters, with a maximum observed in the 200–400 m layer. The distributions of main zooplankton taxa described in epipelagic waters in the eastern Ligurian Sea in autumn were compared with their distribution at surface in the north-western Mediterranean obtained by sampling performed with the Continuous Plankton Recorder in 1997–1999. The analysis of the zooplankton community in CPR samples confirms the dominance of small copepods (Paracalanus spp., Clausocalanus spp., Oithona spp.) and appendicularians in the north-western Mediterranean in late autumn-winter and shows that their distribution is mainly related to the main mesoscale hydrographic features characterising this basin.

Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean)

Long-term variability of the main calycophoran siphonophores was investigated between 1974 and 1999 in a coastal station in the north-western Mediterranean. The data were collected at weekly frequency using a macroplankton net (680 μm mesh size) adapted to quantitatively sample delicate gelatinous plankton. A 3-year collection (1967–1969) of siphonophores from offshore waters using the same methodology showed that the patterns of variability observed inshore were representative of siphonophores’ changes at a regional scale. The aims of the study were: (i) to investigate the patterns of variability that characterised the dominant calycophoran species and assemblages; (ii) to identify the environmental optima that were associated with a significant increase in the dominant siphonophore species and (iii) to verify the influence of hydroclimatic variability on long-term changes of siphonophores. Our results showed that during nearly 3 decades the standing stock of calycophoran siphonophores did not show any significant change, with the annual maximum usually recorded in spring as a result of high densities of the dominant species Lensia subtilis, Muggiaea kochi and Muggiaea atlantica. Nevertheless, major changes in community composition occurred within the calycophoran population. Since the middle 1980s, M. kochi, once the most dominant species, started to decrease allowing other species, the congeneric M. atlantica and Chelophyes appendiculata, to increasingly dominate in spring and summer–autumn, respectively. The comparison of environmental and biotic long-term trends suggests that the decrease of M. kochi was triggered by hydrological changes that occurred in the north-western Mediterranean under the forcing of large-scale climate oscillations. Salinity, water stratification and water temperature were the main hydroclimatic factors associated with a significant increase of siphonophores, different species showing different environmental preferences.

Ocean acidification impacts on nitrogen fixation in the coastal western Mediterranean Sea

The effects of ocean acidification on nitrogen (N2) fixation rates and on the community composition of N2-fixing microbes (diazotrophs) were examined in coastal waters of the North-Western Mediterranean Sea. Nine experimental mesocosm enclosures of ∼50 m3 each were deployed for 20 days during June-July 2012 in the Bay of Calvi, Corsica, France. Three control mesocosms were maintained under ambient conditions of carbonate chemistry. The remainder were manipulated with CO2 saturated seawater to attain target amendments of pCO2 of 550, 650, 750, 850, 1000 and 1250 μatm. Rates of N2 fixation were elevated up to 10 times relative to control rates (2.00 ± 1.21 nmol L-1d-1) when pCO2 concentrations were >1000 μatm and pHT (total scale) < 7.74. Diazotrophic phylotypes commonly found in oligotrophic marine waters, including the Mediterranean, were not present at the onset of the experiment and therefore, the diazotroph community composition was characterised by amplifying partial nifH genes from the mesocosms. The diazotroph community was comprised primarily of cluster III nifH sequences (which include possible anaerobes), and proteobacterial (α and γ) sequences, in addition to small numbers of filamentous (or pseudo-filamentous) cyanobacterial phylotypes. The implication from this study is that there is some potential for elevated N2 fixation rates in the coastal western Mediterranean before the end of this century as a result of increasing ocean acidification. Observations made of variability in the diazotroph community composition could not be correlated with changes in carbon chemistry, which highlights the complexity of the relationship between ocean acidification and these keystone organisms.

Zooplankton associations in a Mediterranean long-term time-series

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.

Predicting future thermal habitat suitability of competing native and invasive fish species: from metabolic scope to oceanographic modelling

Global increase in sea temperatures has been suggested to facilitate the incoming and spread of tropical invaders. The increasing success of these species may be related to their higher physiological performance compared with indigenous ones. Here, we determined the effect of temperature on the aerobic metabolic scope (MS) of two herbivorous fish species that occupy a similar ecological niche in the Mediterranean Sea: the native salema (Sarpa salpa) and the invasive marbled spinefoot (Siganus rivulatus). Our results demonstrate a large difference in the optimal temperature for aerobic scope between the salema (21.8°C) and the marbled spinefoot (29.1°C), highlighting the importance of temperature in determining the energy availability and, potentially, the distribution patterns of the two species. A modelling approach based on a present-day projection and a future scenario for oceanographic conditions was used to make predictions about the thermal habitat suitability (THS, an index based on the relationship between MS and temperature) of the two species, both at the basin level (the whole Mediterranean Sea) and at the regional level (the Sicilian Channel, a key area for the inflow of invasive species from the Eastern to the Western Mediterranean Sea). For the present-day projection, our basin-scale model shows higher THS of the marbled spinefoot than the salema in the Eastern compared with the Western Mediterranean Sea. However, by 2050, the THS of the marbled spinefoot is predicted to increase throughout the whole Mediterranean Sea, causing its westward expansion. Nevertheless, the regional-scale model suggests that the future thermal conditions of Western Sicily will remain relatively unsuitable for the invasive species and could act as a barrier for its spread westward. We suggest that metabolic scope can be used as a tool to evaluate the potential invasiveness of alien species and the resilience to global warming of native species.

