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J. E. Cloern

Publications and source records attributed to J. E. Cloern.

At least 19 recordsLinked to original sources

Drivers of change in estuarine-coastal ecosystems: Discoveries from four decades of study in San Francisco Bay

Poised at the interface of rivers, ocean, atmosphere and dense human settlement, estuaries are driven by a large array of natural and anthropogenic forces. San Francisco Bay exemplifies the fast-paced change occurring in many of the world's estuaries, bays and inland seas in response to these diverse forces. We use observations from this particularly well-studied estuary to illustrate responses to six drivers that are common agents of change where land and sea meet: water consumption and diversion; human modification of sediment supply; introduction of non-native species; sewage input; environmental policy; and climate shifts. In San Francisco Bay, responses to these drivers include, respectively, shifts in the timing and extent of freshwater inflow and salinity intrusion; decreasing turbidity; restructuring of plankton communities; nutrient enrichment; elimination of hypoxia and reduced metal contamination of biota; and food web changes that decrease resistance of the estuary to nutrient pollution. Detection of these changes and discovery of their causes through environmental monitoring have been essential for establishing and measuring outcomes of environmental policies that aim to maintain high water quality and sustain services provided by estuarine-coastal ecosystems. The wide range of variability time scales and the multiplicity of interacting drivers place heavy demands on estuarine monitoring programs. But the San Francisco Bay case study illustrates why the imperative for monitoring has never been greater.

California

Complex seasonal patterns of primary producers at the land-sea interface

Seasonal fluctuations of plant biomass and photosynthesis are key features of the Earth system because they drive variability of atmospheric CO 2 , water and nutrient cycling, and food supply to consumers. There is no inventory of phytoplankton seasonal cycles in nearshore coastal ecosystems where forcings from ocean, land and atmosphere intersect. We compiled time series of phytoplankton biomass (chlorophyll a ) from 114 estuaries, lagoons, inland seas, bays and shallow coastal waters around the world, and searched for seasonal patterns as common timing and amplitude of monthly variability. The data revealed a broad continuum of seasonal patterns, with large variability across and within ecosystems. This contrasts with annual cycles of terrestrial and oceanic primary producers for which seasonal fluctuations are recurrent and synchronous over large geographic regions. This finding bears on two fundamental ecological questions: (1) how do estuarine and coastal consumers adapt to an irregular and unpredictable food supply, and (2) how can we extract signals of climate change from phytoplankton observations in coastal ecosystems where local‐scale processes can mask responses to changing climate?

Ecology Letters

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.

Ecosystems

What is causing the phytoplankton increase in San Francisco Bay?

The largest living component of San Francisco Bay is the phytoplankton, a suspension of microscopic cells that convert sunlight energy into new living biomass through the same process of photosynthesis used by land plants. This primary production is the ultimate source of food for clams, zooplankton, crabs, sardines, halibut, sturgeon, diving ducks, pelicans, and harbor seals. From measurements made in 1980, we estimated that phytoplankton primary production in San Francisco Bay was about 200,000 tons of organic carbon per year (Jassby et al. 1993). This is equivalent to producing the biomass of 5500 adult humpback whales, or the calories to feed 1.8 million people. These numbers may seem large, but primary production in San Francisco Bay is low compared to many other nutrient-enriched estuaries.

California

Phytoplankton community ecology: Principles applied in San Francisco Bay

In his seminal 1961 paper 'The paradox of the plankton' Am Nat 95:137-147, G. E. Hutchinson asked why many species of phytoplankton can coexist while competing for a small number of limiting resources in an unstructured habitat. Hutchinson anticipated the resolution of his paradox, recognizing that communities are organized by processes beyond resource competition including species interactions, habitat variability and dispersal. Since 1961 we have made fundamental discoveries that have revolutionized our conceptual understanding of pelagic ecology, including (1) habitat heterogeneity at all scales relevant to plankton population dynamics, (2) community shifts in response to global climate cycles, (3) fast and selective predation as a powerful top-down force to shape phytoplankton communities, (4) turbulent mixing as a physical process that selects species on the basis of their size and form, (5) mixotrophy that allows some algal species to tap organic nutrient pools and function at multiple trophic levels, (6) taxon-specific life cycles including alternating vegetative and resting stages, and (7) the pelagic as an open system where communities are continually reshaped by species immigration. Here we synthesize these discoveries to show how they validate and amplify Hutchinson's hypothesis that phytoplankton communities are assembled by many processes. Our synthesis is built around observations of phytoplankton species composition from a decade of study in San Francisco Bay, used as a case study to illustrate the contemporary principles of phytoplankton community ecology. We apply these principles to address 2 central questions: (1) What processes assemble phytoplankton communities? (2) How does phytoplankton community composition influence ecosystem functions such as production in pelagic and benthic food webs?

