Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths

Phytoplankton-grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton-grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon:nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light-nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the "paradox of energy enrichment" (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.     

Consumers affect prey biomass and diversity through resource partitioning

Consumer presence and nutrient availability can have contrasting and interactive effects on plant diversity. In a factorial experiment, we manipulated two levels of nutrient supply and the presence of two moderately specialized grazers in different combinations (no grazers, two species in monoculture, and both in combination). We tested how nutrients and grazers regulated the biomass of marine coastal epiphytes and the diversity of algal assemblages, based on the prediction that the effect of consumers on prey diversity depends on productivity and consumer specialization. Nutrient enrichment increased the epiphytic load, while monocultures of single grazer species partly prevented epiphyte growth. However, only the presence of two species with complementary feeding preferences effectively prevented epiphyte overgrowth. The epiphytes comprised micro- and macroalgal species, and the diversity of these algal assemblages differed, depending on grazer identity. For the microalgae, diversity was reduced by nutrient addition when grazer control was inefficient, but not when specialist microalgal grazers were present. Macroalgal diversity was reduced in ambient water with specialist macroalgal grazers compared to the treatment with inefficient ones. These results indicate that grazer composition and productivity are crucial in determining whether consumer pressure will have a positive or negative effect on algal diversity.     

Light-mediated thresholds in stream-water nutrient composition in a river network

The elemental composition of solutes transported by rivers reflects combined influences of surrounding watersheds and transformations within stream networks, yet comparatively little is known about downstream changes in effects of watershed loading vs. in-channel processes. In the forested watershed of a river under a mediterranean hydrologic regime, we examined the influence of longitudinal changes in environmental conditions on water-column nutrient composition during summer base flow across a network of sites ranging from strongly heterotrophic headwater streams to larger, more autotrophic sites downstream. Small streams (0.1-10 km2 watershed area) had longitudinally similar nutrient concentration and composition with low (approximately 2) dissolved nitrogen (N) to phosphorus (P) ratios. Abrupt deviations from this pattern were observed in larger streams with watershed areas > 100 km2 where insolation and algal abundance and production rapidly increased. Downstream, phosphorus and silica concentrations decreased by > 50% compared to headwater streams, and dissolved organic carbon and nitrogen increased by approximately 3-6 times. Decreasing dissolved P and increasing dissolved N raised stream-water N:P to 46 at the most downstream sites, suggesting a transition from N limitation in headwaters to potential P limitation in larger channels. We hypothesize that these changes were mediated by increasing algal photosynthesis and N fixation by benthic algal assemblages, which, in response to increasing light availability, strongly altered stream-water nutrient concentration and stoichiometry in larger streams and rivers.     

Effects of grazer richness and composition on algal biomass in a closed and open marine system

Most natural local systems exchange organisms with a regional pool of species through migration and dispersal. Such metacommunity processes of interconnected multispecies assemblages are likely to affect local dynamics of both species and processes. We present results from an artificial marine outdoor rock pool system in which we investigated the factors of (1) local grazer richness and composition, and (2) connectivity of local patches to a regional species pool, and their effects on algal biomass. Local species richness of six grazers was manipulated in both open and closed pools, which were embedded in a regional species pool containing all six grazers. Grazer richness showed significant net biodiversity effects on grazing in the closed, but not in the open, system. Grazer composition, on the other hand, showed significant effects on grazing in both open and closed systems, depending on which species were initially present. The two most efficient grazers were able to compensate for less efficient grazers in species mixtures, hence ensuring the function of grazing. The efficiency of top-down control of algal biomass in open systems thus depends on which particular species are lost. Further, differences in grazing between the open and closed system changed over time due to temporal dynamics in grazer composition. The results emphasize the importance of including system connectivity in experimental designs to allow an extrapolation of biodiversity ecosystem-functioning relationships to natural systems.     

