Modeling marine ecosystem responses to global climate change: Where are we now and where should we be going?

Modeling of global climate change is moving from global circulation model (GCM)-type projections with coupled biogeochemical models to projections of ecological responses, including food web and upper trophic levels. Marine and coastal ecosystems are highly susceptible to the impacts of global climate change and also produce significant ecosystem services. The effects of global climate change on coastal and marine ecosystems involve a much wider array of effects than the usual temperature, sea level rise, and precipitation. This paper is an overview for a collection of 12 papers that examined various aspects of global climate change on marine ecosystems and comprise this special issue. We summarized the major features of the models and analyses in the papers to determine general patterns. A wide range of ecosystems were simulated using a diverse set of modeling approaches. Models were either 3-dimensional or used a few spatial boxes, and responses to global climate change were mostly expressed as changes from a baseline condition. Three issues were identified from the across-model comparison: (a) lack of standardization of climate change scenarios, (b) the prevalence of site-specific and even unique models for upper trophic levels, and (c) emphasis on hypothesis evaluation versus forecasting. We discuss why these issues are important as global climate change assessment continues to progress up the food chain, and, when possible, offer some initial steps for going forward.

Understanding the Mechanisms of Cell and Population Responses of Coccolithophores to Changing Ocean Chemistry

The calcifying coccolithophores have been proposed as a potentially vulnerable group in the face of increasing surface ocean CO2 levels. A full understanding of the likely responses of this group requires better mechanistic information on pH- and CO2-sensitive processes that underlie cell function at molecular, cellular and population levels. New findings on the mechanisms of pH homeostasis at a molecular and cellular level in both diatoms and coccolithophores are shaping our understanding of how these important groups may respond or acclimate to changing ocean pH. Critical parameters including intracellular pH homeostasis and cell surface pH will be considered. These studies are being carried out in parallel with genetic studies of natural oceanic populations to assess the natural genetic and physiological diversity that will underlie adaptation of populations in the long term.

Long-term responses of North Atlantic calcifying plankton to climate change

The global increase in atmospheric carbon dioxide concentration is potentially threatening marine biodiversity in two ways. First, carbon dioxide and other greenhouse gases accumulating in the atmosphere are causing global warming1. Second, carbon dioxide is altering sea water chemistry, making the ocean more acidic2. Although temperature has a cardinal influence on all biological processes from the molecular to the ecosystem level3, acidification might impair the process of calcification or exacerbate dissolution of calcifying organisms4. Here, we show however that North Atlantic calcifying plankton primarily responded to climate-induced changes in temperatures during the period 1960–2009, overriding the signal from the effects of ocean acidification. We provide evidence that foraminifers, coccolithophores, both pteropod and nonpteropod molluscs and echinoderms exhibited an abrupt shift circa 1996 at a time of a substantial increase in temperature5 and that some taxa exhibited a poleward movement in agreement with expected biogeographical changes under sea temperature warming6,7. Although acidification may become a serious threat to marine calcifying organisms, our results suggest that over the study period the primary driver of North Atlantic calcifying planktonwas oceanic temperature.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

Short-term responses of the cold water coral Lophelia pertusa to predicted rises in atmospheric CO2

Cold-water corals are associated with high local biodiversity, but despite their importance as ecosystem engineers, little is known about how these organisms will respond to projected ocean acidification. Since preindustrial times, average ocean pH has decreased from 8.2 to ~8.1, and predicted CO2 emissions will decrease by up to another 0.3 pH units by the end of the century. This decrease in pH may have a wide range of impacts upon marine life, and in particular upon calcifiers such as cold-water corals. Lophelia pertusa is the most widespread cold-water coral (CWC) species, frequently found in the North Atlantic. Here, we present the first short-term (21 days) data on the effects of increased CO2 (750 ppm) upon the metabolism of freshly collected L. pertusa from Mingulay Reef Complex, Scotland, for comparison with net calcification. Over 21 days, corals exposed to increased CO2 conditions had significantly lower respiration rates (11.4±1.39 SE, µmol O2 g−1 tissue dry weight h−1) than corals in control conditions (28.6±7.30 SE µmol O2 g−1 tissue dry weight h−1). There was no corresponding change in calcification rates between treatments, measured using the alkalinity anomaly technique and 14C uptake. The decrease in respiration rate and maintenance of calcification rate indicates an energetic imbalance, likely facilitated by utilisation of lipid reserves. These data from freshly collected L. pertusa from the Mingulay Reef Complex will help define the impact of ocean acidification upon the growth, physiology and structural integrity of this key reef framework forming species.

Strong responses of Southern Ocean phytoplankton communities to volcanic ash

Volcanic eruptions have been hypothesized as an iron supply mechanism for phytoplankton blooms; however, little direct evidence of stimulatory responses has been obtained in the field. Here we present the results of twenty-one 1–2 day bottle enrichment experiments from cruises in the South Atlantic and Southern Ocean which conclusively demonstrated a photophysiological and biomass stimulation of phytoplankton communities following supply of basaltic or rhyolitic volcanic ash. Furthermore, experiments in the Southern Ocean demonstrated significant phytoplankton community responses to volcanic ash supply in the absence of responses to addition of dissolved iron alone. At these sites, dissolved manganese concentrations were among the lowest ever measured in seawater, and we therefore suggest that the enhanced response to ash may have been a result of the relief of manganese (co)limitation. Our results imply that volcanic ash deposition events could trigger extensive phytoplankton blooms, potentially capable of significant impacts on regional carbon cycling.

