Changes in marine dinoflagellate and diatom abundance under climate change

Marine diatoms and dinoflagellates play a variety of key ecosystem roles as important primary producers (diatoms and some dinoflagellates) and grazers (some dinoflagellates). Additionally some are harmful algal bloom (HAB) species and there is widespread concern that HAB species may be increasing accompanied by major negative socio-economic impacts, including threats to human health and marine harvesting1, 2. Using 92,263 samples from the Continuous Plankton Recorder survey, we generated a 50-year (1960–2009) time series of diatom and dinoflagellate occurrence in the northeast Atlantic and North Sea. Dinoflagellates, including both HAB taxa (for example, Prorocentrum spp.) and non-HAB taxa (for example, Ceratium furca), have declined in abundance, particularly since 2006. In contrast, diatom abundance has not shown this decline with some common diatoms, including both HAB (for example, Pseudo-nitzschia spp.) and non-HAB (for example, Thalassiosira spp.) taxa, increasing in abundance. Overall these changes have led to a marked increase in the relative abundance of diatoms versus dinoflagellates. Our analyses, including Granger tests to identify criteria of causality, indicate that this switch is driven by an interaction effect of both increasing sea surface temperatures combined with increasingly windy conditions in summer.

Long-term changes in abundance and distribution of microzooplankton in the NE Atlantic and North Sea

Long-term changes in mesozooplankton and phytoplankton populations have been well documented in the North Atlantic region, whereas data for microzooplankton are scarce. This neglected component of the plankton is a vital link in marine food-webs, grazing on smaller flagellates and cyanobacteria and in turn providing food for the larger mesozooplankton. We use the latest tintinnid (Ciliophora, Protista) data from the Continuous Plankton Recorder (CPR) survey in the NE Atlantic and North Sea to examine the phenology, distribution and abundance of this important group of ciliates. Presence/absence data came from 167 122 CPR samples collected between 1960 and 2009 and abundance data from 49 662 samples collected between 1996 and 2009. In the North Atlantic the genus Dictyocysta spp. dominated and Parafavella gigantea showed an increase in abundance around Iceland and Greenland. In the North Sea higher densities of Tintinnopsis spp., Favella serrata and Ptychocylis spp. were found. The presence of tintinnids in CPR samples collected in the North Atlantic has increased over the last 50 years and the seasonal window of high abundance has lengthened. Conversely in the North Sea there has been an overall reduction in abundance. We discuss possible drivers for these long-term changes and point the way forward to more holistic studies that examine how ecosystems, rather than just selected taxa, are responding to climate change.

Long-term changes in North Atlantic zooplankton biomass

New measures of zooplankton biomass have been derived from CPR samples in the North Atlantic from 1958 to 2005. The final aim was to investigate how the zooplankton standing stock had varied throughout the last decades, knowing that in different areas of the North Atlantic significant changes in the distribution of the dominant zooplankton species as well as the plankton assemblage have been observed. During the forty-five years of monitoring the contribution of the different groups (e.g. copepods, euphausiids, meroplankton larvae) to the total zooplankton biomass has been evaluated. The changes in the phenology of the biomass were also considered. The relationship between quantity, quality and seasonal timing of plankton and the poor fish recruitment seen in recent years in the North Sea are also discussed.

Long-term changes in the meroplankton of the North Sea

Data from the continuous plankton recorder (CPR) survey collected in the late-1940s to early-1960s indicated that the abundance of decapod larvae was low and the seasonal peak of abundance was late following cold winters. The phenological effect of temperature was shown to be consistent with relationships between both geographical and interannual patterns of variation. Analyses of CPR data collected from the 1940s to the present day reveal large-scale long-term changes in the abundance and phenology of the North Sea meroplankton. Echinoderm larvae, whose peak abundance has advanced by 47 days, show the greatest shift in timing. Echinoderm larvae have also increased in abundance to become the most abundant taxon in North Sea CPR samples. Genetic and morphological analyses of CPR samples show that the variations in echinoderm larvae are mainly attributable to an increasing abundance and earlier occurrence of the larvae of a resident species, Echinocardium cordatum, rather than a change in species composition. The remarkable scale of the changes in abundance and phenology of the meroplankton, which are greater than those seen in the holoplankton, has stimulated the development of further research into the causes and effects of these changes.

