Defining seasonal marine microbial community dynamics

Here we describe, the longest microbial time-series analyzed to date using high-resolution 16S rRNA tag pyrosequencing of samples taken monthly over 6 years at a temperate marine coastal site off Plymouth, UK. Data treatment effected the estimation of community richness over a 6-year period, whereby 8794 operational taxonomic units (OTUs) were identified using single-linkage preclustering and 21 130 OTUs were identified by denoising the data. The Alphaproteobacteria were the most abundant Class, and the most frequently recorded OTUs were members of the Rickettsiales (SAR 11) and Rhodobacteriales. This near-surface ocean bacterial community showed strong repeatable seasonal patterns, which were defined by winter peaks in diversity across all years. Environmental variables explained far more variation in seasonally predictable bacteria than did data on protists or metazoan biomass. Change in day length alone explains >65% of the variance in community diversity. The results suggested that seasonal changes in environmental variables are more important than trophic interactions. Interestingly, microbial association network analysis showed that correlations in abundance were stronger within bacterial taxa rather than between bacteria and eukaryotes, or between bacteria and environmental variables.

How well does the Continuous Plankton Recorder (CPR) sample zooplankton? A comparison with the Longhurst Hardy Plankton Recorder (LHPR) in the northeast Atlantic

The Continuous Plankton Recorder (CPR) survey has collected data on basin- scale zooplankton abundance in the North Atlantic since the 1930s. These data have been used in many studies to elucidate seasonal patterns and long-term change in plankton populations, as well as more recently to validate ecosystem models. There has, however, been relatively little comparison of the data from the CPR with that from other samplers. In this study we compare zooplankton abundance estimated from the CPR in the northeast Atlantic with near-surface samples collected by a Longhurst-Hardy Plankton Recorder (LHPR) at Ocean Weather Station India (59 degree N, 19 degree W) between 1971 and 1975. Comparisons were made for six common copepods in the region: Acartia clausi, Calanus finmarchicus, Euchaeta norvegica, Metridia lucens, Oithona sp. and Pleuromamma robusta. Seasonal cycles based on CPR data were similar to those recorded by the LHPR. Differences in absolute abundances were apparent, however, with the CPR underestimating abundances by a factor of between 5 and 40, with the exception of A. clausi. Active avoidance by zooplankton is thought to be responsible. This avoidance is species specific, so that care must be taken describing communities, as the CPR emphasises those species that are preferentially caught, a problem common to many plankton samplers.

Temporal changes in plankton of the North Sea: community shifts and environmental drivers

This paper analyses long-term and seasonal changes in the North Sea plankton community during the period 1970 to 2008. Based on Continuous Plankton Recorder (CPR) data covering 38 yr, major changes in both phytoplankton and zooplankton abundance and community structure were identified. Regime changes were detected around 1978, 1989 and 1998. The first 2 changes have been discussed in the literature and are defined as a cold episodic event (1978) and a regime shift towards a warm dynamic regime (1989). The effect of these 2 regime changes on plankton indicators was assessed and checked against previous studies. The 1998 change represents a shift in the abundance and seasonal patterns of dinoflagellates and the dominant zooplankton group, the neritic copepods. Furthermore, environmental factors such as air temperature, wind speed and the North Atlantic water inflow were identified as potential drivers of change in seasonal patterns, and the most-likely environmental causes for detected changes were assessed. We suggest that a change in the balance of dissolved nutrients driven by these environmental factors was the cause of the latest change in plankton community structure, which in turn could have affected the North Sea fish community.

Modelling a light-driven phytoplankton succession

We used a numerical model to investigate if and to what extent cellular photoprotective capacity accounts for succession and vertical distribution of marine phytoplankton species/groups. A model describing xanthophyll photoprotective activity in phytoplankton has been implemented in the European Regional Sea Ecosystem Model and applied at the station L4 in the Western English Channel. Primary producers were subdivided into three phytoplankton functional types defined in terms of their capacity to acclimate to different light-specific environments: low light (LL-type), high light (HL-type) and variable light (VL-type) adapted species. The LL-type is assumed to have low cellular level of xanthophyll-cycling pigments (PX) relative to the modelled photosynthetically active pigments (chlorophyll and fucoxanthin (FUCO) = PSP). The HL-type has high PX content relative to PSP while VL-type presents an intermediate PX to PSP ratio. Furthermore, the VL-type is capable of reversibly converting FUCO to PX and synthesizing new PX under high-light stress. In order to reproduce phytoplankton community succession with each of the three groups being dominant in different periods of the year, we had also to assume reduced grazing pressure on HL-adapted species. Model simulations realistically reproduce the observed seasonal patterns of pigments and nutrients highlighting the reasonability of the underpinning assumptions. Our model suggests that pigment-mediated photophysiology plays a primary role in determining the evolution of marine phytoplankton communities in the winter-spring period corresponding to the shoaling of the mixed layer and the increase of light intensity. Grazing selectivity however contributes to the phytoplankton community composition in summer.

