Investigation of benthic community change over a century-wide scale in the western English Channel

Since the early part of the 20th Century the impact of a range of anthropogenic activities in our coastal seas has steadily increased. The effect of such activities is a major cause for concern but in the benthic environment few studies exist that date back more than a few decades. Hence understanding long term changes is a challenge. Within this study we utilized a historic benthic dataset and resurveyed an area west of Eddystone reef in the English Channel previously investigated 112 years ago. The aim of the present work was to describe the current benthic community structure and investigate potential differences between 1895 and 2007. For each of the four major phyla investigated (Polychaeta, Crustacea, Mollusca and Echinodermata), multivariate community analysis showed significant differences between the historic and contemporary surveys. Echinoderm diversity showed a clear reduction between 1895 and 2007. The sea urchins Echinus esculentus, Spatangus purpureus, and Psammechinus miliaris and large star-fish Marthasterias glacialis showed reductions in abundance, in some cases being entirely absent from the survey area in 2007. Polychaetes showed a shift from tubiculous species to small errant and predatory species such as Glycera, Nephtys, and Lumbrineris spp. Within the group Mollusca large species such as Pecten maximus and Laevicardium crassum decreased in abundance while small species increased. Crustaceans in 1895 were dominated by crab species which were present in similar abundances in 2007, but, the order Amphipoda appeared to show a significant increase. While some of the differences observed could stem from differences in methodologies between the surveys, in particular increases of small cryptic species, the loss of large conspicuous species was judged to be genuine. The study area is an important beam trawling and scallop dredging ground; the differences observed are concomitant with changes generally associated with disturbance from demersal fishing activities such as these.

Long-term variability of the siphonophores Muggiaea atlantica and M. kochi in the Western English Channel

We investigated long-term variability of the calycophoran siphonophores Muggiaea atlantica and Muggiaea kochi in the Western English Channel (WEC) between 1930 and 2011. Our aims were to describe long-term changes in abundance and temporal distribution in relation to local environmental dynamics. In order to better understand mechanisms that regulate the species’ populations, we identified periods that were characteristic of in situ population growth and the environmental optima associated with these events. Our results show that between 1930 and the 1960s both M. atlantica and M. kochi were transient components of the WEC ecosystem. In the late 1960s M. atlantica, successfully established a resident population in the WEC, while the occurrence of M. kochi became increasingly sporadic. Once established as a resident species, the seasonal abundance and distribution of M. atlantica increased. Analysis of environmental conditions associated with in situ population growth revealed that temperature and prey were key determinants of the seasonal distribution and abundance of M. atlantica. Salinity was shown to have an indirect effect, likely representing a proxy for water circulation in the WEC. Anomalies in the seasonal cycle of salinity, indicating deviation from the usual circulation pattern in the WEC, were negatively associated with in situ growth, suggesting dispersal of the locally developing M. atlantica population. However, our findings identified complexity in the relationship between characteristics of the environment and M. atlantica variability. The transition from a period of transiency (1930–1968) to residency (1969–2011) was tentatively attributed to structural changes in the WEC ecosystem that occurred under the forcing of wider-scale hydroclimatic changes.

The combined effects of seasonal community succession and adaptive algal physiology on lipid profiles of coastal phytoplankton in the western English channel

Lipids are key constituents of marine phytoplankton, and some fatty acids (key constituents of lipids) are essential dietary components for secondary producers. However, in natural marine ecosystems the interactions of factors affecting seasonal phytoplankton lipid composition are still poorly understood. The aim of this study was to assess the roles of seasonal succession in phytoplankton community composition and nutrient concentrations, on the lipid composition of the phytoplankton community. Fatty acid and polar lipid composition in seston was measured in surface waters at the time series station L4, an inshore station in the Western English Channel, from January to December 2013. Redundancy analyses (RDA) were used to identify factors (abiotic and biotic) that explained the seasonal variability in phytoplankton lipids. RDA demonstrated that nutrients (namely nitrogen) explained the majority of variation in phytoplankton lipid composition, as well as a smaller explanatory contribution from changes in phytoplankton community composition. The physiological adaptations of the phytoplankton community to nutrient deplete conditions during the summer season when the water column was stratified, was further supported by changes in the polar lipid to phytoplankton biomass ratios (also modelled with RDA) and increases in the lipid to chlorophyll a ratios, which are both indicative of nutrient stress. However, the association of key fatty acid markers with phytoplankton groups e.g. 22:6 n-3 and dinoflagellate biomass (predominant in summer), meant there were no clear seasonal differences in the overall degree of fatty acid saturation, as might have been expected from typical nutrient stress on phytoplankton. Based on the use of polyunsaturated fatty acids (PUFA) as markers of ‘food quality’ for grazers, our results suggest that in this environment high food quality is maintained throughout summer, due to seasonal succession towards flagellated phytoplankton species able to maintain PUFA synthesis under surface layer nutrient depletion.

