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James E. Cloern

Publications and source records attributed to James E. Cloern.

At least 19 recordsLinked to original sources

Chlorophyll trends are negative for lakes but positive for estuarine–coastal waters

Nutrient (nitrogen and phosphorus) pollution is an environmental problem of global concern because overenrichment of water bodies increases phytoplankton biomass and ecosystem metabolism, depletes oxygen in bottom waters, and increases the frequency and intensity of harmful algal blooms. These responses to nutrient pollution have motivated policies to reduce anthropogenic nutrient inputs. However, these policies have not been universally implemented and their success at reducing phytoplankton biomass is confounded by other components of global change that regulate the conversion of nutrients into biomass. These regulating processes themselves are changing in the Anthropocene. Our study is an assessment of changing phytoplankton biomass over the period 2000–2019, using chlorophyll a trends measured in 191 lakes and 159 estuarine-coastal sites. Our results show that phytoplankton biomass has decreased in most lakes, increased in most coastal sites, and the largest trends have been predominantly negative for lakes and positive for coastal sites. These results provide evidence of opposing directions of phytoplankton biomass change between lakes and coastal sites in this recent period of unprecedented global change. Nutrient pollution is a knotty environmental problem, and our study indicates that it might be a particularly challenging problem for ecosystems situated in densely populated landscapes where freshwater and sea water meet. Success at overcoming this challenge will require deeper scientific understanding of changes in processes that regulate the conversion of nutrients into phytoplankton biomass, substantial investments of time and resources to reduce nutrient inputs, and a flexible strategy designed to anticipate and adapt to a changing world.

PNAS

On the human appropriation of wetland primary production

Humans are changing the Earth's surface at an accelerating pace, with significant consequences for ecosystems and their biodiversity. Landscape transformation has far-reaching implications including reduced net primary production (NPP) available to support ecosystems, reduced energy supplies to consumers, and disruption of ecosystem services such as carbon storage. Anthropogenic activities have reduced global NPP available to terrestrial ecosystems by nearly 25%, but the loss of NPP from wetland ecosystems is unknown. We used a simple approach to estimate aquatic NPP from measured habitat areas and habitat-specific areal productivity in the largest wetland complex on the USA west coast, comparing historical and modern landscapes and a scenario of wetland restoration. Results show that a 77% loss of wetland habitats (primarily marshes) has reduced ecosystem NPP by 94%, C (energy) flow to herbivores by 89%, and detritus production by 94%. Our results also show that attainment of habitat restoration goals could recover 12% of lost NPP and measurably increase carbon flow to consumers, including at-risk species and their food resources. This case study illustrates how a simple approach for quantifying the loss of NPP from measured habitat losses can guide wetland conservation plans by establishing historical baselines, projecting functional outcomes of different restoration scenarios, and establishing performance metrics to gauge success.

California

Nutrient status of San Francisco Bay and its management implications

Nutrient enrichment has degraded many of the world’s estuaries by amplifying algal production, leading to hypoxia/anoxia, loss of vascular plants and fish/shellfish habitat, and expansion of harmful blooms (HABs). Policies to protect coastal waters from the effects of nutrient enrichment require information to determine if a water body is impaired by nutrients and if regulatory actions are required. We compiled information to inform these decisions for San Francisco Bay (SFB), an urban estuary where the best path toward nutrient management is not yet clear. Our results show that SFB has high nutrient loadings, primarily from municipal wastewater; there is potential for high algal production, but that production is not fully realized; and SFB is not impaired by hypoxia or recurrent HABs. However, our assessment includes reasons for concern: nitrogen and phosphorus concentrations higher than those in other estuaries impaired by nutrient pollution, chronic presences of multiple algal toxins, a recent increase of primary production, and projected future hydroclimatic conditions that could increase the magnitude and frequency of algal blooms. Policymakers thus face the challenge of determining the appropriate protective policy for SFB. We identify three crucial next steps for meeting this challenge: (1) new research to determine if algal toxins can be reduced through nutrient management, (2) establish management goals as numeric targets, and (3) determine the magnitude of nutrient load reduction required to meet those targets. Our case study illustrates how scientific information can be acquired and communicated to inform policymakers about the status of nutrient pollution, its risks, and strategies for minimizing those risks.