Variability in grazer-mediated defensive responses of green and red macroalgae on the south coast of South Africa

Variabilities in the responses of several South African red and green macroalgae to direct grazing and the responses of one green alga to cues from grazers were tested. We used two feeding experiments: (1) testing the induced responses of three red and one green algae to direct grazing by mesograzers and (2) a multi-treatment experiment, in which the direct and indirect effects of one macrograzer species on the green alga Codium platylobium were assessed. Consumption rates were assessed in feeding assays with intact algal pieces and with agar pellets containing non-polar extracts of the test algae. Defensive responses were induced for intact pieces of Galaxaura diessingiana, but were not induced in pellets, suggesting either morphological defence or chemical defence using polar compounds other than polyphenols. In contrast, exposure to grazing stimulated consumption of Gracilaria capensis and Hypnea spicifera by another grazing species. In the multi-treatment experiment, waterborne cues from both grazing and non-grazing snails induced defensive algal traits in C. platylobium. We suggest that inducible defences among macroalgae are not restricted to brown algae, but that both the responses of algae to grazers and of grazers to the defences of macroalgae are intrinsically variable and complex.     

Stoichiometry, growth, and fecundity responses to nutrient enrichment by invertebrate grazers in sub-tropical turtle grass (Thalassia testudinum) meadows

Although the effectiveness of herbivores in mitigating the effects of nutrient enrichment is well documented, few studies have examined the effects of nutrient enrichment on components of consumer fitness. Enclosures were deployed in shallow turtle grass (Thalassia testudinum) beds in Florida Bay, Florida in fall 2003, spring 2004, and fall 2004 to measure the effects of nitrogen and phosphorous enrichment on the growth, fecundity, and stoichiometry of three invertebrate epiphyte grazers commonly associated with T. testudinum. The gastropod Turbo castanea exhibited significantly greater wet weight gain and lower C:P and N:P in enriched than in ambient treatments. Although nutrient enrichment did not have any significant effects on the growth of caridean shrimp (treatment consisted of several different caridean shrimp species), their C:N was significantly lower in enriched treatments. The final size and stoichiometry of the hermit crab Paguristes tortugae was not significantly affected by nutrient enrichment, nor did nutrient enrichment significantly affect the fecundity of P. tortugae, the only grazer in which gravid individuals or egg masses were present. Our study demonstrated that nutrient enrichment of primary producers can positively affect the growth of marine invertebrate grazers and alter their stoichiometry; however, these effects were species-specific and may be dependent upon the life stage, specific diets, and/or compensatory feeding habits of the grazers.     

Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea

The Baltic Sea is a very suitable site for stoichiometric studies, since its subbasins differ in their concentration of elemental components, and primary production can therefore be either nitrogen or phosphorus limited. To reveal if the nutrient limitation of mesozooplankton mirrors that of the primary producers, carbon, nitrogen and phosphorus content of both seston and grazers (Acartia sp., Centropages hamatus, Daphnia cristata, Eurytemora affinis, Limnocalanus macrurus, Temora longicornis) were measured in midsummer in the Baltic proper, the Gulf of Finland and the Gulf of Bothnia. The mineral ratios of the different taxa were equal, apart from L. macrurus with notably higher C:P and N:P ratios. Molar C:N ratios were relatively stable (5.1-6.3), whereas C:P and N:P ratios fluctuated more (41-144 and 6.6-24). However, zooplankton elemental composition and limitation did not depend on the limiting nutrient of the phytoplankton, the seston mineral ratio or the sea area. Both the seston-zooplankton elemental imbalance and the food threshold ratio indicated phosphorus limitation of most of the grazers. While L. macrurus may be C or N limited, the possible P deficiency of the other studied taxa suggests that the Baltic Sea zooplankton may act as a potential phosphorus sink, as the freshwater secondary producers do.     

Herbivore vs. nutrient control of marine primary producers: context-dependent effects

