Comparing copepod time-series in the north of Spain: Spatial autocorrelation of community composition

Four time-series of copepod species biomass in the north of Spain were contrasted to demonstrate spatial autocorrelation of local communities and their responses to short-term local and regional variability in oceanographic conditions. The series represented coastal and oceanic environments along a marked gradient of influence of seasonal upwelling from Galicia to the Mar Cantábrico (S Bay of Biscay), and each one included at least 10 years of continuous data collected at monthly frequency. Community composition (i.e. species number and diversity) was very consistent through the region, but local variations in the presence of new species and the relative proportions of common species allowed for the characterisation of the response to the environment at each site. Small-sized species were more frequent near the coast. A few species, however, captured the main patterns of variability in all series. Calanus helgolandicus and Acartia (mainly Acartia clausi) were generally the main contributors to total biomass, while other species as Paracalanus parvus and Clausocalanus spp. were important only at some locations. Most copepod indices were positively correlated with upwelling, either considering the whole community (biomass, species richness and diversity) or individual species, but only in the coastal series analysed since 1991. Copepods in the nearby ocean, however, showed negative correlations with upwelling in the period 1960–1986. The effects of upwelling may have been modulated by local factors, as showed by the increases in biomass, number of species and diversity in associations with increases in sea surface temperature in Galicia, while in the Mar Cantábrico only the warming-tolerant species increased and those typical of upwelling decreased. Density stratification of the water column was associated with decreases in total copepod biomass in Galicia, while it favoured the increase in species richness in the Mar Cantábrico. Nearly all significant responses of copepods to environmental variability were delayed by up to 5 months, showing the importance of considering time-lags in the analysis of temporal responses of zooplankton.

The influence of phytoplankton productivity, temperature and environmental stability on the control of copepod diversity in the North East Atlantic

The patterns of copepod species richness (S) and their relationship with phytoplankton productivity, temperature and environmental stability were investigated at climatological, seasonal and year-to-year time scales as well as scales along latitudinal and oceanic–neritic gradients using monthly time series of the Continuous Plankton Recorder (CPR) Survey collected in the North East Atlantic between 1958 and 2006. Time series analyses confirmed previously described geographic patterns. Equatorward and towards neritic environments, the climatological average of S increases and the variance explained by the seasonal cycle decreases. The bi-modal character of seasonality increases equatorward and the timing of the seasonal cycle takes place progressive earlier equatorward and towards neritic environments. In the long-term, the climatological average of S decreased significantly (p < 0.001) between 1958 and 2006 in the Bay of Biscay and North Iberian shelf at a rate of ca. 0.04 year−1, and increased at the same rate between 1991 and 2006 in the northernmost oceanic location. The climatological averages of S correlate positively with those of the index of seasonality of phytoplankton productivity (ratio between the minimum and maximum monthly values of surface chlorophyll) and sea surface temperature, and negatively with those of the proxy for environmental stability (monthly frequency of occurrence of daily averaged wind speed exceeding 10 m s−1). The seasonal cycles of S and phytoplankton productivity (surface chlorophyll as proxy) exhibit similar features in terms of shape, timing and explained variance, but the relationship between the climatological averages of both variables is non-significant. From year-to-year, the annual averages of S correlate negatively with those of phytoplankton productivity and positively with those of sea surface temperature along the latitudinal gradient, and negatively with those of environmental stability along the oceanic–neritic gradient. The annual anomalies of S (i.e. factoring out geographic variation) show a unimodal relationship with those of sea surface temperature and environmental stability, with S peaking at intermediate values of the anomalies of these variables. The results evidence the role of seasonality of phytoplankton productivity on the control of copepod species richness at seasonal and climatological scales, giving support to the species richness–productivity hypothesis. Although sea surface temperature (SST) is indeed a good predictor of richness along the latitudinal gradient, it is unable to predict the increase of richness form oceanic to neritic environments, thus lessening the generality of the species richness–energy hypothesis. Meteo-hydrographic disturbances (i.e. SST and wind speed anomalies as proxies), presumably through its role on mixed layer depth dynamics and turbulence and hence productivity, maximise local diversity when occurring at intermediate frequency and or intensity, thus providing support to the intermediate disturbance hypothesis on the control of copepod diversity.