California

Metal uptake by phytoplankton during a bloom in South San Francisco Bay: Implications for metal cycling in estuaries

The 1994 spring phytoplankton bloom in South San Francisco Bay caused substantial reductions in concentrations of dissolved Cd, Ni, and Zn, but not Cu. We estimate that the equivalent of ~60% of the total annual input of Cd, Ni, and Zn from local waste‐water treatment plants is cycled through the phytoplankton in South Bay. The results suggest that processes that affect phytoplankton bloom frequency or intensity in estuaries (e.g. nutrient enrichment) may also affect metal trapping. The bloom was characterized by hydrographic surveys conducted at weekly intervals for 9 weeks. Metal samples were collected from the water column on three occasions, timed to bracket the period when the bloom was predicted. Factors that might confound observations of biological influences, such as freshwater inputs, were relatively constant during the study. Before the bloom, concentrations of dissolved Cd were 0.81 ± 0.02 nmol kg −1 , Zn concentrations were 19.8 ± 1.5 nmol kg −1 , Ni were 42 ± 1.4 nmol kg −1 , and Cu were 37 ± 1.4 nmol kg −1 . These values are elevated relative to riverine and coastal end‐members, reflecting inputs from wastewater and(or) sediments. At the height of the bloom, dissolved Zn, Cd, and Ni were reduced to 19, 50, and 75% of their prebloom concentrations, respectively. Dissolved Cu concentrations increased 20%. The mass of Cd taken up by phytoplankton was similar to the mass of Cd removed from solution if particle settling was considered, and Cd concentrations estimated in phytoplankton were higher than concentrations in suspended particulate material (SPM). Particulate concentrations of Zn and Ni during the bloom appeared to be dominated by the influence of changes in resuspension of Zn and Ni‐rich sediments.

California

Bioavailability of organic matter in a highly disturbed Estuary: The role of detrital and algal resources

The importance of algal and detrital food supplies to the planktonic food web of a highly disturbed, estuarine ecosystem was evaluated in response to declining zooplankton and fish populations. We assessed organic matter bioavailability among a diversity of habitats and hydrologic inputs over 2 years in San Francisco Estuary's Sacramento-San Joaquin River Delta. Results show that bioavailable dissolved organic carbon from external riverine sources supports a large component of ecosystem metabolism. However, bioavailable particulate organic carbon derived primarily from internal phytoplankton production is the dominant food supply to the planktonic food web. The relative importance of phytoplankton as a food source is surprising because phytoplankton production is a small component of the ecosystem's organic-matter mass balance. Our results indicate that management plans aimed at modifying the supply of organic matter to riverine, estuarine, and coastal food webs need to incorporate the potentially wide nutritional range represented by different organic matter sources.

Proceedings of the National Academy of Sciences of

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 phytoplankton biomass can be highly variable within and between nearby habitats of the same type, due to variations in phytoplankton sources, sinks, and transport. Therefore, superficially similar, geographically proximate habitats can function very differently, and that functional variability introduces large uncertainties into the restoration process. Comparative study of existing habitats is one way ecosystem science can elucidate and potentially minimize restoration uncertainties, by identifying processes shaping habitat functionality, including those that can be controlled in the restoration design.