Influence of the Pacific Decadal Oscillation on phytoplankton phenology and community structure in the western North Pacific

Phytoplankton phenology and community structure in the western North Pacific were investigated for 2001–2009, based on satellite ocean colour data and the Continuous Plankton Recorder survey. We estimated the timing of the spring bloom based on the cumulative sum satellite chlorophyll adata, and found that the Pacific Decadal Oscillation (PDO)-related interannual SST anomaly in spring significantly affected phytoplankton phenology. The bloom occurred either later or earlier in years of positive or negative PDO (indicating cold and warm conditions, respectively). Phytoplankton composition in the early summer varied depending on the magnitude of seasonal SST increases, rather than the SST value itself. Interannual variations in diatom abundance and the relative abundance of non-diatoms were positively correlated with SST increases for March–April and May–July, respectively, suggesting that mixed layer environmental factors, such as light availability and nutrient stoichiometry, determine shifts in phytoplankton community structure. Our study emphasised the importance of the interannual variation in climate-induced warm–cool cycles as one of the key mechanisms linking climatic forcing and lower trophic level ecosystems.

Long-Term Oceanographic and Ecological Research in the Western English Channel

Long-term research in the western English Channel, undertaken by the marine laboratories in Plymouth, is described and details of survey methods, sites, and time series given in this chapter. Major findings are summarized and their limitations outlined. Current research, with recent reestablishment and expansion of many sampling programmes, is presented, and possible future approaches are indicated. These unique long-term data sets provide an environmental baseline for predicting complex ecological responses to local, regional, and global environmental change. Between 1888 and the present, investigations have been carried out into the physical, chemical, and biological components (ranging from plankton and fish to benthic and intertidal assemblages) of the western English Channel ecosystem. The Marine Biological Association of the United Kingdom has performed the main body of these observations. More recent contributions come from the Continuous Plankton Recorder Survey, now the Sir Alister Hardy Foundation for Ocean Science, dating from 1957; the Institute for Marine Environmental Research, from 1974 to 1987; and the Plymouth Marine Laboratory, which was formed by amalgamation of the Institute for Marine Environmental Research and part of the Marine Biological Association, from 1988. Together, these contributions constitute a unique data series; one of the longest and most comprehensive samplings of environmental and marine biological variables in the world. Since the termination of many of these time series in 1987-1988 during a reorganisation of UK marine research, there has been a resurgence of interest in long-term environmental change. Many programmes have been restarted and expanded with support from several agencies. The observations span significant periods of warming (1921-1961; 1985-present) and cooling (1962-1980). During these periods of change, the abundance of key species underwent dramatic shifts. The first period of warming saw changes in zooplankton, pelagic fish, and larval fish, including the collapse of an important herring fishery. During later periods of change, shifts in species abundances have been reflected in other assemblages, such as the intertidal zone and the benthic fauna. Many of these changes appear to be related to climate, manifested as temperature changes, acting directly or indirectly. The hypothesis that climate is a forcing factor is widely supported today and has been reinforced by recent studies that show responses of marine organisms to climatic attributes such as the strength of the North Atlantic Oscillation. The long-term data also yield important insights into the effects of anthropogenic disturbances such as fisheries exploitation and pollution. Comparison of demersal fish hauls over time highlights fisheries effects not only on commercially important species but also on the entire demersal community. The effects of acute ("Torrey Canyon" oil spill) and chronic (tributyltin [TBT] antifoulants) pollution are clearly seen in the intertidal records. Significant advances in diverse scientific disciplines have been generated from research undertaken alongside the long-term data series.

Variations in the distributions of the Centropages chierchiae and Temora stylifera (Copepoda: Calanoida) in the north-eastern Atlantic Ocean and western european shelf waters.

Centropages chierchiae and Temora stylifera occurred rarely in the Continuous Plankton Recorder (CPR) survey in the Bay of Biscay, Celtic Sea, and English Channel before 1988. By 2000 they were found frequently and in abundance. The seasonal cycles of abundance of these species differ, C. chierchiae occurring mainly in the summer while T. stylifera was found most frequently in late autumn or winter towards the northern limits of its distribution. The increase in abundance of both species is related to temperature. However, in the years when it was found in the samples, the frequency of occurrence of C. chierchiae was correlated positively with the strength of the shelf edge current and negatively with the North Atlantic Oscillation (NAO) while the reverse was true for T. stylifera.