Pervasive overharvesting of consumers and anthropogenic nutrient loading are changing the strengths of top-down and bottom-up forces in ecosystems worldwide. Thus, identifying the relative and synergistic roles of these forces and how they differ across habitats, ecosystems, or primary-producer types is increasingly important for understanding how communities are structured. We used factorial meta-analysis of 54 field experiments that orthogonally manipulated herbivore pressure and nutrient loading to quantify consumer and nutrient effects on primary producers in benthic marine habitats. Across all experiments and producer types, herbivory and nutrient enrichment both significantly affected primary-producer abundance. They also interacted to create greater nutrient enrichment effects in the absence of herbivores, suggesting that loss of herbivores produces more dramatic effects of nutrient loading. Herbivores consistently had stronger effects than did nutrient enrichment for both tropical macroalgae and seagrasses. The strong effects of herbivory but limited effects of nutrient enrichment on tropical macroalgae suggest that suppression of herbivore populations has played a larger role than eutrophication in driving the phase shift from coral- to macroalgal-dominated reefs in many areas, especially the Caribbean. For temperate macroalgae and benthic microalgae, the effects of top-down and bottom-up forces varied as a function of the inherent productivity of the ecosystem. For these algal groups, nutrient enrichment appeared to have stronger effects in high- vs. low-productivity systems, while herbivores exerted a stronger top-down effect in low-productivity systems. Effects of herbivores vs. nutrients also varied among algal functional groups (crustose algae, upright macroalgae, and filamentous algae), within a functional group between temperate and tropical systems, and according to the metric used to measure producer abundance. These analyses suggest that human alteration of food webs and nutrient availability have significant effects on primary producers but that the effects vary among latitudes and primary producers, and with the inherent productivity of ecosystems.     

Spatial variation in caddisfly grazing regimes within a northern California watershed

Ecologists seek better understanding of why species interactions change across space and time in natural communities. In streams, species effects on resources and community structure may change as physical characteristics of the stream environment change along drainage networks. We examined spatial and seasonal effects of armored grazers using a small-scale exclusion experiment that was replicated in streams of different drainage areas. Effects of grazing varied with stream size and were related to variation in grazer abundance and phenology. We identified three distinct grazing regimes and a stream size (drainage area [DA]) threshold corresponding to a shift from one to two functional trophic levels. In streams with DA < 1 km2, armored grazers did not reduce biomass of algal biofilms. In slightly larger streams (2-3 km2 DA), the armored grazer guild was dominated by bivoltine Glossosoma. These caddisflies persisted and limited algal biofilms throughout the summer in one of these streams. In the largest tributaries (DA > 10 km2), the grazer guild was dominated by univoltine caddisflies, and grazing limited algal biofilms in early summer, but not late summer, after caddisflies pupated. Drainage area is a useful predictor of spatial transitions in food web interactions within and among watersheds. Quantifying the drainage area threshold at which interactions change in catchments with differing geology, vegetation, hydrology, climate, land use, or species pools should help build the understanding we need to forecast ecological responses to environmental change.     

Anthropogenic subsidies alter stream consumer-resource stoichiometry, biodiversity, and food chains.

Urbanization is dramatically changing nutrient and organic matter regimes in streams, yet the community and ecosystem implications often remain obscure. We assessed the consequences of sewage-derived particulate organic matter (SDPOM) for invertebrate community structure and function in a headwater stream. Using stable isotope analyses, we found assimilation of organic SDPOM to double community secondary production, and stoichiometric analyses revealed SDPOM enriched in phosphorus (P) to foster putatively fast-growing, P-rich consumers in the subsidized reach. This altered consumer-resource stoichiometry impacted both community structure and nutrient fluxes through the invertebrate community. Community structure shifted toward significantly reduced diversity and evenness in the subsidized reach and consequently toward shorter food chains. Our integration of ecological stoichiometry with stable isotope analyses and food web ecology expands the previous focus of traditional ecotoxicology and ecophysiology to an ecosystem-level appreciation of pollutant ecology.     

Unintended nutrient imbalance induced by wastewater effluent inputs to receiving water and its ecological consequences

Eutrophication is the most widespread water quality issue globally. To date, most efforts to control eutrophication have focused on reductions of external nutrient inputs, yet importance of nutrient stoichiometry and subsequent shift in plankton composition in aquatic ecosystem has been largely neglected. To address eutrophication, improved sanitation is one of the United Nations Sustainable Development Goals, spurring the constructions of wastewater treatment facilities that have improved water quality in many lakes and rivers. However, control measures are often targeted at and effective in removing a single nutrient from sewage and thus are less effective in removing the others, resulting in the changes of nutrient stoichiometry. In general, more effective phosphorus removal relative to nitrogen has occurred in wastewater treatment leading to substantial increases in N/P ratios in effluent relative to the influent. Unfortunately, high N/P ratios in receiving waters can impose negative influences on ecosystems. Thus, long-term strategies for domestic wastewater management should not merely focus on the total reduction of nutrient discharge but also consider their stoichiometric balances in receiving waters.     

