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.

Nitrogen uptake of phytoplankton assemblages under contrasting upwelling and downwelling conditions: The Ría de Vigo, NW Iberia

Regenerated production (including organic nitrogen) is shown here to be important in the Ria de Vigo (Galicia, NW Iberia) in supporting both harmful algal bloom communities during the downwelling season, but also (to a lesser extent) diatom communities during stratified periods of weak to moderate upwelling. The Galician Rias, situated in the Iberian upwelling system, are regularly affected by blooms of toxic dinoflagellates, which pose serious threats to the local mussel farming industry. These tend to occur towards the end of summer, during the transition from upwelling to downwelling favourable seasons, when cold bottom shelf waters in the rias are replaced by warm surface shelf waters. Nitrate, ammonium and urea uptake rates were measured in the Ria de Vigo during a downwelling event in September 2006 and during an upwelling event in June 2007. In September the ria was well mixed, with a downwelling front observed towards the middle of the ria and relatively high nutrient concentrations (1.0-2.6 mu mol L-1 nitrate; 1.0-5.6 mu mol L-1 ammonium; 0.1-0.8 mu mol L-1 phosphate; 2.0-9.0 mu mol L-1 silicic acid) were present throughout the water column. Ammonium represented more than 80% of the nitrogenous nutrients, and the phytoplankton assemblage was dominated by dinoflagellates and small flagellates. In June the water column was stratified, with nutrient-rich, upwelled water below the thermocline and warm, nutrient-depleted water in the surface. At this time, nitrate represented more than 80% of the nitrogenous nutrients, and a mixed diatom assemblage was present. Primary phytoplankton production during both events was mainly sustained by regenerated nitrogen, with ammonium uptake rates of 0.035-0.063 mu mol N L-1 h(-1) in September and 0.078-0.188 mu mol N L-1 h(-1) in June. Although f-ratios were generally low (<0.2) in both June and September, a maximum of 0.61 was reached in June due to higher nitrate uptake (0.225 mu mol N L-1 h(-1)). Total nitrogen uptake was also higher during the upwelling event (0.153-0.366 in June and 0.053-0.096 mu mol N L-1 h(-1) in September). Nitrogen uptake kinetics demonstrated a strong preference for ammonium and urea over nitrate in June.

Coccolithophore species as indicators of surface oceanographic conditions in the vicinity of Azores islands

During summer 2008 and spring 2009, surface oceanographic surveys were carried out around three islands of the Azores archipelago (Terceira, Sao Miguel and Santa Maria) to assess the phytoplankton distribution and associated physico-chemical processes. The Azores archipelago is a major feature in the biogeochemical North Atlantic Subtropical Gyre (NAST) province although its influence on the productivity of the surrounding ocean is poorly known. Surface phytoplankton was studied by microscopy and HPLC (High Precision Liquid Chromatography). The mean values for biomass proxy Chlorophyll a (Chla) ranged from 0.04 to 0.55 mu g L-1 (Chla maximum = 0.86 mu g L-1) and coccolithophores were the most abundant group, followed by small flagellates, Cyanobacteria, diatoms and dinoflagellates being the least abundant group. The distribution of phytoplankton and coccolithophore species in particular presented seasonal differences and was consistent with the nearshore influence of warm subtropical waters from the south Azores current and colder subpolar waters from the north. The satellite-derived circulation patterns showed southward cold water intrusions off Terceira and northward warm water intrusions off Santa Maria. The warmer waters signal was confirmed by the subtropical coccolithophore assemblage, being Discosphaera tubifera a constant presence under these conditions. The regions of enhanced biomass, either resulting from northern cooler waters or from island induced processes, were characterized by the presence of Emiliania huxleyi. Diatoms and dinoflagellates indicated coastal and regional processes of nutrient enrichment and areas of physical stability, respectively.