Feeding rates and prey selectivity of planktonic decapod larvae in the Western English Channel

Meroplankton are seasonally important contributors to the zooplankton, particularly at inshore sites, yet their feeding ecology is poorly known relative to holoplankton. While several studies have measured feeding in decapod larvae, few studies have examined the feeding rates of decapod larvae on natural prey assemblages throughout the reproductive season. We conducted 8 feeding experiments with Necora puber, Liocarcinus spp. and Upogebia spp. zoea larvae collected from the L4 monitoring site off Plymouth (50°15.00′N, 4°13.02′W) during spring–summer 2009 and 2010. This period spanned moderate-to-high food availability (0.5–1.6 µg chl-a L−1), but a great range in food composition with small cells <20 µm dominating in 2010. Daily rations averaged 17, 60 and 22 % of body C for the 3 respective decapod species. Clearance rates differed according to prey type, and all 3 decapod genera showed evidence of selection of dinoflagellates. Importantly, small cells including nano- and pico-plankton were ingested, this being demonstrated independently by flow cytometric analysis of the feeding experiments and molecular analysis. PCR-based analysis of the haptophyte portion of the diet revealed ingestion of Isochrysis galbana by decapod larvae in the bottle incubations and Isochrysis galbana and Phaeocystis globosa by decapod larvae collected directly from the field. This study has shown that pico- and nano-sized plankton form an important supplement to the diverse and variable diet of decapod larvae.

Feeding selectivity of bivalve larvae on natural plankton assemblages in the Western English Channel

Meroplankton, including bivalve larvae, are an important and yet understudied component of coastal marine food webs. Understanding the baseline of meroplankton ecology is imperative to establish and predict their sensitivity to local and global marine stressors. Over an annual cycle (October 2009–September 2010), bivalve larvae were collected from the Western Channel Observatory time series station L4 (50°15.00′N, 4°13.02′W). The morphologically similar larvae were identified by analysis of the 18S nuclear small subunit ribosomal RNA gene, and a series of incubation experiments were conducted to determine larval ingestion rates on natural plankton assemblages. Complementary gut content analysis was performed using a PCR-based method for detecting prey DNA both from field-collected larvae and those from the feeding experiments. Molecular identification of bivalve larvae showed the community composition to change over the course of the sampling period with domination by Phaxas in winter and higher diversity in autumn. The larvae selected for nanoeukaryotes (2–20 µm) including coccolithophores (<20 µm) which together comprised >75 % of the bivalve larvae diet. Additionally, a small percentage of carbon ingested originated from heterotrophic ciliates (<30 µm). The molecular analysis of bivalve larvae gut content provided increased resolution of identification of prey consumed and demonstrated that the composition of prey consumed established through bottle incubations conferred with that established from in situ larvae. Despite changes in bivalve larvae community structure, clearance rates of each prey type did not change significantly over the course of the experiment, suggesting different bivalve larvae species may consume similar prey.

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.

