The Importance of Regional, System-Wide and Local Spatial Scales in Structuring Temperate Estuarine Fish Communities

An extensive literature base worldwide demonstrates how spatial differences in estuarine fish assemblages are related to those in the environment at (bio)regional, estuary-wide or local (within-estuary) scales. Few studies, however, have examined all three scales, and those including more than one have often focused at the level of individual environmental variables rather than scales as a whole. This study has identified those spatial scales of environmental differences, across regional, estuary-wide and local levels, that are most important in structuring ichthyofaunal composition throughout south-western Australian estuaries. It is the first to adopt this approach for temperate microtidal waters. To achieve this, we have employed a novel approach to the BIOENV routine in PRIMER v6 and a modified global BEST test in an alpha version of PRIMER v7. A combination of all three scales best matched the pattern of ichthyofaunal differences across the study area (rho = 0.59; P = 0.001), with estuary-wide and regional scales accounting for about twice the variability of local scales. A shade plot analysis showed these broader-scale ichthyofaunal differences were driven by a greater diversity of marine and estuarine species in the permanently-open west coast estuaries and higher numbers of several small estuarine species in the periodically-open south coast estuaries. When interaction effects were explored, strong but contrasting influences of local environmental scales were revealed within each region and estuary type. A quantitative decision tree for predicting the fish fauna at any nearshore estuarine site in south-western Australia has also been produced. The estuarine management implications of the above findings are highlighted.

Long-term iceshelf-covered meiobenthic communities of the Antarctic continental shelf resemble those of the deep sea

Since strong regional warming has led to the disintegration of huge parts of the Larsen A and B ice shelves east of the Antarctic Peninsula in 1995 and 2002, meiofaunal communities covered by ice shelves for thousands of years could be investigated for the first time. Based on a dataset of more than 230,000 individuals, meiobenthic higher taxa diversity and composition of Larsen continental shelf stations were compared to those of deep-sea stations in the Western Weddell Sea to see whether the food-limiting conditions in the deep sea and the food-poor shelf regime at times of iceshelf coverage has resulted in similar meiobenthic communities, on the premises that food availability is the main driver of meiobenthic assemblages. We show here that this is indeed the case; in terms of meiobenthic communities, there is greater similarity between the deep sea and the inner Larsen embayments than there is similarity between the deep sea and the former Larsen B iceshelf edge and the open continental shelf. We also show that resemblance to Antarctic deep-sea meiofaunal communities was indeed significantly higher for communities of the innermost Larsen B area than for those from intermediate parts of Larsen A and B. Similarity between communities from intermediate parts and the deep sea was again higher than between those of the ice-edge and the open shelf. Meiofaunal densities were low at the inner parts of Larsen A and B, and comparable to deep-sea densities, again likely owing to the low food supply at both habitats. We suggest that meiobenthic communities have not yet recovered from the food-limiting conditions present at the time of iceshelf coverage. Meiofaunal diversity on the other hand seemed driven by sediment structure, being higher in coarser sediments.

Disturbance to conserved bacterial communities in the cold water gorgonian coral Eunicella verrucosa

The bacterial communities associated with healthy and diseased colonies of the cold-water gorgonian coral Eunicella verrucosa at three sites off the south-west coast of England were compared using denaturing gradient gel electrophoresis (DGGE) and clone libraries. Significant differences in community structure between healthy and diseased samples were discovered, as were differences in the level of disturbance to these communities at each site; this correlated with depth and sediment load. The majority of cloned sequences from healthy coral tissue affiliated with the Gammaproteobacteria. The stability of the bacterial community and dominance of specific genera found across visibly healthy colonies suggest the presence of a specific microbial community. Affiliations included a high proportion of Endozoicomonas sequences, which were most similar to sequences found in tropical corals. This genus has been found in a number of invertebrates and is suggested to have a role in coral health and in the metabolisation of dimethylsulfoniopropionate (DMSP) produced by zooxanthellae. However, screening of colonies for the presence of zooxanthellae produced a negative result. Diseased colonies showed a decrease in affiliated clones and an increase in clones related to potentially harmful/transient microorganisms but no increase in a particular pathogen. This study demonstrates that a better understanding of these bacterial communities, the factors that affect them and their role in coral health and disease will be of critical importance in predicting future threats to temperate gorgonian communities.

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.

