Lisa V. Lucas

Does the Sverdrup critical depth model explain bloom dynamics in estuaries

In this paper we use numerical models of coupled biological-hydrodynamic processes to search for general principles of bloom regulation in estuarine waters. We address three questions: What are the dynamics of stratie cation in coastal systems as ine uenced by variable freshwater input and tidal stirring? How does phytoplankton growth respond to these dynamics? Can the classical Sverdrup Critical Depth Model (SCDM) be used to predict the timing of bloom events in shallow coastal domains such as estuaries? We present results of simulation experiments which assume that vertical transport and net phytoplankton growth rates are horizontally homogeneous. In the present approach the temporally and spatially varying turbulent diffusivities for various stratie cation scenarios are calculated using a hydrodynamic code that includes the Mellor-Yamada 2.5 turbulence closure model. These diffusivities are then used in a time- and depth-dependent advection-diffusion equation, incorporating sources and sinks, for the phytoplankton biomass. Our modeling results show that, whereas persistent stratie cation greatly increases the probability of a bloom, semidiurnal periodic stratie cation does not increase the likelihood of a phytoplankton bloom over that of a constantly unstratie ed water column. Thus, for phytoplankton blooms, the physical regime of periodic stratie cation is closer to complete mixing than to persistent stratie cation. Furthermore, the details of persistent stratie cation are important: surface layer depth, thickness of the pycnocline, vertical density difference, and tidal current speed all weigh heavily in producing conditions which promote the onset of phytoplankton blooms. Our model results for shallow tidal systems do not conform to the classical concepts of stratie cation and blooms in deep pelagic systems. First, earlier studies (Riley, 1942, for example) suggest a monotonic increase in surface layer production as the surface layer shallows. Our model results suggest, however, a nonmonotonic relationship between phytoplankton population growth and surface layer depth, which results from a balance between several ‘ ‘ competing’ ’ processes, including the interaction of sinking with turbulent mixing and average net growth occurring within the surface layer. Second, we show that the traditional SCDM must be ree ned for application to energetic shallow systems or for systems in which surface layer mixing is not strong enough to counteract the sinking loss of phytoplankton. This need for ree nement arises because of the leakage of phytoplankton from the surface layer by turbulent diffusion and sinking, processes not considered in the classical SCDM. Our model shows that, even for low sinking rates and small turbulent

Functional variability of habitats within the Sacramento-San Joaquin Delta: Restoration implications

We have now entered an era of large-scale attempts to restore ecological functions and biological communities in impaired ecosystems. Our knowledge base of complex ecosystems and interrelated functions is limited, so the outcomes of specific restoration actions are highly uncertain. One approach for exploring that uncertainty and anticipating the range of possible restoration outcomes is comparative study of existing habitats similar to future habitats slated for construction. Here we compare two examples of one habitat type targeted for restoration in the Sacramento–San Joaquin River Delta. We compare one critical ecological function provided by these shallow tidal habitats—production and distribution of phytoplankton biomass as the food supply to pelagic consumers. We measured spatial and short-term temporal variability of phytoplankton biomass and growth rate and quantified the hydrodynamic and biological processes governing that variability. Results show that the production and distribution of phytopl...

Ecological Values of Shallow-Water Habitats: Implications for the Restoration of Disturbed Ecosystems

A presumed value of shallow-habitat enhanced pelagic productivity derives from the principle that in nutrient-rich aquatic systems phytoplankton growth rate is controlled by light availability, which varies inversely with habitat depth. We measured a set of biological indicators across the gradient of habitat depth within the Sacramento–San Joaquin River Delta (California) to test the hypothesis that plankton biomass, production, and pelagic energy flow also vary systematically with habitat depth. Results showed that phytoplankton biomass and production were only weakly related to phytoplankton growth rates whereas other processes (transport, consumption) were important controls. Distribution of the invasive clam Corbicula fluminea was patchy, and heavily colonized habitats all supported low phytoplankton biomass and production and functioned as food sinks. Surplus primary production in shallow, uncolonized habitats provided potential subsidies to neighboring recipient habitats. Zooplankton in deeper habitats, where grazing exceeded phytoplankton production, were likely supported by significant fluxes of phytoplankton biomass from connected donor habitats. Our results provide three important lessons for ecosystem science: (a) in the absence of process measurements, derived indices provide valuable information to improve our mechanistic understanding of ecosystem function and to benefit adaptive management strategies; (b) the benefits of some ecosystem functions are displaced by water movements, so the value of individual habitat types can only be revealed through a regional perspective that includes connectedness among habitats; and (c) invasive species can act as overriding controls of habitat function, adding to the uncertainty of management outcomes.

