Claire E. Widdicombe

Data integration for European marine biodiversity research: creating a database on benthos and plankton to study large-scale patterns and long-term changes.

The general aim of setting up a central database on benthos and plankton was to integrate long-, medium- and short-term datasets on marine biodiversity. Such a database makes it possible to analyse species assemblages and their changes on spatial and temporal scales across Europe. Data collation lasted from early 2007 until August 2008, during which 67 datasets were collected covering three divergent habitats (rocky shores, soft bottoms and the pelagic environment). The database contains a total of 4,525 distinct taxa, 17,117 unique sampling locations and over 45,500 collected samples, representing almost 542,000 distribution records. The database geographically covers the North Sea (221,452 distribution records), the North-East Atlantic (98,796 distribution records) and furthermore the Baltic Sea, the Arctic and the Mediterranean. Data from 1858 to 2008 are presented in the database, with the longest time-series from the Baltic Sea soft bottom benthos. Each delivered dataset was subjected to certain quality control procedures, especially on the level of taxonomy. The standardisation procedure enables pan-European analyses without the hazard of taxonomic artefacts resulting from different determination skills. A case study on rocky shore and pelagic data in different geographical regions shows a general overestimation of biodiversity when making use of data before quality control compared to the same estimations after quality control. These results prove that the contribution of a misspelled name or the use of an obsolete synonym is comparable to the introduction of a rare species, having adverse effects on further diversity calculations. The quality checked data source is now ready to test geographical and temporal hypotheses on a large scale.

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.

Fine-scale variability in phytoplankton community structure and inherent optical properties measured from an autonomous underwater vehicle

The relationship between inherent optical properties (IOPs), phytoplankton community structure and the abundance of suspended particles in the size range 3–500 Am was studied near the Isles of Scilly (UK) in May 2000. Autosub, an autonomous submersible vehicle specifically designed for science missions, was used as an instrument-positioning platform. It carried a CTD system, an ac-9+ dual tube spectrophotometer, a prototype submersible flow cytometer and an Aqua-monitor water sampler. The vehicle made a 10-km transect at constant depth across a boundary between water masses with contrasting remote sensing reflectance, which was located using SeaWiFs ocean colour imagery. This boundary corresponded to a sharp (1 km) transition between one phytoplankton community consisting of coccolithophores, flagellates and dinoflagellates, and a second community dominated by diatoms and flagellates. Inherent optical properties measured along the autonomous underwater vehicle (AUV) track showed marked changes in magnitudes, ratios, spectral shapes and fine-scale spatial variability. These changes were well correlated with variations in the composition of the suspended particle assemblage measured by microscopy and in situ flow cytometry. D 2003 Elsevier B.V. All rights reserved.

The influence of balanced and imbalanced resource supply on biodiversity–functioning relationship across ecosystems

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity–ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity–ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity–productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity–functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.

Ecological equivalence of species within phytoplankton functional groups

1.There are tens of thousands of species of phytoplankton found throughout the tree of life. Despite this diversity, phytoplankton are often aggregated into a few functional groups according to metabolic traits or biogeochemical role. We investigate the extent to which phytoplankton species dynamics are neutral within functional groups. 2.Seasonal dynamics in many regions of the ocean are known to affect phytoplankton at the functional group level leading to largely predictable patterns of seasonal succession. It is much more difficult to make general statements about the dynamics of individual species. 3.We use a 7 year time-series at station L4 in the Western English Channel with 57 diatom and 17 dinoflagellate species enumerated weekly to test if the abundance of diatom and dinoflagellate species vary randomly within their functional group envelope or if each species is driven uniquely by external factors. 4.We show that the total biomass of the diatom and dinoflagellate functional groups is well predicted by irradiance and temperature and quantify trait values governing the growth rate of both functional groups. The biomass dynamics of the functional groups are not neutral and each has their own distinct responses to environmental forcing. Compared to dinoflagellates, diatoms have faster growth rates, and grow faster under lower irradiance, cooler temperatures, and higher nutrient conditions. 5.The biomass of most species vary randomly within their functional group biomass envelope, most of the time. As a consequence, modelers will find it difficult to predict the biomass of most individual species. Our analysis supports the approach of using a single set of traits for a functional group and suggests that it should be possible to determine these traits from natural communities.