Global latitudinal variations in marine copepod diversity and environmental factors

Latitudinal gradients in diversity are among the most striking features in ecology. For terrestrial species, climate (i.e. temperature and precipitation) is believed to exert a strong influence on the geographical distributions of diversity through its effects on energy availability. Here, we provide the first global description of geographical variation in the diversity of marine copepods, a key trophic link between phytoplankton and fish, in relation to environmental variables. We found a polar-tropical difference in copepod diversity in the Northern Hemisphere where diversity peaked at subtropical latitudes. In the Southern Hemisphere, diversity showed a tropical plateau into the temperate regions. This asymmetry around the Equator may be explained by climatic conditions, in particular the influence of the Inter-Tropical Convergence Zone, prevailing mainly in the northern tropical region. Ocean temperature was the most important explanatory factor among all environmental variables tested, accounting for 54 per cent of the variation in diversity. Given the strong positive correlation between diversity and temperature, local copepod diversity, especially in extra-tropical regions, is likely to increase with climate change as their large-scale distributions respond to climate warming.

Nematode diversity in different microhabitats in a mangrove region

Mangroves are highly productive environments that play important ecological and socioeconomic roles; however, they have been impacted to different degrees in most countries worldwide. The knowledge of which organisms inhabit this environment and their ecological interactions is the first step towards its conservation. The natural variability of environmental factors in mangroves provides numerous niches available to different species. Meiofauna have patchy patterns of distribution that are related to the availability of resources. Hence, meiofauna are expected to present a high diversity of different taxa occupying the different microhabitats offered by mangroves. This work aims to test the hypothesis that the assemblage structure of Nematoda varies significantly among mangrove microhabitats and to contribute knowledge on the meiofauna diversity in mangrove environments. This work was carried out in a mangrove region at Pernambuco state, Northeastern Brazil. Qualitative samples were collected in nine microhabitats which show different characteristics mainly in terms of presence of vegetation or another organism and sediment grain size. Univariate and multivariate analysis were applied to Nematoda genera abundance data. Our results demonstrate the existence of significant differences among microhabitats regarding nematode assemblage structure corroborating the hypothesis. Different Nematoda assemblages are present in at least seven microhabitats. These assemblages are composed of nematode genera with different trophic and morphological features, demonstrating a strong relationship between morphological diversity and ecological plasticity. Furthermore, this study also demonstrates the importance of the conservation of this ecosystem and its attributes.

Temporal and spatial variation in the Nazaré Canyon (Western Iberian margin): Inter-annual and canyon heterogeneity effects on meiofauna biomass and diversity

The Nazaré Canyon on the Portuguese Margin (NE Atlantic) was sampled during spring-summer for three consecutive years (2005–2007), permitting the first inter-annual study of the meiofaunal communities at the Iberian Margin at two abyssal depths (~3500 m and ~4400 m). Using new and already published data, the meiofauna standing stocks (abundance and biomass) and nematode structural and functional diversity were investigated in relation to the sediment biogeochemistry (e.g. organic carbon, nitrogen, chlorophyll a, phaeopigments) and grain size. A conspicuous increase in sand content from 2005 to 2006 and decrease of phytodetritus at both sites, suggested the occurrence of one or more physical disturbance events. Nematode standing stocks and trophic diversity decreased after these events, seemingly followed by a recovery/recolonisation period in 2007, which was strongly correlated with an increase in the quantity and bioavailability of phytodetrital organic matter supplied. Changes in meiofauna assemblages, however, also differed between stations, likely because of the contrasting hydrodynamic and food supply conditions. Higher meiofauna and nematode abundances, biomass and trophic complexity were found at the shallowest canyon station, where the quantity, quality and bioavailability of food material were higher than at the deeper site. The present results suggest that even though inter-annual variations in the sedimentary environment can regulate the meiofauna in the abyssal Nazaré Canyon, heterogeneity between sampling locations in the canyon were more pronounced.

Frequent locations of oceanic fronts as an indicator of pelagic diversity: application to marine protected areas and renewables

Frequent locations of thermal fronts in UK shelf seas were identified using an archive of 30,000 satellite images acquired between 1999 and 2008, and applied as a proxy for pelagic diversity in the designation of Marine Protected Areas (MPAs). Networks of MPAs are required for conservation of critical marine habitats within Europe, and there are similar initiatives worldwide. Many pelagic biodiversity hotspots are related to fronts, for example cetaceans and basking sharks around the Isle of Man, Hebrides and Cornwall, and hence remote sensing can address this policy need in regions with insufficient species distribution data. This is the first study of UK Continental Shelf front locations to use a 10-year archive of full-resolution (1.1 km) AVHRR data, revealing new aspects of their spatial and seasonal variability. Frontal locations determined at sea or predicted by ocean models agreed closely with the new frequent front maps, which also identified many additional frontal zones. These front maps were among the most widely used datasets in the recommendation of UK MPAs, and would be applicable to other geographic regions and to other policy drivers such as facilitating the deployment of offshore renewable energy devices with minimal environmental impact.