Effects of tidal shallowing and deepening on phytoplankton production dynamics: A modeling study

Processes influencing estuarine phytoplankton growth occur over a range of time scales, but many conceptual and numerical models of estuarine phytoplankton production dynamics neglect mechanisms occurring on the shorter (e.g., intratidal) time scales. We used a numerical model to explore the influence of short time-scale variability in phytoplankton sources and sinks on long-term growth in an idealized water column that shallows and deepens with the semidiurnal tide. Model results show that tidal fluctuations in water surface elevation can determine whether long-term phytoplankton growth is positive or negative. Hourly-scale interactions influencing weekly-scale to monthly-scale phytoplankton dynamics include intensification of the depth-averaged benthic grazing effect by water column shallowing and enhancement of water column photosynthesis when solar noon coincides with low tide. Photosynthesis and benthic consumption may modulate over biweekly time scales due to spring-neap fluctuations in tidal range and the 15-d cycle of solar noon-low tide phasing. If tidal range is a large fraction of mean water depth, then tidal shallowing and deepening may significantly influence net phytoplankton growth. In such a case, models or estimates of long-term phytoplankton production dynamics that neglect water surface fluctuations may overestimate or underestimate net growth and could even predict the wrong sign associated with net growth rate.

Intradaily Variability of Water Quality in a Shallow Tidal Lagoon: Mechanisms and Implications

Although surface water quality and its underlying processes vary over time scales ranging from seconds to decades, they have historically been studied at the lower (weekly to interannual) frequencies. The aim of this study was to investigate intradaily variability of three water quality parameters in a small freshwater tidal lagoon (Mildred Island, California). High frequency time series of specific conductivity, water temperature, and chlorophylla at two locations within the habitat were analyzed in conjunction with supporting hydrodynamic, meteorological, biological, and spatial mapping data. All three constituents exhibited large amplitude intradaily (e.g., semidiurnal tidal and diurnal) oscillations, and periodicity varied across constituents, space, and time. Like other tidal embayments, this habitat is influenced by several processes with distinct periodicities including physical controls, such as tides, solar radiation, and wind, and biological controls, such as photosynthesis, growth, and grazing. A scaling approach was developed to estimate individual process contributions to the observed variability. Scaling results were generally consistent with observations and together with detailed examination of time series and time derivatives, revealed specific mechanisms underlying the observed periodicities, including interactions between the tidal variability, heating, wind, and biology. The implications for monitoring were illustrated through subsampling of the data set. This exercise demonstrated how quantities needed by scientists and managers (e.g., mean or extreme concentrations) may be misrepresented by low frequency data and how short-duration high frequency measurements can aid in the design and interpretation of temporally coarser sampling programs. The dispersive export of chlorophylla from the habitat exhibited a fortnightly variability corresponding to the modulation of semidiurnal tidal currents with the diurnal cycle of phytoplankton variability, demonstrating how high frequency interactions can govern long-term trends. Process identification, as through the scaling analysis here, can help us anticipate changes in system behavior and adapt our own interactions with the system.

Changing restoration rules: exotic bivalves interact with residence time and depth to control phytoplankton productivity.