Past, present and future nutrient loads of the North Sea: causes and consequences

Comprehensive, aggregate nutrient budgets were established for two compartments of the North Sea, the shallow coastal and deeper open regions, and for three different periods, representing pre-eutrophication (∼1950), eutrophication (∼1990) and contemporary (∼2000) phases. The aim was to quantify the major budget components, to identify their sources of variability, to specify the anthropogenic components, and to draw implications for past and future policy. For all three periods, open North Sea budgets were dominated (75%) by fluxes from and to the North-East Atlantic; sediment exchange was of secondary importance (18%). For the coastal North Sea, fluxes during the eutrophication period were dominated by sediment exchange (49% of all inputs), followed by exchange with the open sea (21%), and anthropogenic inputs (19%). Between 1950 and 1990, N-loading of coastal waters increased by a factor of 1.62 and P-loading by 1.45. These loads declined after 1990. Interannual variability in Atlantic inflow was found to correspond to a variability of 11% in nutrient load to the open North Sea. Area-specific external loads of both the open and coastal North Sea were below Vollenweider-type critical loads when expressed relative to depth and flushing. External area-specific load of the coastal North Sea has declined since 1990 from 1.8 to about 1.4 g P m−2 y−1 in 2000, which is close to the estimate of 1.3 for 1950. N-load declined less, leading to an increase in N/P ratio.

Marine biodiversity and ecosystem function relationships:the potential for practical monitoring applications

There is an increasing demand for environmental assessments of the marine environment to include ecosystem function. However, existing schemes are predominantly based on taxonomic (i.e. structural) measures of biodiversity. Biodiversity and Ecosystem Function (BEF) relationships are suggested to provide a mechanism for converting taxonomic information into surrogates of ecosystem function. This review assesses the evidence for marine BEF relationships and their potential to be used in practical monitoring applications (i.e. operationalized). Five key requirements were identified for the practical application of BEF relationships: (1) a complete understanding of strength, direction and prevalence of marine BEF relationships, (2) an understanding of which biological components are influential within specific BEF relationships, (3) the biodiversity of the selected biological components can be measured easily, (4) the ecological mechanisms that are the most important for generating marine BEF relationships, i.e. identity effects or complementarity, are known and (5) the proportion of the overall functional variance is explained by biodiversity, and hence BEF relationships, has been established. Numerous positive and some negative BEF relationships were found within the literature, although many reproduced poorly the natural species richness, trophic structures or multiple functions of real ecosystems (requirement 1). Null relationships were also reported. The consistency of the positive and negative relationships was often low that compromised the ability to generalize BEF relationships and confident application of BEF within marine monitoring. Equally, some biological components and functions have received little or no investigation. Expert judgement was used to attribute biological components using spatial extent, presence and functional rate criteria (requirement 2). This approach highlighted the main biological components contributing the most to specific ecosystem functions, and that many of the particularly influential components were found to have received the least amount of research attention. The need for biodiversity to be measureable (requirement 3) is possible for most biological components although difficult within the functionally important microbes. Identity effects underpinned most marine BEF relationships (requirement 4). As such, processes that translated structural biodiversity measures into functional diversity were found to generate better BEF relationships. The analysis of the contribution made by biodiversity, over abiotic influences, to the total expression of a particular ecosystem function was rarely measured or considered (requirement 5). Hence it is not possible to determine the overall importance of BEF relationships within the total ecosystem functioning observed. In the few studies where abiotic factors had been considered, it was clear that these modified BEF relationships and have their own direct influence on functional rate. Based on the five requirements, the information required for immediate ‘operationalization’ of BEF relationships within marine functional monitoring is lacking. However, the concept of BEF inclusion within practical monitoring applications, supported by ecological modelling, shows promise for providing surrogate indicators of functioning.

Tillin, H. & Tyler-Walters, H., 2014. Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with marine activities. Phase 2 Report – Literature review and sensitivity assessments for ecological groups for circalittoral and offshore Level 5 biotopes. JNCC Report No. 512B, 260 pp.