California

Machine learning identifies a strong association between warming and reduced primary productivity in an oligotrophic ocean gyre

Phytoplankton play key roles in the oceans by regulating global biogeochemical cycles and production in marine food webs. Global warming is thought to affect phytoplankton production both directly, by impacting their photosynthetic metabolism, and indirectly by modifying the physical environment in which they grow. In this respect, the Bermuda Atlantic Time-series Study (BATS) in the Sargasso Sea (North Atlantic gyre) provides a unique opportunity to explore effects of warming on phytoplankton production across the vast oligotrophic ocean regions because it is one of the few multidecadal records of measured net primary productivity (NPP). We analysed the time series of phytoplankton primary productivity at BATS site using machine learning techniques (ML) to show that increased water temperature over a 27-year period (1990–2016), and the consequent weakening of vertical mixing in the upper ocean, induced a negative feedback on phytoplankton productivity by reducing the availability of essential resources, nitrogen and light. The unbalanced availability of these resources with warming, coupled with ecological changes at the community level, is expected to intensify the oligotrophic state of open-ocean regions that are far from land-based nutrient sources.

Scientific Reports

Blurred lines: Multiple freshwater and marine algal toxins at the land-sea interface of San Francisco Bay, California

San Francisco Bay (SFB) is a eutrophic estuary that harbors both freshwater and marine toxigenic organisms that are responsible for harmful algal blooms. While there are few commercial fishery harvests within SFB, recreational and subsistence harvesting for shellfish is common. Coastal shellfish are monitored for domoic acid and paralytic shellfish toxins (PSTs), but within SFB there is no routine monitoring for either toxin. Dinophysis shellfish toxins (DSTs) and freshwater microcystins are also present within SFB, but not routinely monitored. Acute exposure to any of these toxin groups has severe consequences for marine organisms and humans, but chronic exposure to sub-lethal doses, or synergistic effects from multiple toxins, are poorly understood and rarely addressed. This study documents the occurrence of domoic acid and microcystins in SFB from 2011 to 2016, and identifies domoic acid, microcystins, DSTs, and PSTs in marine mussels within SFB in 2012, 2014, and 2015. At least one toxin was detected in 99% of mussel samples, and all four toxin suites were identified in 37% of mussels. The presence of these toxins in marine mussels indicates that wildlife and humans who consume them are exposed to toxins at both sub-lethal and acute levels. As such, there are potential deleterious impacts for marine organisms and humans and these effects are unlikely to be documented. These results demonstrate the need for regular monitoring of marine and freshwater toxins in SFB, and suggest that co-occurrence of multiple toxins is a potential threat in other ecosystems where freshwater and seawater mix.

California

Why large cells dominate estuarine phytoplankton

Surveys across the world oceans have shown that phytoplankton biomass and production are dominated by small cells (picoplankton) where nutrient concentrations are low, but large cells (microplankton) dominate when nutrient-rich deep water is mixed to the surface. I analyzed phytoplankton size structure in samples collected over 25 yr in San Francisco Bay, a nutrient-rich estuary. Biomass was dominated by large cells because their biomass selectively grew during blooms. Large-cell dominance appears to be a characteristic of ecosystems at the land–sea interface, and these places may therefore function as analogs to oceanic upwelling systems. Simulations with a size-structured NPZ model showed that runs of positive net growth rate persisted long enough for biomass of large, but not small, cells to accumulate. Model experiments showed that small cells would dominate in the absence of grazing, at lower nutrient concentrations, and at elevated (+5°C) temperatures. Underlying these results are two fundamental scaling laws: (1) large cells are grazed more slowly than small cells, and (2) grazing rate increases with temperature faster than growth rate. The model experiments suggest testable hypotheses about phytoplankton size structure at the land–sea interface: (1) anthropogenic nutrient enrichment increases cell size; (2) this response varies with temperature and only occurs at mid-high latitudes; (3) large-cell blooms can only develop when temperature is below a critical value, around 15°C; (4) cell size diminishes along temperature gradients from high to low latitudes; and (5) large-cell blooms will diminish or disappear where planetary warming increases temperature beyond their critical threshold.