The Mediterranean Sea Regime Shift at the End of the 1980s, and Intriguing Parallelisms with Other European Basins

Regime shifts are abrupt changes encompassing a multitude of physical properties and ecosystem variables, which lead to new regime conditions. Recent investigations focus on the changes in ecosystem diversity and functioning associated to such shifts. Of particular interest, because of the implication on climate drivers, are shifts that occur synchronously in separated basins. In this work we analyze and review long-term records of Mediterranean ecological and hydro-climate variables and find that all point to a synchronous change in the late 1980s. A quantitative synthesis of the literature (including observed oceanic data, models and satellite analyses) shows that these years mark a major change in Mediterranean hydrographic properties, surface circulation, and deep water convection (the Eastern Mediterranean Transient). We provide novel analyses that link local, regional and basin scale hydrological properties with two major indicators of large scale climate, the North Atlantic Oscillation index and the Northern Hemisphere Temperature index, suggesting that the Mediterranean shift is part of a large scale change in the Northern Hemisphere. We provide a simplified scheme of the different effects of climate vs. temperature on pelagic ecosystems.

Mediterranean marine copepods: basin-scale trends of the calanoid Centropages typicus

The Mediterranean Sea is located in a crossroad of mid-latitude and subtropical climatic modes that enhance contrasting environmental conditions over both latitudinal and longitudinal ranges. Here, we show that the large-scale environmental forcing is reflected in the basin scale trends of the adult population of the calanoid copepod Centropages typicus. The species is distributed over the whole Mediterranean basin, and maximal abundances were found in the north-western basin associated to oceanic fronts, and in the Adriatic Sea associated to shallow and semi enclosed waters. The peak of main abundances of C. typicus correlates with the latitudinal temperature gradient and the highest seasonal abundances occurred in spring within the 14–18°C temperature window. Such thermal cline may define the latitudinal geographic region where C. typicus seasonally dominates the >200 μm-sized spring copepod community in the Mediterranean Sea. The approach used here is generally applicable to investigate the large-scale spatial patterns of other planktonic organisms and to identify favourable environmental windows for population development.

Satellite estimates of net community production indicate predominance of net autotrophy in the Atlantic Ocean

There is ongoing debate as to whether the oligotrophic ocean is predominantly net autotrophic and acts as a CO2 sink, or net heterotrophic and therefore acts as a CO2 source to the atmosphere. This quantification is challenging, both spatially and temporally, due to the sparseness of measurements. There has been a concerted effort to derive accurate estimates of phytoplankton photosynthesis and primary production from satellite data to fill these gaps; however there have been few satellite estimates of net community production (NCP). In this paper, we compare a number of empirical approaches to estimate NCP from satellite data with in vitro measurements of changes in dissolved O2 concentration at 295 stations in the N and S Atlantic Ocean (including the Antarctic), Greenland and Mediterranean Seas. Algorithms based on power laws between NCP and particulate organic carbon production (POC) derived from 14C uptake tend to overestimate NCP at negative values and underestimate at positive values. An algorithm that includes sea surface temperature (SST) in the power function of NCP and 14C POC has the lowest bias and root-mean square error compared with in vitro measured NCP and is the most accurate algorithm for the Atlantic Ocean. Nearly a 13 year time series of NCP was generated using this algorithm with SeaWiFS data to assess changes over time in different regions and in relation to climate variability. The North Atlantic subtropical and tropical Gyres (NATL) were predominantly net autotrophic from 1998 to 2010 except for boreal autumn/winter, suggesting that the northern hemisphere has remained a net sink for CO2 during this period. The South Atlantic subtropical Gyre (SATL) fluctuated from being net autotrophic in austral spring-summer, to net heterotrophic in austral autumn–winter. Recent decadal trends suggest that the SATL is becoming more of a CO2 source. Over the Atlantic basin, the percentage of satellite pixels with negative NCP was 27%, with the largest contributions from the NATL and SATL during boreal and austral autumn–winter, respectively. Variations in NCP in the northern and southern hemispheres were correlated with climate indices. Negative correlations between NCP and the multivariate ENSO index (MEI) occurred in the SATL, which explained up to 60% of the variability in NCP. Similarly there was a negative correlation between NCP and the North Atlantic Oscillation (NAO) in the Southern Sub-Tropical Convergence Zone (SSTC),which explained 90% of the variability. There were also positive correlations with NAO in the Canary Current Coastal Upwelling (CNRY) and Western Tropical Atlantic (WTRA)which explained 80% and 60% of the variability in each province, respectively. MEI and NAO seem to play a role in modifying phases of net autotrophy and heterotrophy in the Atlantic Ocean.