Human activities within the marine environment give rise to a number of pressures on seabed habitats. Improved understanding of the sensitivity of subtidal sedimentary habitats is required to underpin the management advice provided for Marine Protected Areas, as well as supporting other UK marine monitoring and assessment work. The sensitivity of marine sedimentary habitats to a range of pressures induced by human activities has previously been systematically assessed using approaches based on expert judgement for Defra Project MB0102 (Tillin et al. 2010). This previous work assessed sensitivity at the level of the broadscale habitat and therefore the scores were typically expressed as a range due to underlying variation in the sensitivity of the constituent biotopes. The objective of this project was to reduce the uncertainty around identifying the sensitivity of selected subtidal sedimentary habitats by assessing sensitivity, at a finer scale and incorporating information on the biological assemblage, for 33 Level 5 circalittoral and offshore biotopes taken from the Marine Habitat Classification of Britain and Ireland (Connor et al. 2004). Two Level 6 sub-biotopes were also included in this project as these contain distinctive characterising species that differentiate them from the Level 5 parent biotope. Littoral, infralittoral, reduced and variable salinity sedimentary habitats were excluded from this project as the scope was set for assessment of circalittoral and offshore sedimentary communities. This project consisted of three Phases. • Phase 1 - define ecological groups based on similarities in the sensitivity of characterising species from the Level 5 and two Level 6 biotopes described above. • Phase 2 - produce a literature review of information on the resilience and resistance of characterising species of the ecological groups to pressures associated with activities in the marine environment. • Phase 3 - to produce sensitivity assessment ‘proformas’ based on the findings of Phase 2 for each ecological group. This report outlines results of Phase 2. The Tillin et al., (2010) sensitivity assessment methodology was modified to use the best available scientific evidence that could be collated within the project timescale. An extensive literature review was compiled, for peer reviewed and grey literature, to examine current understanding about the effects of pressures from human activities on circalittoral and offshore sedimentary communities in UK continental shelf waters, together with information on factors that contribute to resilience (recovery) of marine species. This review formed the basis of an assessment of the sensitivity of the 16 ecological groups identified in Phase 1 of the project (Tillin & Tyler-Walters 2014). As a result: • the state of knowledge on the effects of each pressure on circalittoral and offshore benthos was reviewed; • the resistance, resilience and, hence, sensitivity of sixteen ecological groups, representing 96 characteristic species, were assessed for eight separate pressures; • each assessment was accompanied by a detailed review of the relevant evidence; Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with human activities • knowledge gaps and sources of uncertainty were identified for each group; • each assessment was accompanied by an assessment of the quality of the evidence, its applicability to the assessment and the degree of concordance (agreement) between the evidence, to highlight sources of uncertainty as an assessment of the overall confidence in the sensitivity assessment, and finally • limitations in the methodology and the application of sensitivity assessments were outlined. This process demonstrated that the ecological groups identified in Phase 1 (Tillin & Tyler-Walters 2014) were viable groups for sensitivity assessment, and could be used to represent the 33 circalittoral and offshore sediments biotopes identified at the beginning of the project. The results of the sensitivity assessments show: • the majority of species and hence ecological groups in sedimentary habitats are sensitive to physical change, especially loss of habitat and sediment extraction, and change in sediment type; • most sedimentary species are sensitive to physical damage, e.g. abrasion and penetration, although deep burrowing species (e.g. the Dublin Bay prawn - Nephrops norvegicus and the sea cucumber - Neopentadactyla mixta) are able to avoid damaging effects to varying degrees, depending on the depth of penetration and time of year; • changes in hydrography (wave climate, tidal streams and currents) can significantly affect sedimentary communities, depending on whether they are dominated by deposit, infaunal feeders or suspension feeders, and dependant on the nature of the sediment, which is itself modified by hydrography and depth; • sedentary species and ecological groups that dominate the top-layer of the sediment (either shallow burrowing or epifaunal) remain the most sensitive to physical damage; • mobile species (e.g. interstitial and burrowing amphipods, and perhaps cumaceans) are the least sensitive to physical change or damage, and hydrological change as they are already adapted to unstable, mobile substrata; • sensitivity to changes in organic enrichment and hence oxygen levels, is variable between species and ecological groups, depending on the exact habitat preferences of the species in question, although most species have at least a medium sensitivity to acute deoxygenation; • there is considerable evidence on the effects of bottom-contact fishing practices and aggregate dredging on sedimentary communities, although not all evidence is directly applicable to every ecological group; • there is lack of detailed information on the physiological tolerances (e.g. to oxygenation, salinity, and temperature), habitat preferences, life history and population dynamics of many species, so that inferences has been made from related species, families, or even the same phylum; • there was inadequate evidence to assess the effects of non-indigenous species on most ecological groups, and Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with human activities • there was inadequate evidence to assess the effects of electromagnetic fields and litter on any ecological group. The resultant report provides an up-to-date review of current knowledge about the effects of pressures resulting from human activities of circalittoral and offshore sedimentary communities. It provides an evidence base to facilitate and support the provision of management advice for Marine Protected Areas, development of UK marine monitoring and assessment, and conservation advice to offshore marine industries. However, such a review will require at least annual updates to take advantage of new evidence and new research as it becomes available. Also further work is required to test how ecological group assessments are best combined in practice to advise on the sensitivity of a range of sedimentary biotopes, including the 33 that were originally examined.

