Marine foams impede metabolic and behavioural traits in the rough periwinkle Littorina saxatilis.

Foams are a ubiquitous feature of marine environments. They can have major economic, societal and ecological consequences through their accumulation on the shore. Despite their pervasive nature and evidence that stable foam deposits play a pivotal role in the ecology of soft shore and estuaries, very limited amounts of information are available on their contribution to the structure and function at play in rocky intertidal ecosystems. This study shows that the metabolic rate of the high-shore gastropod Littorina saxatilis is significantly higher in individuals exposed to foams. Behavioural assays conducted under laboratory-controlled conditions further show that this species detects foam-born infochemicals both indirectly or directly, hence rely on both airborne and contact chemosensory cues. L. saxatilis also actively avoid areas covered in foam, and increase their activity in the presence of foam. These observations are interpreted in terms of foam-induced increased metabolic stress and increases behavioural anxiety and vigilance. They are further discussed in relation to the occurrence of two phytoplankton species known to produce repellent and/or toxic compounds such as domoic acid and dimethylsulfoniopropionate, the diatom Pseudo-nitzschia multistriata and the haptophyte Phaeocystis globosa, with the latter occurring at unusually high density. Taken together, these results suggest that the accumulation of foams on intertidal rocky shores may have major implications on taxa relying on both airborne and contact chemosensory cues to navigate, find food and mating partners. Specifically, the observed increased behavioural activity coupled with increased metabolic demands may impact species fitness and highlight potentially large ecological consequences in rocky intertidal ecosystems characterized by strong hydrodynamism and elevated organic matter content leading to the presence of long-lived foam.

On the conditions promoting Pseudo-nitzschia spp. blooms in the eastern English Channel and southern North Sea.

This study investigated the drivers of the blooms of Pseudo-nitzschia seriata and Pseudo-nitzschia delicatissima complexes in the eastern English Channel and southern North Sea. Phytoplankton data series acquired from 1992 to 2020 were analyzed with a multivariate statistical approach based on Hutchinson's niche concept. P. seriata and P. delicatissima complexes were found to be typically present year round, but they bloomed at different periods because they occupied different realized ecological niches. P. delicatissima complex occupied a more marginal niche and was less tolerant than P. seriata complex. P. delicatissima complex typically bloomed in April-May at the same time as Phaeocystis globosa while P. seriata complex blooms were more frequently observed in June during the decline of low intensity P. globosa blooms. P. delicatissima and P. seriata complexes were both favored by low-silicate environments and relatively low turbulence but they responded differently to water temperature, light, ammonium, phosphate and nitrite + nitrate conditions. Niche shifts and biotic interactions played important roles in the control of the blooms of P. delicatissima and P. seriata complexes. The two complexes occupied different sub-niches during their respective low abundance and bloom periods. The phytoplankton community structure and the number of other taxa presenting a niche overlapping the niches of P. delicatissima and P. seriata complexes also differed between these periods. P. globosa was the taxa contributing the most to the dissimilarity in community structure. P. globosa interacted positively with P. delicatissima complex and negatively with P. seriata complex.

Stochastic and Deterministic Processes Regulate Phytoplankton Assemblages in a Temperate Coastal Ecosystem.