Quantifying ecosystem-level consequences of ocean acidification and warming by scaling up individual level responses of a predator snail and its trophic interactions

Understanding long‐term, ecosystem‐level impacts of climate change is challenging because experimental research frequently focuses on short‐term, individual‐level impacts in isolation. We address this shortcoming first through an interdisciplinary ensemble of novel experimental techniques to investigate the impacts of 14‐month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterize a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual‐level responses, while acidification had a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual‐level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large‐scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local‐environmental conditions and resource availability. Such changes in macro‐scale distributions cannot be predicted by investigating individual‐level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long‐term, multiscale responses to multiple stressors, in an ecosystem context.

Multiple environmental controls on phytoplankton growth strategies determine adaptive responses of the N : P ratio

The controls on the 'Redfield' N:P stoichiometry of marine phytoplankton and hence the N:P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource-allocation trade-offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per-ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N:P ratios and N:P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.

Volcanic ash supply to the surface ocean – remote sensing of biological responses and their wider biogeochemical significance

Transient micronutrient enrichment of the surface ocean can enhance phytoplankton growth rates and alter microbial community structure with an ensuing spectrum of biogeochemical feedbacks. Strong phytoplankton responses to micronutrients supplied by volcanic ash have been reported recently. Here we: (i) synthesize findings from these recent studies; (ii) report the results of a new remote sensing study of ash fertilization; and (iii) calculate theoretical bounds of ash-fertilized carbon export. Our synthesis highlights that phytoplankton responses to ash do not always simply mimic that of iron amendment; the exact mechanisms for this are likely biogeochemically important but are not yet well understood. Inherent optical properties of ash-loaded seawater suggest rhyolitic ash biases routine satellite chlorophyll-a estimation upwards by more than an order of magnitude for waters with <0.1 mg chlorophyll-a m-3, and less than a factor of 2 for systems with >0.5 mg chlorophyll-a m-3. For this reason post-ash-deposition chlorophyll-a changes in oligotrophic waters detected via standard Case 1 (open ocean) algorithms should be interpreted with caution. Remote sensing analysis of historic events with a bias less than a factor of 2 provided limited stand-alone evidence for ash-fertilization. Confounding factors were poor coverage, incoherent ash dispersal, and ambiguity ascribing biomass changes to ash supply over other potential drivers. Using current estimates of iron release and carbon export efficiencies, uncertainty bounds of ash-fertilized carbon export for 3 events are presented. Patagonian iron supply to the Southern Ocean from volcanic eruptions is less than that of windblown dust on thousand year timescales but can dominate supply at shorter timescales. Reducing uncertainties in remote sensing of phytoplankton response and nutrient release from ash are avenues for enabling assessment of the oceanic response to large-scale transient nutrient enrichment.

Are Fast Responses More Random? Testing the Effect of Response Time on Scale in an Online Choice Experiment

Scepticism over stated preference surveys conducted online revolves around the concerns over “professional respondents” who might rush through the questionnaire without sufficiently considering the information provided. To gain insight on the validity of this phenomenon and test the effect of response time on choice randomness, this study makes use of a recently conducted choice experiment survey on ecological and amenity effects of an offshore windfarm in the UK. The positive relationship between self-rated and inferred attribute attendance and response time is taken as evidence for a link between response time and cognitive effort. Subsequently, the generalised multinomial logit model is employed to test the effect of response time on scale, which indicates the weight of the deterministic relative to the error component in the random utility model. Results show that longer response time increases scale, i.e. decreases choice randomness. This positive scale effect of response time is further found to be non-linear and wear off at some point beyond which extreme response time decreases scale. While response time does not systematically affect welfare estimates, higher response time increases the precision of such estimates. These effects persist when self-reported choice certainty is controlled for. Implications of the results for online stated preference surveys and further research are discussed.

Responses in Arctic marine carbon cycle processes: conceptual scenarios & implications for ecosystem.

The Arctic Ocean is one of the fastest changing oceans, plays an important role in global carbon cycling and yet is a particularly challenging ocean to study. Hence, observations tend to be relatively sparse in both space and time. How the Arctic functions, geophysically, but also ecologically, can have significant consequences for the internal cycling of carbon, and subsequently influence carbon export, atmospheric CO2 uptake and food chain productivity. Here we assess the major carbon pools and associated processes, specifically summarizing the current knowledge of each of these processes in terms of data availability and ranges of rates and values for four geophysical Arctic Ocean domains originally described by Carmack & Wassmann (2006): inflow shelves, which are Pacific-influenced and Atlantic-influenced; interior, river-influenced shelves; and central basins. We attempt to bring together knowledge of the carbon cycle with the ecosystem within each of these different geophysical settings, in order to provide specialist information in a holistic context. We assess the current state of models and how they can be improved and/or used to provide assessments of the current and future functioning when observational data are limited or sparse. In doing so, we highlight potential links in the physical oceanographic regime, primary production and the flow of carbon within the ecosystem that will change in the future. Finally, we are able to highlight priority areas for research, taking a holistic pan-Arctic approach.