Climate-induced changes in the North Sea Decapoda over the last 60 years

In the mid-1980s the North Sea ecosystem experienced a climate-induced regime shift that has favoured decapods and detritivores in the benthos and jellyfish in the plankton over commercial fisheries. Here, we investigate changes among the Decapoda in the North Sea plankton over the last 60 yr. Decapods are important predators in the plankton and the benthos where they can influence productivity and structure communities. In the North Sea it has been suggested that a climate-driven increase in decapod abundance has been important in propagating the climate signal through the North Sea food web. We show that climate-induced changes in the Decapoda in the central and southern North Sea include the presence of new warm-water taxa, changes in the abundance and proportions of commercial species of shrimp, and an earlier occurrence of decapod larvae in the plankton compared with the period 1981–1983. Notable amongst the warm-water taxa appearing in the North Sea is the predatory swimming crab Polybius henslowii that can swarm in large numbers when conditions are favourable and that is known to exhibit range shifts in response to fluctuations in hydroclimatic forcing. We suggest that climate-induced changes among North Sea decapods have played an important role in the trophic amplification of a climate signal and the development of the new North Sea dynamic regime. Understanding these changes is likely to be imperative for a successful ecosystem-based approach to the future management of North Sea fisheries at a time of climate change.

Empirically modelling the potential effects of changes in temperature and prey availability on the growth of cod larvae in UK shelf seas

It has been hypothesized that changes in zooplankton community structure over the past four decades led to reduced growth and survival of prerecruit Atlantic cod (Gadus morhua) and that this was a key factor underlying poor year classes, contributing to stock collapse, and inhibiting the recovery of stocks around the UK. To evaluate whether observed changes in plankton abundance, species composition and temperature could have led to periods of poorer growth of cod larvae, we explored the effect of prey availability and temperature on early larval growth using an empirical trophodynamic model. Prey availability was parameterized using species abundance data from the Continuous Plankton Recorder. Our model suggests that the observed changes in plankton community structure in the North Sea may have had less impact on cod larval growth, at least for the first 40 days following hatching, than previously suggested. At least in the short term, environmental and prey conditions should be able to sustain growth of cod larvae and environmental changes acting on this early life stage should not limit stock recovery.

Have climate induced changes in zooplankton communities led to poor conditions for larval fish growth? A test on cod larvae on the UK shelf

Climate effects have been shown to be at least partly responsible for the reorganisation in the plankton ecosystem on the shelf seas of NW Europe over the last 50 years. Most fish larvae feed primarily on zooplankton, so changes in zooplankton quantity, quality and seasonal timing have been hypothesized to be a key factor affecting their survival. To investigate this we have implemented a 1-dimensional trophodynamic growth model of cod larvae for the waters around the UK covering the period 1960 to 2003. Larval growth is modelled as the difference between the amount of food absorbed by the larva and its various metabolic costs. Prey availability is based upon the biomass and size of available preys (i.e. adults and nauplii copepods and cladocerans) taken from the Continuous Plankton Recorder dataset. Temperature and wind forcing are also taken into account. Results suggest that observed changes in plankton community structure may have had less impact than previously suggested. This is because changes in prey availability may be compensated for by increased temperatures resulting in little overall impact on potential larval growth. Stock recovery, at least in the short term is likely to be more dependent upon conserving the year classes recruited to allow spawning stock biomass to rebuild. If as our model suggests, the larvae are still able to survive in the changing environment, reduction in fishing on the adults is needed to allow the stock to recover.

Region-wide changes in marine ecosystem dynamics: state-space models to distinguish trends from step changes

Regime shifts are sudden changes in ecosystem structure that can be detected across several ecosystem components. The concept that regime shifts are common in marine ecosystems has gained popularity in recent years. Many studies have searched for the step-like changes in ecosystem state expected under a simple interpretation of this idea. However, other kinds of change, such as pervasive trends, have often been ignored. We assembled over 300 ecological time series from seven UK marine regions, covering two to three decades. We developed state-space models for the first principal component of the time series in each region, a common measure of ecosystem state. Our models allowed both trends and step changes, possibly in combination. We found trends in three of seven regions and step changes in two of seven regions. Gradual and sudden changes are therefore important trajectories to consider in marine ecosystems.