Non-native species are a prevalent ecosystem stressor that can interact with other stressors to confound resource management and restoration. We examine how interactions between physical habitat attributes and a particular category of non-native species (invasive bivalves) influence primary production in aquatic ecosystems. Using mathematical models, we show how intuitive relationships between phytoplankton productivity and controllable physical factors (water depth, hydraulic transport time) that hold in the absence of bivalves can be complicated—and even reversed—by rapid bivalve grazing. In light-limited environments without bivalves, shallow, hydrodynamically “slow” habitats should generally have greater phytoplankton biomass and productivity than deeper, “faster” habitats. But shallower, slower environments can be less productive than deeper, faster ones if benthic grazing is strong. Moreover, shallower and slower waters exhibit a particularly broad range of possible productivity outcomes that can depend on whether bivalves are present. Since it is difficult to predict the response of non-native bivalves to habitat restoration, outcomes for new shallow, slow environments can be highly uncertain. Habitat depth and transport time should therefore not be used as indicators of phytoplankton biomass and production where bivalve colonization is possible. This study provides for ecosystem management a particular example of a broad lesson: abiotic ecosystem stressors should be managed with explicit consideration of interactions with other major (including biotic) stressors. We discuss the applicability and management implications of our models and results for a range of aquatic system types, with a case study focused on the Sacramento-San Joaquin Delta (California, USA). Simple mathematical models like those used here can illuminate interactions between ecosystem stressors and provide process-based guidance for resource managers as they develop strategies to augment valued populations, restore habitats, and manipulate ecosystem functions.

Bivalve Grazing Can Shape Phytoplankton Communities

The ability of bivalve filter feeders to limit phytoplankton biomass in shallow waters is well-documented, but the role of bivalves in shaping phytoplankton communities is not. The coupled effect of bivalve grazing at the sediment-water interface and sinking of phytoplankton cells to that bottom filtration zone could influence the relative biomass of sinking (diatoms) and non-sinking phytoplankton. Simulations with a pseudo-2D numerical model showed that benthic filter feeding can interact with sinking to alter diatom:non-diatom ratios. Cases with the smallest proportion of diatom biomass were those with the fastest sinking speeds and strongest bivalve grazing rates. Hydrodynamics modulated the coupled sinking-grazing influence on phytoplankton communities. For example, in simulations with persistent stratification, the non-sinking forms accumulated in the surface layer away from bottom grazers while the sinking forms dropped out of the surface layer toward bottom grazers. Tidal-scale stratification also influenced vertical gradients of the two groups in opposite ways. The model was applied to Suisun Bay, a low-salinity habitat of the San Francisco Bay system that was transformed by the introduction of the exotic clam Potamocorbula amurensis. Simulation results for this Bay were similar to (but more muted than) those for generic habitats, indicating that P. amurensis grazing could have caused a disproportionate loss of diatoms after its introduction. Our model simulations suggest bivalve grazing affects both phytoplankton biomass and community composition in shallow waters. We view these results as hypotheses to be tested with experiments and more complex modeling approaches.

Population density, biomass, and age-class structure of the invasive clam Corbicula fluminea in rivers of the lower San Joaquin River watershed, California.

Abstract Corbicula fluminea is well known as an invasive filter-feeding freshwater bivalve with a variety of effects on ecosystem processes. However, C. fluminea has been relatively unstudied in the rivers of the western United States. In June 2003, we sampled C. fluminea at 16 sites in the San Joaquin River watershed of California, which was invaded by C. fluminea in the 1940s. Corbicula fluminea was common in 2 tributaries to the San Joaquin River, reaching densities of 200 clams · m−2, but was rare in the San Joaquin River. Biomass followed a similar pattern. Clams of the same age were shorter in the San Joaquin River than in the tributaries. Distribution of clams was different in the 2 tributaries, but the causes of the difference are unknown. The low density and biomass of clams in the San Joaquin River was likely due to stressful habitat or to water quality, because food was abundant. The success of C. fluminea invasions and subsequent effects on trophic processes likely depends on multiple factors. As C. fluminea continues to expand its range around the world, questions regarding invasion success and effects on ecosystems will become important in a wide array of environmental settings.