Polychaete burrows harbour distinct microbial communities in oil-contaminated coastal sediments

Previous studies have shown that the bioturbating polychaete Hediste (Nereis) diversicolor can affect the composition of bacterial communities in oil-contaminated sediments, but have not considered diversity specifically within bioturbator burrows or the impact on microbial eukaryotes. We tested the hypothesis that H. diversicolor burrows harbour different eukaryotic and bacterial communities compared with un-bioturbated sediment, and that bioturbation stimulates oil degradation. Oil-contaminated sediment was incubated with or without H. diversicolor for 30 days, after which sediment un-affected by H. diversicolor and burrow DNA/RNA samples were analysed using quantitative reverse transcription PCR (Q-RT-PCR) and high-throughput sequencing. Fungi dominated both burrow and un-bioturbated sediment sequence libraries; however, there was significant enrichment of bacterivorous protists and nematodes in the burrows. There were also significant differences between the bacterial communities in burrows compared with un-bioturbated sediment. Increased activity and relative abundance of aerobic hydrocarbon-degrading bacteria in the burrows coincided with the significant reduction in hydrocarbon concentration in the bioturbated sediment. This study represents the first detailed assessment of the effect of bioturbation on total microbial communities in oil-contaminated sediments. In addition, it further shows that bioturbation is a significant factor in determining microbial diversity within polluted sediments and plays an important role in stimulating bioremediation.

Biological Communities at Marine Shallow-Water Vent and Seep Sites

The review compiles, for the first time, data on the communities at 62 shallow-water hydrothermal vent and cold seep sites. ‘Shallow sites’ are defined as sites no deeper than 200 m. The communities at these sites are also compared with communities in reducing sediments at similar depths. Below 200 m, vent and seep obligate species tend to dominate the fauna living in areas where reducing fluids are released from the seabed. At the shallow sites, vent and seep obligate species of fauna are rare, only eight having being reported from shallow vents. No definite seep obligates have been found. Shallow vents and seeps are colonized by communities that consist of a subset of the background fauna, especially those species that are less sensitive to hydrogen sulphide toxicity. Conversely the zones directly surrounding shallow vent and seeps sites with varied topography, substrate type and food supply, often have a higher species diversity than the background area. The reasons for these differences are discussed.

Nanoplankton and picoplankton in the Western English Channel: abundance and seasonality from 2007–2013

The nano- and picoplankton community at Station L4 in the Western English Channel was studied between 2007 and 2013 by flow cytometry to quantify abundance and investigate seasonal cycles within these communities. Nanoplankton included both photosynthetic and heterotrophic eukaryotic single-celled organisms while the picoplankton included picoeukaryote phytoplankton, Synechococcus sp. cyanobacteria and heterotrophic bacteria. A Box–Jenkins Transfer Function climatology analysis of surface data revealed that Synechococcus sp., cryptophytes, and heterotrophic flagellates had bimodal annual cycles. Nanoeukaryotes and both high and low nucleic acid-containing bacteria (HNA and LNA, respectively) groups exhibited unimodal annual cycles. Phaeocystis sp., whilst having clearly defined abundance maxima in spring was not detectable the rest of the year. Coccolithophores exhibited a weak seasonal cycle, with abundance peaks in spring and autumn. Picoeukaryotes did not exhibit a discernable seasonal cycle at the surface. Timings of maximum group abundance varied through the year. Phaeocystis sp. and heterotrophic flagellates peaked in April/May. Nanoeukaryotes and HNA bacteria peaked in June/July and had relatively high abundance throughout the summer. Synechococcus sp., cryptophytes and LNA bacteria all peaked from mid to late September. The transfer function model techniques used represent a useful means of identifying repeating annual cycles in time series data with the added ability to detect trends and harmonic terms at different time scales from months to decades.

Effect of habitat fragmentation on the macroinvertebrate infaunal communities associated with the seagrass Zostera marina