Determination of a limiting nutrient regulating algal biomass using in situ experiments of nutrient enrichment bioassay (NEB) and empirical relations of nutrients and chlorophyll-a

Long-term nutrient contents and nutrient ratios indicated that phosphorus was a potential limiting element for algal growth. In situ experiments of nutrient enrichment bioassay supported the evidence of P-limitation. However, regression analyses of log10-transformed chlorophyll-a (CHL) against TP (R2 values < 0.25) showed that seasonal CHL was not closely related to flux of phosphorus during all seasons. Also, two dimensional graphical approach of Trophic State Index (TSI) showed that most values of TSI (CHL) -TSI (TP) and TSI (CHL) -TSI (SD) were less than zero, indicating factors other than phosphorus limited algal biomass (CHL -TP < 0), and that non-algal particles dominated light attenuation (CHL -SD < 0). The weak empirical relations and trophic deviations were explained well by the experiment of NEB-II that was conduced during a period of high inorganic turbidity. Overall results suggest that phosphorus is the primary element regulating the system productivity, but the system also were highly influenced by rapid flushing and high inorganic turbidity.     

Dynamic stoichiometric response to food quality fluctuations in the heterotrophic dinoflagellate Oxyrrhis marina

With respect to nutrients, plants are rather non-homoeostatic while most metazoans have much more confined ranges of nutrient ratios. It was recently highlighted that the homoeostatic ability of microzooplankters lies in between these two extremes. Nevertheless, we know very little on the dynamics of stoichiometric changes. Hence, we investigated how the stoichiometry of the heterotrophic dinoflagellate Oxyrrhis marina is affected (1) during a starvation period and (2) when fed nutrient deplete Rhodomonas salina after having been pre-conditioned on nutrient replete algae and vice versa. We observed that the dinoflagellate was able to maintain its N:P ratio constant over 78 h of starvation. We inferred that under starvation, nitrogen-limited O. marina mainly used fat as energy source while nitrogen-rich individuals also used proteins as fuel in cellular respiration. Further, we showed that O. marina presents resistance to nutrient limitation, with stronger regulation against P-limitation than against N-limitation. This resilience in microzooplankton stoichiometry following food quality stress would have great implications for both top-down (nutrient remineralisation) and bottom-up controls (quality as food).     

Regulation of algal community development in a Macrocystis pyrifera forest

The effects of small and large-scale roughness, overstory development, competition for space with sessile animals, and grazing on algal community development in a subtidal Macrocystis pyrifera forest were examined using specially prepared concrete blocks as substrata. Variation in small-scale roughness (crevices and grooves in the order of 0.1 to 3 mm width and depth) had no significant effects on community composition. However, M. pyrifera colonization, algal diversity, and sessile animal biomass were higher near the upper horizontal edges of blocks and concrete prisms. This “edge” effect may result from a combination of increased spore and larval settlement and enhanced growth of plants and animals associated with the turbulent eddies formed around these obstructions. Natural and experimentally produced variations in the algal overstory demonstrated that the presence of an overstory can reduce algal diversity and cover beneath. Caging experiments suggest that predatory fishes and sea-stars indirectly affect the algal community by removing sessile animals (primarily bryozoans) which compete with the algae for space. The exclusion of grazers resulted in increased growth of Gigartina spp. Selective grazing on this genus may account for its reduced abundance in the study area.     

Origins and scales of hypoxia on the Louisiana shelf: Importance of seasonal plankton dynamics and river nutrients and discharge