Limnology and Oceanography

The land-sea interface: A source of high-quality phytoplankton to support secondary production

Coastal-estuarine systems are among the most productive marine ecosystems and their special role in producing harvestable fish and shellfish has been attributed to high primary production fueled by nutrient runoff from land and efficient trophic transfer. Here we ask if phytoplankton species composition and their food quality based on the percentage of long-chain essential fatty acids (LCEFA) is another factor contributing to high secondary production in these ecosystems. We used long-term measurements of major phytoplankton taxonomic groups and estimated their content of LCEFA along the salinity gradient in coastal-estuarine ecosystems, with emphasis on Chesapeake Bay and the Baltic Sea, and an oceanic transect. Our data show that cyanobacteria with low nutritional quality often dominate at low-salinity regions, while intermediate to higher salinity regions produce diatoms and dinoflagellates that have a higher content of LCEFA and are thus a higher-quality food resource for consumers. Higher salinity regions have less pronounced seasonal changes in the percentage of phytoplankton LCEFA compared to low salinity regions, providing a stable supply of nutritious phytoplankton to consumers. The phytoplankton LCEFA content is similarly high in coastal upwelling systems and it decreases further offshore in oligotrophic oceanic regions dominated by picophytoplankton. Our results from a broad range of coastal-ecosystem types show that ecosystems at the land-sea interface provide a valuable service by producing phytoplankton enriched in the biochemicals essential for consumers. High primary production, coupled with high quality of that production, explain why the production of fish and shellfish is high where land and sea meet.

Limnology and Oceanography

Water quality measurements in San Francisco Bay by the U.S. Geological Survey, 1969–2015

The U.S. Geological Survey (USGS) maintains a place-based research program in San Francisco Bay (USA) that began in 1969 and continues, providing one of the longest records of water-quality measurements in a North American estuary. Constituents include salinity, temperature, light extinction coefficient, and concentrations of chlorophyll- a , dissolved oxygen, suspended particulate matter, nitrate, nitrite, ammonium, silicate, and phosphate. We describe the sampling program, analytical methods, structure of the data record, and how to access all measurements made from 1969 through 2015. We provide a summary of how these data have been used by USGS and other researchers to deepen understanding of how estuaries are structured and function differently from the river and ocean ecosystems they bridge.

California

Ecosystem variability along the estuarine salinity gradient: Examples from long-term study of San Francisco Bay

The salinity gradient of estuaries plays a unique and fundamental role in structuring spatial patterns of physical properties, biota, and biogeochemical processes. We use variability along the salinity gradient of San Francisco Bay to illustrate some lessons about the diversity of spatial structures in estuaries and their variability over time. Spatial patterns of dissolved constituents (e.g., silicate) can be linear or nonlinear, depending on the relative importance of river-ocean mixing and internal sinks (diatom uptake). Particles have different spatial patterns because they accumulate in estuarine turbidity maxima formed by the combination of sinking and estuarine circulation. Some constituents have weak or no mean spatial structure along the salinity gradient, reflecting spatially distributed sources along the estuary (nitrate) or atmospheric exchanges that buffer spatial variability of ecosystem metabolism (dissolved oxygen). The density difference between freshwater and seawater establishes stratification in estuaries stronger than the thermal stratification of lakes and oceans. Stratification is strongest around the center of the salinity gradient and when river discharge is high. Spatial distributions of motile organisms are shaped by species-specific adaptations to different salinity ranges (shrimp) and by behavioral responses to environmental variability (northern anchovy). Estuarine spatial patterns change over time scales of events (intrusions of upwelled ocean water), seasons (river inflow), years (annual weather anomalies), and between eras separated by ecosystem disturbances (a species introduction). Each of these lessons is a piece in the puzzle of how estuarine ecosystems are structured and how they differ from the river and ocean ecosystems they bridge.

California

Novel analyses of long-term data provide a scientific basis for chlorophyll-a thresholds in San Francisco Bay

San Francisco Bay (SFB), USA, is highly enriched in nitrogen and phosphorus, but has been resistant to the classic symptoms of eutrophication associated with over-production of phytoplankton . Observations in recent years suggest that this resistance may be weakening, shown by: significant increases of chlorophyll- a ( chl-a ) and decreases of dissolved oxygen (DO), common occurrences of phytoplankton taxa that can form Harmful Algal Blooms (HAB), and algal toxins in water and mussels reaching levels of concern. As a result, managers now ask: what levels of chl-a in SFB constitute tipping points of phytoplankton biomass beyond which water quality will become degraded, requiring significant nutrient reductions to avoid impairments? We analyzed data for DO, phytoplankton species composition, chl-a, and algal toxins to derive quantitative relationships between three indicators (HAB abundance, toxin concentrations, DO) and chl-a . Quantile regressions relating HAB abundance and DO to chl-a were significant, indicating SFB is at increased risk of adverse HAB and low DO levels if chl-a continues to increase. Conditional probability analysis (CPA) showed chl-a of 13 mg m −3 as a “protective” threshold below which probabilities for exceeding alert levels for HAB abundance and toxins were reduced. This threshold was similar to chl-a of 13–16 mg m −3 that would meet a SFB-wide 80% saturation Water Quality Criterion (WQC) for DO. Higher “at risk” chl-a thresholds from 25 to 40 mg m −3 corresponded to 0.5 probability of exceeding alert levels for HAB abundance, and for DO below a WQC of 5.0 mg L −1 designated for lower South Bay (LSB) and South Bay (SB). We submit these thresholds as a basis to assess eutrophication status of SFB and to inform nutrient management actions. This approach is transferrable to other estuaries to derive chl-a thresholds protective against eutrophication.