Ocean acidification impacts on nitrogen fixation in the coastal western Mediterranean Sea

The effects of ocean acidification on nitrogen (N2) fixation rates and on the community composition of N2-fixing microbes (diazotrophs) were examined in coastal waters of the North-Western Mediterranean Sea. Nine experimental mesocosm enclosures of ∼50 m3 each were deployed for 20 days during June-July 2012 in the Bay of Calvi, Corsica, France. Three control mesocosms were maintained under ambient conditions of carbonate chemistry. The remainder were manipulated with CO2 saturated seawater to attain target amendments of pCO2 of 550, 650, 750, 850, 1000 and 1250 μatm. Rates of N2 fixation were elevated up to 10 times relative to control rates (2.00 ± 1.21 nmol L-1d-1) when pCO2 concentrations were >1000 μatm and pHT (total scale) < 7.74. Diazotrophic phylotypes commonly found in oligotrophic marine waters, including the Mediterranean, were not present at the onset of the experiment and therefore, the diazotroph community composition was characterised by amplifying partial nifH genes from the mesocosms. The diazotroph community was comprised primarily of cluster III nifH sequences (which include possible anaerobes), and proteobacterial (α and γ) sequences, in addition to small numbers of filamentous (or pseudo-filamentous) cyanobacterial phylotypes. The implication from this study is that there is some potential for elevated N2 fixation rates in the coastal western Mediterranean before the end of this century as a result of increasing ocean acidification. Observations made of variability in the diazotroph community composition could not be correlated with changes in carbon chemistry, which highlights the complexity of the relationship between ocean acidification and these keystone organisms.

Complex coastal oceanographic fields can be described by universal multifractals

Characterization of chlorophyll and sea surface temperature (SST) structural heterogeneity using their scaling properties can provide a useful tool to estimate the relative importance of key physical and biological drivers. Seasonal, annual, and also instantaneous spatial distributions of chlorophyll and SST, determined from satellite measurements, in seven different coastal and shelf-sea regions around the UK have been studied. It is shown that multifractals provide a very good approximation to the scaling properties of the data: in fact, the multifractal scaling function is well approximated by universal multifractal theory. The consequence is that all of the statistical information about data structure can be reduced to being described by two parameters. It is further shown that also bathymetry scales in the studied regions as multifractal. The SST and chlorophyll multifractal structures are then explained as an effect of bathymetry and turbulence.

Defining scenarios of future vectors of change in marine life and associated economic sectors

Addressing the multitude of challenges in marine policy requires an integrated approach that considers the multitude of drivers, pressures, and interests, from several disciplinary angles. Scenarios are needed to harmonise the analyses of different components of the marine system, and to deal with the uncertainty and complexity of the societal and biogeophysical dynamics in the system. This study considers a set of socio-economic scenarios to (1) explore possible futures in relation to marine invasive species, outbreak forming species, and gradual changes in species distribution and productivity; and (2) harmonise the projection modelling performed within associated studies. The exercise demonstrates that developing interdisciplinary scenarios as developed in this study is particularly complicated due to (1) the wide variety in endogeneity or exogeneity of variables in the different analyses involved; (2) the dual role of policy decisions as variables in a scenario or decisions to be evaluated and compared to other decisions; and (3) the substantial difference in time scale between societal and physical drivers.