Assessing the relative contributions of the interacting deterministic and stochastic ecological processes for phytoplankton community assembly is crucial in understanding and predicting community organization and succession at different temporal and spatial scales. In this study, we hypothesized that deterministic and stochastic ecological processes regulating phytoplankton, present seasonal and repeating patterns. This hypothesis was explored during a 5-year survey (287 samples) conducted at a small spatial scale (~15km) in a temperate coastal ecosystem (eastern English Channel). Microscopy and flow cytometry quantified phytoplankton abundance and biomass, while metabarcoding data allowed an extended evaluation of diversity and the exploration of the ecological processes regulating phytoplankton using null model analysis. Alpha diversity of phytoplankton was governed by the effect of environmental conditions (environmental filtering). Temporal community turnover (beta diversity) evidenced a consistent interannual pattern that determined the phytoplankton seasonal structure. In winter and early spring (from January to March), determinism (homogeneous selection) was the major process in the phytoplankton community assembly. The overall mean in the year was 38%. Stochastic processes (ecological drift) prevailed during the rest of the year from April to December, where the overall mean for the year was 55%. The maximum values were recorded in late spring and summer, which often presented recurrent and transient monospecific phytoplankton peaks. Overall, the prevalence of stochastic processes rendered less predictable seasonal dynamics of phytoplankton communities to future environmental change. IMPORTANCE While ecological deterministic processes are conducive to modeling, stochastic ones are far less predictable. Understanding the overall assembly processes of phytoplankton is critical in tracking and predicting future changes. The novelty of this study was that it addressed a long-posed question, on a pluriannual scale. Was seasonal phytoplankton succession influenced by deterministic processes (e.g., abiotic environment) or by stochastic ones (e.g., dispersal, or ecological drift)? Our results provided strong support for a seasonal and repeating pattern with stochastic processes (drift) prevailing during most of the year and periods with monospecific phytoplankton peaks.

Diversity and potential activity patterns of planktonic eukaryotic microbes in a mesoeutrophic coastal area (eastern English Channel).

The diversity of planktonic eukaryotic microbes was studied at a coastal station of the eastern English Channel (EEC) from March 2011 to July 2015 (77 samples) using high throughput sequencing (454-pyrosequencing and Illumina) of the V2-V3 hypervariable region of the 18S SSU rDNA gene. Similar estimations of OTU relative abundance and taxonomic distribution for the dominant higher taxonomic groups (contributing >1% of the total number of OTUs) were observed with the two methods (Kolmogorov-Smirnov p-value = 0.22). Eight super-groups were identified throughout all samples: Alveolata, Stramenopiles, Opisthokonta, Hacrobia, Archeaplastida, Apusozoa, Rhizaria, and Amoebozoa (ordered by decreasing OTU richness). To gain further insight into microbial activity in the EEC, ribosomal RNA was extracted for samples from 2013-2015 (30 samples). Analysis of 18S rDNA and rRNA sequences led to the detection of 696 and 700 OTUs, respectively. Cluster analysis based on OTUs' abundance indicated three major seasonal groups that were associated to spring, winter/autumn, and summer conditions. The clusters inferred from rRNA data showed a clearer seasonal representation of the community succession than the one based on rDNA. The rRNA/rDNA ratio was used as a proxy for relative cell activity. When all OTUs were considered, the average rRNA:rDNA ratio showed a linear trend around the 1:1 line, suggesting a linear relation between OTU abundance (rDNA) and activity (rRNA). However, this ratio was highly variable over time when considering individual OTUs. Interestingly, the OTU affiliated with P. globosa displayed rRNA:rDNA ratio that allowed to delimit high vs low abundance and high vs low activity periods. It unveiled quite well the Phaeocystis bloom dynamic regarding cell proliferation and activity, and could even be used as early indicator of an upcoming bloom.

Realized niche analysis of phytoplankton communities involving HAB: Phaeocystis spp. as a case study.

The link between harmful algal blooms, phytoplankton community dynamics and global environmental change is not well understood. To tackle this challenging question, a new method was used to reveal how phytoplankton communities responded to environmental change with the occurrence of an harmful algae, using the coastal waters of the eastern English Channel as a case study. The great interannual variability in the magnitude and intensity of Phaeocystis spp. blooms, along with diatoms, compared to the ongoing gradual decrease in anthropogenic nutrient concentration and rebalancing of nutrient ratios; suggests that other factors, such as competition for resources, may also play an important role. A realized niche approach was used with the Outlying Mean Index analysis and the dynamics of the species' realized subniches were estimated using the Within Outlying Mean Indexes calculations under low (L) and high (H) contrasting Phaeocystis spp. abundance. The Within Outlying Mean Indexes allows the decomposition of the realized niche into realized subniches, found within the subset of habitat conditions and constrained by a subset of a biotic factor. The two contrasting scenarios were characterized by significantly different subsets of environmental conditions and diatom species (BV-step analysis), and different seasonality in salinity, turbidity, and nutrients. The subset L environmental conditions were potentially favorable for Phaeocystis spp. but it suffered from competitive exclusion by key diatom species such as Skeletonema spp., Thalassiosira gravida, Thalassionema nitzschioides and the Pseudo-nitzchia seriata complex. Accordingly, these diatoms species occupied 81% of Phaeocystis spp.'s existing fundamental subniche. In contrast, the greater number of diatoms, correlated with the community trend, within subset H exerted a weaker biological constraint and favored Phaeocystis spp. realized subniche expansion. In conclusion, the results strongly suggest that both abiotic and biotic interactions should be considered to understand Phaeocystis spp. blooms with greater consideration of the preceeding diatoms. HABs needs must therefore be studied as part of the total phytoplankton community.