Abundance of a chlorophyll a precursor and the oxidation product hydroxychlorophyll a during seasonal phytoplankton community progression in the Western English Channel

This study presents the first in-situ measurements of the chlorophyll a oxidation product, hydroxychlorophyll a as well as the chlorophyll a precursor, chlorophyll aP276 conducted over an annual cycle. Chlorophyll a oxidation products, such as hydroxychlorophyll a may be associated with the decline of algal populations and can act as an initial step in the degradation of chlorophyll a into products which can be found in the geochemical record, important for studying past climate change events. Here, hydroxychlorophyll a and chlorophyll aP276 were measured at the long-term monitoring station L4, Western Channel Observatory (UK, www.westernchannelobservatory.org) over an annual cycle (2012). Weekly measurements of phytoplankton species composition and abundance enabled detailed analysis of possible sources of hydroxychlorophyll a. Dinoflagellates, 2 diatom species, the prymnesiophyte Phaeocystis spp. and the coccolithophorid Emiliania huxleyi were all associated with hydroxychlorophyll a occurrence. However, during alternate peaks in abundance of the diatoms, no association with hydroxychlorophyll a occurred, indicating that the oxidation of chlorophyll a was dependant not only on species but also on additional factors such as the mode of mortality, growth limiting factor (i.e. nutrient concentration) or phenotypic plasticity. Surface sediment samples contained 10 times more hydroxychlorophyll a (relative to chlorophyll a) than pelagic particulate samples, indicating that more chlorophyll a oxidation occurred during sedimentation or at the sediment–water interface, than in the pelagic environment. In addition, chlorophyll aP276 correlated with chl-a concentration, thus supporting its assignment as a chl-a precursor.

Dynamic responses of the benthic bacterial community at the Western English Channel observatory site L4 are driven by deposition of fresh phytodetritus

The impact of the seasonal deposition of phytoplankton and phytodetritus on surface sediment bacterial abundance and community composition was investigated at the Western English Channel site L4. Sediment and water samples were collected from January to September in 2012, increasing in frequency during periods of high water column phytoplankton abundance. Compared to the past two decades, the spring bloom in 2012 was both unusually long in duration and contained higher than average biomass. Within spring months, the phytoplankton bloom was well mixed through the water column and showed accumulations near the sea bed, as evidenced by flow cytometry measurements of nanoeukaryotes, water column chlorophyll a and the appearance of pelagic phytoplankton at the sediment. Measurements of chlorophyll and chlorophyll degradation products indicated phytoplankton material was heavily degraded after it reached the sediment surface: the nature of the chlorophyll degradation products (predominantly pheophorbide, pyropheophorbide and hydroxychlorophyllone) was indicative of grazing activity. The abundance of bacterial 16S rRNA genes g−1 sediment (used as a proxy for bacterial biomass) increased markedly with the onset of the phytoplankton bloom, and correlated with measurements of chlorophyll at the surface sediment. Together, this suggests that bacteria may have responded to nutrients released via grazing activity. In depth sequencing of the 16S rRNA genes indicated that the composition of the bacterial community shifted rapidly through-out the prolonged spring bloom period. This was primarily due to an increase in the relative sequence abundance of Flavobacteria.

Decrease in diatom palatability contributes to bloom formation in the Western English Channel

The aim of this paper is to investigate the role of phytoplankton nutritional status in the formation of the spring bloom regularly observed at the station L4 in the Western English Channel. Using a modelling approach, we tested the hypothesis that the increase in light from winter to spring induces a decrease in diatom nutritional status (i.e., an increase in the C:N and C:P ratios), thereby reducing their palatability and allowing them to bloom. To this end, a formulation describing the Stoichiometric Modulation of Predation (SMP) has been implemented in a simplified version of the European Regional Seas Ecosystem Model (ERSEM). The model was coupled with the General Ocean Turbulence Model (GOTM), implemented at the station L4 and run for 10 years (2000-.2009). Simulated carbon to nutrient ratios in diatoms were analysed in relation to microzooplankton biomass, grazing and assimilation efficiency. The model reproduced in situ data evolutions and showed the importance of microzooplankton grazing in controlling the early onset of the bloom. Simulation results supported our hypothesis and provided a conceptual model explaining the formation of the diatom spring bloom in the investigated area. However, additional data describing the microzooplankton grazing impact and the variation of carbon to nutrient ratios inside phytoplanktonic cells are required to further validate the proposed mechanisms.