1. The effect of habitat fragmentation was investigated in two adjacent, yet separate, intertidal Zostera marina beds in the Salcombe Estuary, Devon, UK. The seagrass bed on the west bank comprised a continuous meadow of ca. 2.3 ha, whilst the bed on the east bank of the estuary was fragmented into patches of 6–9 m2.2. Three 10 cm diameter core samples for infaunal macroinvertebrates were taken from three stations within each bed. No significant difference was found in univariate community parameters between beds, or in measured seagrass parameters. However, multivariate analysis revealed a significant difference in community composition, due mainly to small changes in species abundance rather than differences in the species present.3. The species contributing most to the dissimilarity between the two communities were polychaetes generally associated with unvegetated habitats (e.g. Magelona mirabilis) and found to be more common in the fragmented bed.4. A significant difference in median grain size and sorting coefficient was recorded between the two beds, and median grain size was found to be the variable best explaining multivariate community patterns.5. The results of the study provide evidence for the effects of habitat fragmentation on the communities associated with seagrass beds, habitats which are of high conservation importance. As the infaunal community is perhaps intuitively the component least likely to be affected by fragmentation at the scale observed, the significant difference in community composition recorded has consequences for more sensitive and high-profile parts of the biota (e.g. fish), and thus for the conservation of seagrass habitats and their associated communities.

Probabilistic Models to Describe the Dynamics of Migrating Microbial Communities

In all but the most sterile environments bacteria will reside in fluid being transported through conduits and some of these will attach and grow as biofilms on the conduit walls. The concentration and diversity of bacteria in the fluid at the point of delivery will be a mix of those when it entered the conduit and those that have become entrained into the flow due to seeding from biofilms. Examples include fluids through conduits such as drinking water pipe networks, endotracheal tubes, catheters and ventilation systems. Here we present two probabilistic models to describe changes in the composition of bulk fluid microbial communities as they are transported through a conduit whilst exposed to biofilm communities. The first (discrete) model simulates absolute numbers of individual cells, whereas the other (continuous) model simulates the relative abundance of taxa in the bulk fluid. The discrete model is founded on a birth-death process whereby the community changes one individual at a time and the numbers of cells in the system can vary. The continuous model is a stochastic differential equation derived from the discrete model and can also accommodate changes in the carrying capacity of the bulk fluid. These models provide a novel Lagrangian framework to investigate and predict the dynamics of migrating microbial communities. In this paper we compare the two models, discuss their merits, possible applications and present simulation results in the context of drinking water distribution systems. Our results provide novel insight into the effects of stochastic dynamics on the composition of non-stationary microbial communities that are exposed to biofilms and provides a new avenue for modelling microbial dynamics in systems where fluids are being transported.

Photoacclimation of natural phytoplankton communities

Phytoplankton regulate internal pigment concentrations in response to light and nutrient availability. Chlorophyll to carbon ratios (Chl:Cphyto) are commonly reported as a function of growth irradiance (Eg) for evaluating the photoacclimation response of phytoplankton. In contrast to most culture experiments, natural phytoplankton communities experience fluctuating environmental conditions making it difficult to compare field and lab observations. Observing and understanding photoacclimation in nature is important for deciphering changes in Chl:Cphyto resulting from environmental forcings and for accurately estimating net primary production (NPP) in models which rely on a parameterized description of photoacclimation. Here we employ direct analytical measurements of Cphyto and parallel high-resolution biomass estimates from particulate backscattering (bbp) and flow cytometry to investigate Chl:Cphyto in natural phytoplankton communities. Chl:Cphyto observed over a wide range of Eg in the field was consistent with photoacclimation responses inferred from satellite observations. Field-based photoacclimation observations for a mixed natural community contrast with laboratory results for single species grown in continuous light and nutrient replete conditions. Applying a carbon-based net primary production (NPP) model to our field data for a north-south transect in the Atlantic Ocean results in estimates that closely match 14C depth-integrated NPP for the same cruise and with historical records for the distinct biogeographic regions of the Atlantic Ocean. Our results are consistent with previous satellite and model observations of cells growing in natural or fluctuating light and showcase how direct measurements of Cphyto can be applied to explore phytoplankton photophysiology, growth rates, and production at high spatial resolution in-situ.

Dip in the gene pool: metagenomic survey of natural coccolithovirus communities

Despite the global oceanic distribution and recognised biogeochemical impact of coccolithoviruses (EhV), their diversity remains poorly understood. Here we employed a metagenomic approach to study the occurrence and progression of natural EhV community genomic variability. Analysis of EhV metagenomes from the early and late stages of an induced bloom led to three main discoveries. First, we observed resilient and specific genomic signatures in the EhV community associated with the Norwegian coast, which reinforce the existence of limitations to the capacity of dispersal and genomic exchange among EhV populations. Second, we identified a hyper-variable region (approximately 21kbp long) in the coccolithovirus genome. Third, we observed a clear trend for EhV relative amino-acid diversity to reduce from early to late stages of the bloom. This study validated two new methodological combinations, and proved very useful in the discovery of new genomic features associated with coccolithovirus natural communities.