Environmental control on fish and macrocrustacean spring community-structure, on an intertidal sandy beach.

The inter-annual variability of the fish and macrocrustacean spring community on an intertidal sandy beach near the Canche estuary (North of France) was studied from 2000 to 2013 based on weekly spring sampling over an 11-year period. Twenty-eight species representing 21 families were collected during the course of the study. The community was dominated by a few abundant species accounting for > 99% of the total species densities. Most individuals caught were young-of-the-year indicating the importance of this ecosystem for juvenile fishes and macrocrustaceans. Although standard qualitative community ecology metrics (species composition, richness, diversity, evenness and similarity) indicated notable stability over the study period, community structure showed a clear change since 2009. Densities of P. platessa, P. microps and A. tobianus decreased significantly since 2009, whereas over the period 2010-2013, the contribution of S. sprattus to total species density increased 4-fold. Co-inertia and generalised linear model analyses identified winter NAO index, water temperature, salinity and suspended particular matter as the major environmental factors explaining these changes. Although the recurrent and dense spring blooms of the Prymnesiophyte Phaeocystis globosa is one of the main potential threats in shallow waters of the eastern English Channel, no negative impact of its temporal change was detected on the fish and macrocrustacean spring community structure.

Microplanktonic community structure in a coastal system relative to a Phaeocystis bloom inferred from morphological and tag pyrosequencing methods.

BACKGROUND: Massive phytoplankton blooms, like the recurrent Phaeocystis proliferation observed every year in the Eastern English Channel (EEC), have a significant influence on the overall planktonic community structure and their food web dynamics. As well as being an important area for local fisheries, the EEC is an ideal ecosystem for work on microbial diversity. This is because, although its environmental context is relatively complex, it is reasonably well understood due to several years of monitoring and morphological observations of its planktonic organisms. The objective of our study was to better understand the under-explored microbial eukaryotic diversity relative to the Phaeocystis bloom. METHODOLOGY AND PRINCIPAL FINDINGS: The community structure of microplankton (diatoms, haptophytes, ciliates and dinoflagellates) was studied through morphological observations and tag pyrosequencing. During the annual Phaeocystis spring bloom, the phytoplankton biomass increased by 34-fold, while the microzooplankton biomass showed a 4-fold increase, representing on average about 4.6% of the biomass of their phytoplankton prey. Tag pyrosequencing unveiled an extensive diversity of Gymnodiniaceae, with G. spirale and G. fusiformis representing the most abundant reads. An extended diversity of Phaeocystales, with partial 18S rDNA genes sequence identity as low as 85% was found, with taxa corresponding to P. globosa, but also to unknown Phaeocystaceae. CONCLUSIONS: Morphological analyses and pyrosequencing were generally in accordance with capturing frequency shifts of abundant taxa. Tag pyrosequencing allowed highlighting the maintenance of microplankton diversity during the Phaeocystis bloom and the increase of the taxa presenting low number of reads (minor taxa) along with the dominant ones in response to biotic and/or abiotic changing conditions. Although molecular approaches have enhanced our perception on diversity, it has come to light that the challenge of modelling and predicting ecological change requires the use of different complementary approaches, to link taxonomic data with the functional roles of microbes in biogeochemical cycles.