California

Estuary-ocean connectivity: Fast physics, slow biology

Estuaries are connected to both land and ocean so their physical, chemical, and biological dynamics are influenced by climate patterns over watersheds and ocean basins. We explored climate-driven oceanic variability as a source of estuarine variability by comparing monthly time series of temperature and chlorophyll- a inside San Francisco Bay with those in adjacent shelf waters of the California Current System (CCS) that are strongly responsive to wind-driven upwelling. Monthly temperature fluctuations inside and outside the Bay were synchronous, but their correlations weakened with distance from the ocean. These results illustrate how variability of coastal water temperature (and associated properties such as nitrate and oxygen) propagates into estuaries through fast water exchanges that dissipate along the estuary. Unexpectedly, there was no correlation between monthly chlorophyll- a variability inside and outside the Bay. However, at the annual scale Bay chlorophyll- a was significantly correlated with the Spring Transition Index (STI) that sets biological production supporting fish recruitment in the CCS. Wind forcing of the CCS shifted in the late 1990s when the STI advanced 40 days. This shift was followed, with lags of 1–3 years, by 3- to 19-fold increased abundances of five ocean-produced demersal fish and crustaceans and 2.5-fold increase of summer chlorophyll- a in the Bay. These changes reflect a slow biological process of estuary–ocean connectivity operating through the immigration of fish and crustaceans that prey on bivalves, reduce their grazing pressure, and allow phytoplankton biomass to build. We identified clear signals of climate-mediated oceanic variability in this estuary and discovered that the response patterns vary with the process of connectivity and the timescale of ocean variability. This result has important implications for managing nutrient inputs to estuaries connected to upwelling systems, and for assessing their responses to changing patterns of upwelling timing and intensity as the planet continues to warm.

California

Primary production in the Delta: Then and now

To evaluate the role of restoration in the recovery of the Delta ecosystem, we need to have clear targets and performance measures that directly assess ecosystem function. Primary production is a crucial ecosystem process, which directly limits the quality and quantity of food available for secondary consumers such as invertebrates and fish. The Delta has a low rate of primary production, but it is unclear whether this was always the case. Recent analyses from the Historical Ecology Team and Delta Landscapes Project provide quantitative comparisons of the areal extent of 14 habitat types in the modern Delta versus the historical Delta (pre-1850). Here we describe an approach for using these metrics of land use change to: (1) produce the first quantitative estimates of how Delta primary production and the relative contributions from five different producer groups have been altered by large-scale drainage and conversion to agriculture; (2) convert these production estimates into a common currency so the contributions of each producer group reflect their food quality and efficiency of transfer to consumers; and (3) use simple models to discover how tidal exchange between marshes and open water influences primary production and its consumption. Application of this approach could inform Delta management in two ways. First, it would provide a quantitative estimate of how large-scale conversion to agriculture has altered the Delta's capacity to produce food for native biota. Second, it would provide restoration practitioners with a new approach—based on ecosystem function—to evaluate the success of restoration projects and gauge the trajectory of ecological recovery in the Delta region.

California

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.