California

Spatial and temporal variability of picocyanobacteria Synechococcus sp. in San Francisco Bay

We collected samples monthly, from April to August 1998, to measure the abundance of autotrophic picoplankton in San Francisco Bay. Samples taken along a 160-km transect showed that picocyanobacteria (Synechococcus sp.) was a persistent component of the San Francisco Bay phytoplankton in all the estuarine habitats, from freshwater to seawater and during all months of the spring-summer transition. Abundance ranged from 4.6 X 106 to 5.2 X 108 cells L-1, with peak abundance during the spring bloom (April and May) and during July with a persistent spatial pattern of smallest abundance near the coastal ocean and highest abundance in the landward domains of the estuary. The picocyanobacterial component (as estimated percentage of chlorophyll a concentration) was, on average, 15% of total phytoplankton biomass during the summer-autumn nonbloom periods and only 2% of chlorophyll biomass during the spring bloom. This result is consistent with the emerging concept of a gradient of increasing importance of picocyanobacteria along the gradient of decreasing nutrient concentrations from estuaries to the open ocean.

Limnology and Oceanography

Dynamics of nutrient cycling and related benthic nutrient and oxygen fluxes during a spring phytoplankton bloom in South San Francisco Bay (USA)

Benthic oxygen uptake and nutrient releases of N, P and Si were measured weekly at 2 sites in South San Francisco Bay around the 1996 spring bloom. Exchanges across the sediment-water interface were estimated from whole core incubations performed in the laboratory at in situ temperature and in dark. Fluxes changed significantly on a weekly time scale. Over a period of 15 wk the fluxes of dissolved inorganic N, P and Si ranged from -40 to +200, 0 to 13 and from 30 to 400 µmol m -2 h -1 respectively. Sediment oxygen demand increased from 10 before to 64 mg O 2 m -2 h -1 just after the bloom period. During the bloom, nutrient fluxes represented about 20, 16 and 9% of the Si, P and N requirements for primary production. Before and after the bloom period, Si fluxes contributed up to 30 and >100% of this requirement and P and N fluxes up to 15 and 50% respectively. Simple empirical models explain most of the spatial-temporal variability of benthic fluxes of Si, P and NH 4 (but not NO 3 ) from 3 predictor variables: sediment porosity, nutrient concentration in bottom waters and chlorophyll content of surficial sediments. These models show that algal blooms influence benthic-pelagic nutrient exchange through 2 processes: (1) depletion of nutrients from the water column (which enhances gradient-driven transports across the sediment-water interface) and (2) sedimentation of labile phytodetritus (which promotes remineralization in or on the surficial sediments). Rates and patterns of nutrient cycling were very different at the shallow and deep study sites, illustrating the challenge of extrapolating measurements of coupled algae-nutrient dynamics to whole ecosystems.

Marine Ecology Progress Series

The relative importance of light and nutrient limitation of phytoplankton growth: A simple index of coastal ecosystem sensitivity to nutrient enrichment

Anthropogenic nutrient enrichment of the coastal zone is now a well-established fact. However, there is still uncertainty about the mechanisms through which nutrient enrichment can disrupt biological communities and ecosystem processes in the coastal zone. For example, while some estuaries exhibit classic symptoms of acute eutrophication, including enhanced production of algal biomass, other nutrient-rich estuaries maintain low algal biomass and primary production. This implies that large differences exist among coastal ecosystems in the rates and patterns of nutrient assimilation and cycling. Part of this variability comes from differences among ecosystems in the other resource that can limit algal growth and production - the light energy required for photosynthesis. Complete understanding of the eutrophication process requires consideration of the interacting effects of light and nutrients, including the role of light availability as a regulator of the expression of eutrophication. A simple index of the relative strength of light and nutrient limitation of algal growth can be derived from models that describe growth rate as a function of these resources. This index can then be used as one diagnostic to classify the sensitivity of coastal ecosystems to the harmful effects of eutrophication. Here I illustrate the application of this diagnostic with light and nutrient measurements made in three California estuaries and two Dutch estuaries.

Aquatic Ecology

Processes governing phytoplankton blooms in estuaries. I: The local production-loss balance