Metabolically active, non-nitrogen fixing, Trichodesmium in UK coastal waters during winter

Trichodesmium, a colonial cyanobacterium typically associated with tropical waters, was observed between January and April 2014 in the western English Channel. Sequencing of the heterocyst differentiation (hetR) and 16S rRNA genes placed this community within the Clade IV Trichodesmium, an understudied clade previously found only in low numbers in warmer waters. Nitrogen fixation was not detected although measurable rates of nitrate uptake and carbon fixation were observed. Trichodesmium RuBisCO transcript abundance relative to gene abundance suggests the potential for viable and potentially active Trichodesmium carbon fixation. Observations of Trichodesmium when coupled with a numerical advection model indicate that Trichodesmium communities can remain viable for >3.5 months at temperatures lower than previously expected. The results suggest that Clade IV Trichodesmium occupies a different niche to other Trichodesmium species, and is a cold- or low-light-adapted variant.

Effects of elevated CO2 on phytoplankton community biomass and species composition during a spring Phaeocystis spp. bloom in the western English Channel

A 21-year time series of phytoplankton community structure was analysed in relation to Phaeocystis spp. to elucidate its contribution to the annual carbon budget at station L4 in the western English Channel (WEC). Between 1993-2014 Phaeocystis spp. contributed ∼4.6% of the annual phytoplankton carbon and during the March - May spring bloom, the mean Phaeocystis spp. biomass constituted 17% with a maximal contribution of 47% in 2001. Upper maximal weekly values above the time series mean ranged from 63 to 82% of the total phytoplankton carbon (∼42-137mg carbon (C)m-3) with significant inter-annual variability in Phaeocystis spp. Maximal biomass usually occurred by the end of April, although in some cases as early as mid-April (2007) and as late as late May (2013). The effects of elevated pCO2 on the Phaeocystis spp. spring bloom were investigated during a fifteen-day semi-continuous microcosm experiment. The phytoplankton community biomass was estimated at ∼160mgCm-3 and was dominated by nanophytoplankton (40%, excluding Phaeocystis spp.), Phaeocystis spp. (30%) and cryptophytes (12%). The smaller fraction of the community biomass comprised picophytoplankton (9%), coccolithophores (3%), Synechococcus (3%), dinoflagellates (1.5%), ciliates (1%) and diatoms (0.5%). Over the experimental period, total biomass increased significantly by 90% to ∼305mgCm-3 in the high CO2 treatment while the ambient pCO2 control showed no net gains. Phaeocystis spp. exhibited the greatest response to the high CO2 treatment, increasing by 330%, from ∼50mgCm-3 to over 200mgCm-3 and contributing ∼70% of the total biomass. Taken together, the results of our microcosm experiment and analysis of the time series suggest that a future high CO2 scenario may favour dominance of Phaeocystis spp. during the spring bloom. This has significant implications for the formation of hypoxic zones and the alteration of food web structure including inhibitory feeding effects and lowered fecundity in many copepod species.

Seasonality of Oithona similis and Calanus helgolandicus reproduction and abundance: contrasting responses to environmental variation at a shelf site

The pelagic copepods Oithona similis and Calanus helgolandicus have overlapping geographic ranges, yet contrast in feeding mode, reproductive strategy, and body size. We investigate how these contrasting traits influence the seasonality of copepod abundance and reproductive output under environmental variation, using time series data collected over 25 years at the Western Channel Observatory station L4. The proportional change in Egg Production Rate (EPR, eggs female-1 d-1) over the annual cycle was ~10-fold and similar for both species, although EPR of O. similis was only ~ 11% that of C. helgolandicus. The timing of EPR maxima for O. similis coincided with increased Sea Surface Temperature (SST) in summer, likely due to a temperature-dependent brooding period. Conversely, EPR of broadcast spawning C. helgolandicus was more strongly related to Net Heat Flux (NHF) and diatom biomass, both parameters associated with the spring phytoplankton bloom. In both species, female body mass negatively correlated with SST, with a 7.5% reduction in body mass per °C in C. helgolandicus compared to just 2.3% in O. similis. Finally, seasonality of EPR and adult and copepodite abundance was strongly decoupled in both species, suggesting that optimum conditions for reproduction and abundance occur at different times of the year.