California

Human activities and climate variability drive fast-paced change across the world's estuarine-coastal ecosystems

Time series of environmental measurements are essential for detecting, measuring and understanding changes in the Earth system and its biological communities. Observational series have accumulated over the past 2–5 decades from measurements across the world's estuaries, bays, lagoons, inland seas and shelf waters influenced by runoff. We synthesize information contained in these time series to develop a global view of changes occurring in marine systems influenced by connectivity to land. Our review is organized around four themes: (i) human activities as drivers of change; (ii) variability of the climate system as a driver of change; (iii) successes, disappointments and challenges of managing change at the sea-land interface; and (iv) discoveries made from observations over time. Multidecadal time series reveal that many of the world's estuarine–coastal ecosystems are in a continuing state of change, and the pace of change is faster than we could have imagined a decade ago. Some have been transformed into novel ecosystems with habitats, biogeochemistry and biological communities outside the natural range of variability. Change takes many forms including linear and nonlinear trends, abrupt state changes and oscillations. The challenge of managing change is daunting in the coastal zone where diverse human pressures are concentrated and intersect with different responses to climate variability over land and over ocean basins. The pace of change in estuarine–coastal ecosystems will likely accelerate as the human population and economies continue to grow and as global climate change accelerates. Wise stewardship of the resources upon which we depend is critically dependent upon a continuing flow of information from observations to measure, understand and anticipate future changes along the world's coastlines.

Global Change Biology

Estuarine fish communities respond to climate variability over both river and ocean basins

Estuaries are dynamic environments at the land–sea interface that are strongly affected by interannual climate variability. Ocean–atmosphere processes propagate into estuaries from the sea, and atmospheric processes over land propagate into estuaries from watersheds. We examined the effects of these two separate climate-driven processes on pelagic and demersal fish community structure along the salinity gradient in the San Francisco Estuary, California, USA. A 33-year data set (1980–2012) on pelagic and demersal fishes spanning the freshwater to marine regions of the estuary suggested the existence of five estuarine salinity fish guilds: limnetic (salinity = 0–1), oligohaline (salinity = 1–12), mesohaline (salinity = 6–19), polyhaline (salinity = 19–28), and euhaline (salinity = 29–32). Climatic effects propagating from the adjacent Pacific Ocean, indexed by the North Pacific Gyre Oscillation (NPGO), affected demersal and pelagic fish community structure in the euhaline and polyhaline guilds. Climatic effects propagating over land, indexed as freshwater outflow from the watershed (OUT), affected demersal and pelagic fish community structure in the oligohaline, mesohaline, polyhaline, and euhaline guilds. The effects of OUT propagated further down the estuary salinity gradient than the effects of NPGO that propagated up the estuary salinity gradient, exemplifying the role of variable freshwater outflow as an important driver of biotic communities in river-dominated estuaries. These results illustrate how unique sources of climate variability interact to drive biotic communities and, therefore, that climate change is likely to be an important driver in shaping the future trajectory of biotic communities in estuaries and other transitional habitats.

California

Phytoplankton blooms in estuarine and coastal waters: Seasonal patterns and key species

Phytoplankton blooms are dynamic phenomena of great importance to the functioning of estuarine and coastal ecosystems. We analysed a unique (large) collection of phytoplankton monitoring data covering 86 coastal sites distributed over eight regions in North America and Europe, with the aim of investigating common patterns in the seasonal timing and species composition of the blooms. The spring bloom was the most common seasonal pattern across all regions, typically occurring early (February–March) at lower latitudes and later (April–May) at higher latitudes. Bloom frequency, defined as the probability of unusually high biomass, ranged from 5 to 35% between sites and followed no consistent patterns across gradients of latitude, temperature, salinity, water depth, stratification, tidal amplitude or nutrient concentrations. Blooms were mostly dominated by a single species, typically diatoms (58% of the blooms) and dinoflagellates (19%). Diatom-dominated spring blooms were a common feature in most systems, although dinoflagellate spring blooms were also observed in the Baltic Sea. Blooms dominated by chlorophytes and cyanobacteria were only common in low salinity waters and occurred mostly at higher temperatures. Key bloom species across the eight regions included the diatoms Cerataulina pelagica and Dactyliosolen fragilissimus and dinoflagellates Heterocapsa triquetra and Prorocentrum cordatum . Other frequent bloom-forming taxa were diatom genera Chaetoceros , Coscinodiscus , Skeletonema , and Thalassiosira . Our meta-analysis shows that these 86 estuarine-coastal sites function as diatom-producing systems, the timing of that production varies widely, and that bloom frequency is not associated with environmental factors measured in monitoring programs. We end with a perspective on the limitations of conclusions derived from meta-analyses of phytoplankton time series, and the grand challenges remaining to understand the wide range of bloom patterns and processes that select species as bloom dominants in coastal waters.

Estuarine, Coastal and Shelf Science