The formation and spatial distribution of phytoplankton blooms in estuaries are controlled by (1) local mechanisms, which determine the production-loss balance for a water column at a particular spatial location (i.e. control if a bloom is possible), and (2) transport-related mechanisms, which govern biomass distribution (i.e. control if and where a bloom actually occurs). In this study, the first of a 2-paper series, we use a depth-averaged numerical model as a theoretical tool to describe how interacting local conditions (water column height, light availability, benthic grazing) influence the local balance between phytoplankton sources and sinks. We also explore trends in the spatial variability of the production-loss balance across the topographic gradients between deep channels and lateral shoals which are characteristic of shallow estuaries. For example, under conditions of high turbidity and slow benthic grazing the highest rates of phytoplankton population growth are found in the shallowest regions. On the other hand, with low turbidity and rapid benthic grazing the highest growth rates occur in the deeper areas. We also explore the effects of semidiurnal tidal variation in water column height, as well as spring-neap variability. Local population growth in the shallowest regions is very sensitive to tidal-scale shallowing and deepening of the water column, especially in the presence of benthic grazing. A spring-neap signal in population growth rate is also prominent in the shallow areas. Population growth in deeper regions is less sensitive to temporal variations in tidal elevation. These results show that both shallow and deep regions of estuaries can act as sources or sinks for phytoplankton biomass, depending on the local conditions of mean water column height, tidal amplitude, light-limited growth rate, and consumption by grazers.

Marine Ecology Progress Series

Processes governing phytoplankton blooms in estuaries. II: The role of horizontal transport

The development and distribution of phytoplankton blooms in estuaries are functions of both local conditions (i.e. the production-loss balance for a water column at a particular spatial location) and large-scale horizontal transport. In this study, the second of a 2-paper series, we use a depth-averaged hydrodynamic-biological model to identify transport-related mechanisms impacting phytoplankton biomass accumulation and distribution on a system level. We chose South San Francisco Bay as a model domain, since its combination of a deep channel surrounded by broad shoals is typical of drowned-river estuaries. Five general mechanisms involving interaction of horizontal transport with variability in local conditions are discussed. Residual (on the order of days to weeks) transport mechanisms affecting bloom development and location include residence time/export, import, and the role of deep channel regions as conduits for mass transport. Interactions occurring on tidal time scales, i.e. on the order of hours) include the phasing of lateral oscillatory tidal flow relative to temporal changes in local net phytoplankton growth rates, as well as lateral sloshing of shoal-derived biomass into deep channel regions during ebb and back into shallow regions during flood tide. Based on these results, we conclude that: (1) while local conditions control whether a bloom is possible, the combination of transport and spatial-temporal variability in local conditions determines if and where a bloom will actually occur; (2) tidal-time-scale physical-biological interactions provide important mechanisms for bloom development and evolution. As a result of both subtidal and tidal-time-scale transport processes, peak biomass may not be observed where local conditions are most favorable to phytoplankton production, and inherently unproductive areas may be regions of high biomass accumulation.

Marine Ecology Progress Series

A CALFED-supported study of the Delta’s foodweb base

Scientists from six universities and the US Geological Survey (USGS) have begun a new project to characterize the food resource which supports secondary production in different habitats of the Sacramento-San Joaquin Delta. The project is supported by CALFED Category III, USGS, and the IEP; it is designed to answer basic questions about the organic matter which supports biological production at the lowest trophic levels. We know, from the excellent long-term records of IEP agencies, that the abundance of zooplankton (especially native species) has declined dramatically in the past three decades. We do not understand the underlying mechanisms of these declines, but it is likely that they are the result of multiple, interacting stressors. One potential limitation of secondary production could be a sub-optimal food resource, manifested either as low quantity or poor nutritional quality of the available pools of organic matter. Some past studies support this possibility, especially for the case of benthic macroinvertebrates. The project was conceived to do the following: Identify the potential sources of organic matter (for example, phytoplankton, vascular plants, agricultural sources) in different habitats; Determine the relative food quality of each of these different sources for secondary producers, either directly or after microbial processing; and Assess the habitats and environmental conditions that provide the maximum food resources for secondary producers.

Interagency Ecological Program Newsletter

Water-quality variability in San Francisco Bay: general patterns of change during 1997

The 1997 Annual Report is the fifth Annual Report from the Regional Monitoring Program for Trace Substances (RMP) and contains a comprehensive description of RMP results from the 1997 monitoring year. As in previous years, the report includes results from the Base Program (water, sediment, and bivalve monitoring) and results from Pilot and Special Studies completed in 1997, in addition to an update on the RMP Five-Year Review implementation. It also includes papers contributed by RMP investigators and other scientists. These articles address related monitoring activities, and help to provide additional insight into contaminant patterns and the impacts of those contaminants on the San Francisco Estuary. The 1997 monitoring year proved to be an unusual one, with record-setting precipitation in December and January followed by unusually dry weather in February and March. These weather patterns had a visible effect on RMP results, frequently creating sharp contrasts in results between the first two sampling cruises of the year, and higher than normal contaminant concentrations at many RMP sampling sites in February. These results, and results from the other aspects of the RMP, are summarized below.