Long-term dynamics in a soft-bottom amphipod community and the influence of the pelagic environment

The processes and patterns seen in coastal benthic communities can be strongly influenced by the overlying pelagic environmental conditions. Integrating long-term biological and environmental data (both benthic and pelagic) can give insight into the specific relationships between key benthic functional groups and natural temporal changes in the marine environment. The identity and abundance of amphipod species found at Station L4 (Western English Channel) were tracked for 7 years (2008-2014), whilst simultaneously, annual changes in phytoplankton biomass, water temperature, salinity and chlorophyll a concentration were also characterized. The main species were persistent and showed little variability along the study period. Overall, however, there were significant changes in the structure of the whole community between sampling times, highlighting the importance of less numerically-dominant species in driving temporal variability. Surprisingly, the current study did not detect a significant influence of the phytoplankton biomass on benthic amphipod dynamics. On the other hand, there was a clear and constant correlation between bottom water temperatures and amphipod abundance. This pattern is different from that observed in other detritivorous species at L4, highlighting the complexity of benthic-pelagic coupling and the high variability of the response to pelagic conditions among different groups. As a result of the biogeographic position of the Western English Channel, the key role of amphipods in benthic communities, the influence of the temperature in their populations dynamics, as well as the solid baseline provided here and in previous studies, the monitoring of long-term amphipod dynamics in the English Channel could be a valuable tool to evaluate the biological effect of climate change over marine benthic communities.

Effects of elevated CO 2 and temperature on phytoplankton community biomass, species composition and photosynthesis during an autumn bloom in the Western English Channel

The combined effects of elevated pCO 2 and temperature were investigated during an autumn phytoplankton bloom in the Western English Channel (WEC). A full factorial 36-day microcosm experiment was conducted under year 2100 predicted temperature (+4.5 °C) and pCO 2 levels (800 μatm). The starting phytoplankton community biomass was 110.2 (±5.7 sd) mg carbon (C) m −3 and was dominated by dinoflagellates (~ 50 %) with smaller contributions from nanophytoplankton (~ 13 %), cryptophytes (~ 11 %)and diatoms (~ 9 %). Over the experimental period total biomass was significantly increased by elevated pCO 2 (20-fold increase) and elevated temperature (15-fold increase). In contrast, the combined influence of these two factors had little effect on biomass relative to the ambient control. The phytoplankton community structure shifted from dinoflagellates to nanophytoplankton at the end of the experiment in all treatments. Under elevated pCO 2 nanophytoplankton contributed 90% of community biomass and was dominated by Phaeocystis spp., while under elevated temperature nanophytoplankton contributed 85 % of the community biomass and was dominated by smaller nano-flagellates. Under ambient conditions larger nano-flagellates dominated while the smallest nanophytoplankton contribution was observed under combined elevated pCO 2 and temperature (~ 40 %). Dinoflagellate biomass declined significantly under the individual influences of elevated pCO 2 , temperature and ambient conditions. Under the combined effects of elevated pCO 2 and temperature, dinoflagellate biomass almost doubled from the starting biomass and there was a 30-fold increase in the harmful algal bloom (HAB) species, Prorocentrum cordatum . Chlorophyll a normalised maximum photosynthetic rates (P B m ) increased > 6-fold under elevated pCO 2 and > 3-fold under elevated temperature while no effect on P B m was observed when pCO 2 and temperature were elevated simultaneously. The results suggest that future increases in temperature and pCO 2 do not appear to influence coastal phytoplankton productivity during autumn in the WEC which would have a negative feedback on atmospheric CO 2 .