Oithona similislikes it cool: evidence from two long-term time series

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.

Ocean Net Heat Flux Influences Seasonal to Interannual Patterns of Plankton Abundance

Changes in the net heat flux (NHF) into the ocean have profound impacts on global climate. We analyse a long-term plankton time-series and show that the NHF is a critical indicator of ecosystem dynamics. We show that phytoplankton abundance and diversity patterns are tightly bounded by the switches between negative and positive NHF over an annual cycle. Zooplankton increase before the transition to positive NHF in the spring but are constrained by the negative NHF switch in autumn. By contrast bacterial diversity is decoupled from either NHF switch, but is inversely correlated (r=-0.920) with the magnitude of the NHF. We show that the NHF is a robust mechanistic tool for predicting climate change indicators such as spring phytoplankton bloom timing and length of the growing season.

Seasonal movements and behaviour of basking sharks from archival tagging: No evidence of winter hibernation

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer-feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of continental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hibernate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (> 750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete productivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

Patterns of jellyfish abundance in the North Atlantic

A number of explanations have been advanced to account for the increased frequency and intensity at which jellyfish (pelagic cnidarians and ctenophores) blooms are being observed, most of which have been locally directed. Here, we investigate seasonal and inter-annual patterns in abundance and distribution of jellyfish in the North Atlantic Ocean to determine if there have been any system-wide changes over the period 1946–2005, by analysing records of the presence of coelenterates from the Continuous Plankton Recorder (CPR) survey. Peaks in jellyfish abundance are strongly seasonal in both oceanic and shelf areas: oceanic populations have a mid-year peak that is more closely related to peaks in phyto- and zooplankton, whilst the later peak of shelf populations mirrors changes in SST and reflects processes of advection and aggregation. There have been large amplitude cycles in the abundance of oceanic and shelf jellyfish (although not synchronous) over the last 60 years, with a pronounced synchronous increase in abundance in both areas over the last 10 years. Inter-annual variations in jellyfish abundance in oceanic areas are related to zooplankton abundance and temperature changes, but not to the North Atlantic Oscillation or to a chlorophyll index. The long-term inter-annual abundance of jellyfish on the shelf could not be explained by any environmental variables investigated. As multi-decadal cycles and more recent increase in jellyfish were obvious in both oceanic and shelf areas, we conclude that these are likely to reflect an underlying climatic signal (and bottom-up control) rather than any change in fishing pressure (top-down control). Our results also highlight the role of the CPR data in investigating long-term changes in jellyfish, and suggest that the cnidarians sampled by the CPR are more likely to be holoplanktic hydrozoans and not the much larger meroplanktic scyphozoans as has been suggested previously.

Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea

We study the spatial and seasonal variability of phytoplankton biomass (as phytoplankton color) in relation to the environmental conditions in the North Sea using data from the Continuous Plankton Recorder survey. By using only environmental fields and location as predictor variables we developed a nonparametric model (generalized additive model) to empirically explore how key environmental factors modulate the spatio-temporal patterns of the seasonal cycle of algal biomass as well as how these relate to the ,1988 North Sea regime shift. Solar radiation, as manifest through changes of sea surface temperature (SST), was a key factor not only in the seasonal cycle but also as a driver of the shift. The pronounced increase in SST and in wind speed after the 1980s resulted in an extension of the season favorable for phytoplankton growth. Nutrients appeared to be unimportant as explanatory variables for the observed spatio-temporal pattern, implying that they were not generally limiting factors. Under the new climatic regime the carrying capacity of the whole system has been increased and the southern North Sea, where the environmental changes have been more pronounced, reached a new maximum.

Seasonal plankton dynamics along a cross-shelf gradient off northern Spain

The development of population models able to reproduce the dynamics of zooplankton is a central issue when trying to understand how a changing environment would affect zooplankton in the future. Using 10 years of monthly data on phytoplankton and zooplankton abundance in the Bay of Biscay from the IEO's RADIALES time-series programme, we built non-parametric Generalized Additive Models (GAMs) able to reproduce the dynamics of plankton on the basis of environmental factors (nutrients, temperature, upwelling and photoperiod). We found that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors are non-linear. With the inclusion of the environmental variability, the main seasonal and inter-annual dynamic patterns observed within the studied plankton assemblage indicate the prevalence of bottom-up regulatory control. The statistically deduced models were used to simulate the dynamics of the phytoplankton and zooplankton. A good agreement between observations and simulations was obtained, especially for zooplankton. We are presently developing spatio-temporal GAM models for the North Sea based on the Continuous Plankton Recorder database.