A methodology for the assessment of local-scale changes in marine environmental benefits and its application

Local-scale planning decisions are required by the existing Environmental Impact Assessment process to take account of the implications of a development on a range of environmental and social factors, and could therefore be supported by an ecosystem services approach. However, empirical assessments at a local scale within the marine environment have focused on only a single or limited set of services. This paper tests the applicability of the ecosystem services approach to environmental impact appraisal by considering how the identification and quantification of a comprehensive suite of benefits provided at a local scale might proceed in practice. A methodology for conducting an Environmental Benefits Assessment (EBA) is proposed, the underlying framework for which follows the recent literature by placing the emphasis on ecosystem benefits, as opposed to services. The EBA methodology also proposes metrics that can be quantified at local scale, and is tested using a case study of a hypothetical tidal barrage development in the Taw Torridge estuary in North Devon, UK. By suggesting some practical steps for assessing environmental benefits, this study aims to stimulate discussion and so advance the development of methods for implementing ecosystem service approaches at a local scale.

Vulnerability of coastal ecosystems to changes in harmful algal bloom distribution in response to climate change: projections based on model analysis

Harmful algal blooms (HABs), those proliferations of algae that can cause fish kills, contaminate seafood with toxins, form unsightly scums, or detrimentally alter ecosystem function have been increasing in frequency, magnitude, and duration worldwide. Here, using a global modeling approach, we show, for three regions of the globe, the potential effects of nutrient loading and climate change for two HAB genera, pelagic Prorocentrum and Karenia, each with differing physiological characteristics for growth. The projections (end of century, 2090-2100) are based on climate change resulting from the A1B scenario of the Intergovernmental Panel on Climate Change Institut Pierre Simon Laplace Climate Model (IPCC, IPSL-CM4), applied in a coupled oceanographic-biogeochemical model, combined with a suite of assumed physiological 'rules' for genera-specific bloom development. Based on these models, an expansion in area and/or number of months annually conducive to development of these HABs along the NW European Shelf-Baltic Sea system and NE Asia was projected for both HAB genera, but no expansion (Prorocentrum spp.), or actual contraction in area and months conducive for blooms (Karenia spp.), was projected in the SE Asian domain. The implications of these projections, especially for Northern Europe, are shifts in vulnerability of coastal systems to HAB events, increased regional HAB impacts to aquaculture, increased risks to human health and ecosystems, and economic consequences of these events due to losses to fisheries and ecosystem services.

Variation in elemental stoichiometry of the marine diatomThalassiosira weissflogii(Bacillariophyceae) in response to combined nutrient stress and changes in carbonate chemistry

The combined consequences of the multi-stressors of pH and nutrient availability upon the growth of a marine diatom were investigated. Thalassiosira weissflogii was grown in N- or P-limited batch culture in sealed systems, with pH commencing at 8.2 (extant conditions) or 7.6 (ocean acidification [OA] conditions), and then pH was allowed to either drift with growth, or was held fixed. Results indicated that within the pH range tested, the stability of environmental pH rather than its value (i.e., OA vs. extant) fundamentally influenced biomass accumul-ation and C:N:P stoichiometry. Despite large changes in total alkalinity in the fixed pH systems, final biomass production was consistently greater in these systems than that in drifting pH systems. In drift systems, pH increased to exceed pH 9.5, a level of alkalinity that was inhibitory to growth. No statis-tically significant differences between pH treatments were measured for N:C, P:C or N:P ratios during nutrient-replete growth, although the diatom expre-ssed greater plasticity in P:C and N:P ratios than in N:C during this growth phase. During nutrient-deplete conditions, the capacity for uncoupled carbon fixa-tion at fixed pH was considerably greater than that measured in drift pH systems, leading to strong contrasts in C:N:P stoichiometry between these treatments. Whether environmental pH was stable or drifted directly influenced the extent of physiological stress. In contrast, few distinctions could be drawn between extant versus OA conditions for cell physiology.