California

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 stratification in coastal systems as influenced 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 stratification 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 stratification greatly increases the probability of a bloom, semidiurnal periodic stratification does not increase the likelihood of a phytoplankton bloom over that of a constantly unstratified water column. Thus, for phytoplankton blooms, the physical regime of periodic stratification is closer to complete mixing than to persistent stratification. Furthermore, the details of persistent stratification 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 stratification 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 refined 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 refinement 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 diffusivities, a significant % of the phytoplankton biomass produced in the surface layer can be lost by these processes.

Journal of Marine Research

Changes in production and respiration during a spring phytoplankton bloom in San Francisco Bay, California, USA: Implications for net ecosystem metabolism

We present results of an intensive sampling program designed to measure weekly changes in ecosystem respiration (oxygen consumption in the water column and sediments) around the 1996 spring bloom in South San Francisco Bay, California, USA. Measurements were made at a shallow site (2 m, where mean photic depth was 60% of the water column height) and a deep site (15 m, mean photic depth was only 20% of the water column). We also estimated phytoplankton primary production weekly at both sites to develop estimates of net oxygen flux as the sum of pelagic production (PP), pelagic respiration (PR) and benthic respiration (BR). Over the 14 wk period from February 5 to May 14, PP ranged from 2 to 210, PR from 9 to 289, and BR from 0.1 to 48 mmol O2 m-2 d-1, illustrating large variability of estuarine oxygen fluxes at the weekly time scale. Pelagic production exceeded total respiration at the shallow site, but not at the deep site, demonstrating that the shallow domains are net autotrophic but the deep domains are net heterotrophic, even during the period of the spring bloom. If we take into account the potential primary production by benthic microalgae, the estuary as a whole is net autotrophic during spring, net heterotrophic during the nonbloom seasons, and has a balanced net metabolism over a full annual period. The seasonal shift from net autotrophy to heterotrophy during the transition from spring to summer was accompanied by a large shift from dominance by pelagic respiration to dominance by benthic respiration. This suggests that changes in net ecosystem metabolism can reflect changes in the pathways of energy flow in shallow coastal ecosystems.

California

Water quality variability in San Francisco Bay, Some gGeneral lessons from 1996 sampling: 1996 annual report, San Francisco estuary regional monitoring program for trace substances

This report describes the results from the 1996 Regional Monitoring Program for Trace Substances (RMP). It is the fourth Annual Report from the RMP which began in 1993 and attempts to synthesize the most obvious data patterns from the last four years. This report includes data from Base Program monitoring activities, as well as results of Pilot and Special Studies conducted or completed in 1996. Additionally, several articles contributed by RMP investigators and others, are included. These articles provide perspective and insight on important contaminant issues identified by the RMP. This summary addresses which kinds of pollutants measured by the RMP appear to be at levels that warrant concern, what kinds of trends may be discerned, and which stations have consistently shown elevated contaminant levels. The goals or general objectives of the RMP are: 1. To obtain high quality baseline data describing the concentrations of toxic and potentially toxic trace elements and organic contaminants in the water and sediment of the San Francisco Estuary. 2. To determine seasonal and annual trends in chemical and biological water quality in the San Francisco Estuary. 3. To continue to develop a data set that can be used to determine long-term trends in the concentrations of toxic and potentially toxic trace elements and organic contaminants in the water and sediments of the San Francisco Estuary. 4. To determine whether water quality and sediment quality in the Estuary at large are in compliance with objectives established by the Basin Plan (the regulatory planning document used by the Regional Water Quality Control Board). 5. To provide a database on water and sediment quality in the Estuary which is compatible with data being developed in other ongoing studies, including wasteload allocation studies and model development, sediment quality objectives development, in-bay studies of dredged material disposal, Interagency Ecological Program (IEP) water quality studies, primary productivity studies, local effects biomonitoring programs, and state and federal mussel watch programs.

California