Bacterioviridins: novel sedimentary chlorins providing evidence for oxidative processes affecting palaeobacterial communities

Of novel sedimentary transformation products of bacteriochlorophyll a, 7,8-didehydro derivatives (Bacterioviridins), has been identified using high performance liquid chromatography with photodiode array detection and atmospheric pressure chemical ionisation liquid chromatography-multistage mass spectrometry. The derivatives occur in a range of sediments including Priest Pot (UK), les Salines de la Trinital, Ebre delta (Spain), Sutton-on-the-Forest (UK) and Certes (France). The Bacterioviridins are products of oxidative transformation of bacteriochlorophyll a, and their presence in sediments indicates exposure of the bacterial community to oxygen.

An unusually large phytoplankton spring bloom drives rapid changes in benthic diversity and ecosystem function

In 2012, the Western English Channel experienced an unusually large and long-lived phytoplankton spring bloom. When compared with data from the past 20 years, average phytoplankton biomass at Station L4 (part of the Western Channel Observatory) was approximately 3× greater and lasted 50% longer than any previous year. Regular (mostly weekly) box core samples were collected from this site before, during and after the bloom to determine its impact on macrofaunal abundance, diversity, biomass, community structure and function. The spring bloom of 2012 was shown to support a large and rapid response in the majority of benthic taxa and functional groups. However, key differences in the precise nature of this response, as well as in its timing, was observed between different macrofauna feeding groups. Deposit feeders responded almost instantly at the start of the bloom, primarily thorough an increase in abundance. Suspension feeders and opportunistic/predatory/carnivorous taxa responded slightly more slowly and primarily with an increase in biomass. At the end of the bloom a rapid decline in macrobenthic abundance, diversity and biomass closely followed the decline in phytoplankton biomass. With suspension feeders showing evidence of this decline a few weeks before deposit feeders, it was concluded that this collapse in benthic communities was driven primarily by food availability and competition. However, it is possible that environmental hypoxia and the presence of toxic benthic cyanobacteria could also have contributed to this decline. This study shows evidence for strong benthic–pelagic coupling at L4; a shallow (50 m), coastal, fine-sand habitat. It also demonstrates that in such habitats, it is not just planktonic organisms that demonstrate clear community phenology. Different functional groups within the benthic assemblage will respond to the spring bloom in specific manner, with implications for key ecosystem functions and processes, such as secondary production and bioturbation. Only by taking integrated benthic and pelagic observations over such fine temporal scales (weekly) was the current study able to identify the intimate structure of the benthic response. Similar studies from other habitats and under different bloom conditions are urgently needed to fully appreciate the strength of benthic–pelagic coupling in shallow coastal environments.

Modelling the future biogeography of North Atlantic zooplankton communities in response to climate change

Advances in habitat and climate modelling allow us to reduce uncertainties of climate change impacts on species distribution. We evaluated the impacts of future climate change on community structure, diversity, distribution and phenology of 14 copepod species in the North Atlantic. We developed and validated habitat models for key zooplankton species using continuous plankton recorder (CPR) survey data collected at mid latitudes of the North Atlantic. Generalized additive models (GAMs) were applied to relate the occurrence of species to environmental variables. Models were projected to future (2080–2099) environmental conditions using coupled hydroclimatix–biogeochemical models under the Intergovernmental Panel on Climate Change (IPCC) A1B climate scenario, and compared to present (2001–2020) conditions. Our projections indicated that the copepod community is expected to respond substantially to climate change: a mean poleward latitudinal shift of 8.7 km per decade for the overall community with an important species range variation (–15 to 18 km per decade); the species seasonal peak is expected to occur 12–13 d earlier for Calanus finmarchicus and C. hyperboreus; and important changes in community structure are also expected (high species turnover of 43–79% south of the Oceanic Polar Front). The impacts of the change expected by the end of the century under IPCC global warming scenarios on copepods highlight poleward shifts, earlier seasonal peak and changes in biodiversity spatial patterns that might lead to alterations of the future North Atlantic pelagic ecosystem. Our model and projections are supported by a temporal validation undertaken using the North Atlantic climate regime shift that occurred in the 1980s: the habitat model built in the cold period (1970–1986) has been validated in the warm period (1987–2004).