Management plans for the Mississippi River Basin call for reductions in nutrient concentrations up to 40% or more to reduce hypoxia in the Gulf of Mexico (GOM), while at the same time the government is considering new farm subsidies to promote development of biofuels from corn. Thus there are possibilities of both increasing and decreasing river nutrients depending on national priorities. River flow rates which also influence the extent of hypoxia on the shelf may be altered by global climate change. We have therefore developed a series of simulations to forecast ecosystem response to alterations in nutrient loading and river flow. We simulate ecosystem response and hypoxia events using a linked model consisting of multiple phytoplankton groups competing for nitrogen, phosphorus and light, zooplankton grazing that is influenced by prey edibility and stoichiometry, sub-pycnocline water-column metabolism that is influenced by sinking fecal pellets and algal cells, and multi-element sediment diagenesis. This model formulation depicts four areas of increasing salinity moving westward away from the Mississippi River point of discharge, where the surface mixed layer, four bottom layers and underlying sediments are represented in each area. The model supports the contention that a 40% decrease in river nutrient will substantially reduce the duration and areal extent of hypoxia on the shelf. But it also suggests that in low and middle salinity areas the hypoxia response is saturated with respect to nutrients, and that in high salinity regions small increases in nutrient and river flow will have disproportionally large effects on GOM hypoxia. The model simulations also suggest that river discharge is a stronger factor influencing hypoxia than river nutrients in the Mississippi River plume. Finally, the model simulations suggest that primary production in the low salinity regions is light limited while primary production in the higher salinity zones is phosphate limited during the May to October period when hypoxia is prevalent in the Mississippi River plume.     

Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.     

Microbial nitrogen limitation increases decomposition

With anthropogenic nutrient inputs to ecosystems increasing globally, there are long-standing, fundamental questions about the role of nutrients in the decomposition of organic matter. We tested the effects of exogenous nitrogen and phosphorus inputs on litter decomposition across a broad suite of litter and soil types. In one experiment, C mineralization was compared across a wide array of plants individually added to a single soil, while in the second, C mineralization from a single substrate was compared across 50 soils. Counter to basic stoichiometric decomposition theory, low N availability can increase litter decomposition as microbes use labile substrates to acquire N from recalcitrant organic matter. This "microbial nitrogen mining" is consistently suppressed by high soil N supply or substrate N concentrations. There is no evidence for phosphorus mining as P fertilization increases short- and long-term mineralization. These results suggest that basic stoichiometric decomposition theory needs to be revised and ecosystem models restructured accordingly in order to predict ecosystem carbon storage responses to anthropogenic changes in nutrient availability.     

Too much of a good thing: on stoichiometrically balanced diets and maximal growth

Nutritional imbalances are of great interest in the ecological stoichiometry literature, in which researchers have focused almost exclusively on cases where nutrients are available in low amounts relative to energy (carbon), and animal growth is impaired due to insufficient nutrient intake. Little attention has been given to situations where food elemental content is higher than the level that satisfies animal requirements. However, most animals are strongly homeostatic with respect to the elemental composition of their body; hence they must excrete the excess of elements that are not in short supply. To date, stoichiometric theory has assumed that excretion of superfluous elements does not come with a cost and, thus, that consumption of food with surplus nutrients does not impair performance. Here we challenge this assumption, based on a compilation of several examples involving food phosphorus content that show that the performance of a wide variety of animals decreases when supplied with food containing high concentrations of (potentially) limiting nutrients. We discuss possible mechanisms for this phenomenon, and suggest that animals most vulnerable to effects of high food nutrient content are those that normally feed on low- quality (low-nutrient: C) food, and have a relatively low body nutrient content themselves, such as herbivores and detritivores.     

Does biodiversity of estuarine phytoplankton depend on hydrology?

Phytoplankton growth in estuaries is controlled by factors such as flushing, salinity tolerance, light, nutrients and grazing. Here, we show that biodiversity of estuarine phytoplankton is related to flushing, and illustrate this for some European estuaries.The implications for the definition of reference conditions for quality elements in estuaries of different types are examined, leading to the conclusion that constraints on the number of estuarine and coastal types that may be defined for management purposes require that quality classes take into account natural variability within types, in order to be ecologically meaningful. We develop a screening model to predict the growth rate required for a phytoplankton species to be present under different flushing conditions and apply it to estuaries in the EU and US to show how changes in physical forcing may alter biodiversity. Additional results are presented on the consequences for eutrophication, showing that changes in residence time may interact with species-specific nutrient uptake rates to cause shifts in species composition, potentially leading to effects such as harmful algal blooms.We discuss applications for integrated coastal zone management, and propose an approach to normalization of estuarine phytoplankton composition as regards species numbers.