Late winter under ice pelagic microbial communities in the high Arctic Ocean and the impact of short-term exposure to elevated CO2 levels

Polar Oceans are natural CO2 sinks because of the enhanced solubility of CO2 in cold water. The Arctic Ocean is at additional risk of accelerated ocean acidification (OA) because of freshwater inputs from sea ice and rivers, which influence the carbonate system. Winter conditions in the Arctic are of interest because of both cold temperatures and limited CO2 venting to the atmosphere when sea ice is present. Earlier OA experiments on Arctic microbial communities conducted in the absence of ice cover, hinted at shifts in taxa dominance and diversity under lowered pH. The Catlin Arctic Survey provided an opportunity to conduct in situ, under-ice, OA experiments during late Arctic winter. Seawater was collected from under the sea ice off Ellef Ringnes Island, and communities were exposed to three CO2 levels for 6 days. Phylogenetic diversity was greater in the attached fraction compared to the free-living fraction in situ, in the controls and in the treatments. The dominant taxa in all cases were Gammaproteobacteria but acidification had little effect compared to the effects of containment. Phylogenetic net relatedness indices suggested that acidification may have decreased the diversity within some bacterial orders, but overall there was no clear trend. Within the experimental communities, alkalinity best explained the variance among samples and replicates, suggesting subtle changes in the carbonate system need to be considered in such experiments. We conclude that under ice communities have the capacity to respond either by selection or phenotypic plasticity to heightened CO2 levels over the short term.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

Phytoplankton phenology indices in coral reef ecosystems: Application to ocean-color observations in the Red Sea

Phytoplankton, at the base of the marine food web, represent a fundamental food source in coral reef ecosystems. The timing (phenology) and magnitude of the phytoplankton biomass are major determinants of trophic interactions. The Red Sea is one of the warmest and most saline basins in the world, characterized by an arid tropical climate regulated by the monsoon. These extreme conditions are particularly challenging for marine life. Phytoplankton phenological indices provide objective and quantitative metrics to characterize phytoplank- ton seasonality. The indices i.e. timings of initiation, peak, termination and duration are estimated here using 15 years (1997–2012) of remote sensing ocean-color data from the European Space Agency (ESA) Climate Change Initiative project (OC-CCI) in the entire Red Sea basin. The OC-CCI product, comprising merged and bias-corrected observations from three independent ocean-color sensors (SeaWiFS, MODIS and MERIS), and processed using the POLYMER algorithm (MERIS period), shows a significant increase in chlorophyll data cover- age, especially in the southern Red Sea during the months of summer NW monsoon. In open and reef-bound coastal waters, the performance of OC-CCI chlorophyll data is shown to be comparable with the performance of other standard chlorophyll products for the global oceans. These features have permitted us to investigate phytoplankton phenology in the entire Red Sea basin, and during both winter SE monsoon and summer NW monsoon periods. The phenological indices are estimated in the four open water provinces of the basin, and further examined at six coral reef complexes of particular socio-economic importance in the Red Sea, including Siyal Islands, Sharm El Sheikh, Al Wajh bank, Thuwal reefs, Al Lith reefs and Farasan Islands. Most of the open and deeper waters of the basin show an apparent higher chlorophyll concentration and longer duration of phyto- plankton growth during the winter period (relative to the summer phytoplankton growth period). In contrast, most of the reef-bound coastal waters display equal or higher peak chlorophyll concentrations and equal or lon- ger duration of phytoplankton growth during the summer period (relative to the winter phytoplankton growth period). The ecological and biological significance of the phytoplankton seasonal characteristics are discussed in context of ecosystem state assessment, and particularly to support further understanding of the structure and functioning of coral reef ecosystems in the Red Sea.

Impact of sub-seabed CO2 leakage on macrobenthic community structure and diversity

A sub-seabed release of carbon dioxide (CO2) was conducted to assess the potential impacts of leakage from sub-seabed geological CO2 Capture and Storage CCS) on benthic macrofauna. CO2 gas was released 12 m below the seabed for 37 days, causing significant disruption to sediment carbonate chemistry. Regular macrofauna samples were collected from within the area of active CO2 leakage (Zone 1) and in three additional reference areas, 25 m, 75 m and 450 m from the centre of the leakage (Zones 2, 3 and 4 respectively). Macrofaunal community structure changed significantly in all zones during the study period. However, only the changes in Zone 1 were driven by the CO2 leakage with the changes in reference zones appearing to reflect natural seasonal succession and stochastic weather events. The impacts in Zone 1 occurred rapidly (within a few days), increased in severity through the duration of the leak, and continued to worsen after the leak had stopped. Considerable macrofaunal recovery was seen 18 days after the CO2 gas injection had stopped. In summary, small short-term CCS leakage events are likely to cause highly localised impacts on macrofaunal communities and there is the potential for rapid recovery to occur, depending on the characteristics of the communities and habitats impacted.