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Nutrient and carbon limitation drive broad-scale patterns of mixotrophy in the ocean

Posted by mmaheigan

· Tuesday, May 14th, 2019

In the ocean, unicellular eukaryotes are often mixotrophic, which means they photosynthesize and also consume prey. In recent decades, it has become clear that mixotrophs are ubiquitous in sunlit ocean habitats. Additionally, models predict that mixotrophs have important impacts on productivity, nutrient cycling, carbon export, and food web structure. However, there is little understanding of the environmental conditions that select for a mixotrophic lifestyle, and it is unclear how mixotrophs succeed in competition with autotrophic and heterotrophic specialists. A recent study in PNAS that synthesized measurements of mixotrophic nanoflagellates showed that mixotrophs are more abundant in stratified, well-lit, low latitude environments (Figure 1A). They are also more abundant, relative to pure heterotrophs, in productive coastal environments (Figure 1B). A trait-based model analysis revealed that the success of mixotrophs depends on the fact that they are less nutrient-limited than autotrophs (due to prey-derived nutrients) and less carbon-limited than heterotrophs (due to photosynthesis). This synergy requires sufficient light, leading to success in low latitude environments. Similarly, a greater supply of dissolved nutrients relative to prey, as commonly observed in coastal environments, favors mixotrophs relative to heterotrophs. One implication of these results is that carbon fixation at lower latitudes may be enhanced by mixotrophy, while limiting nutrients may be more efficiently transferred to higher trophic levels.

Figure 1. Estimated abundance of autotrophic, mixotrophic, and heterotrophic nanoflagellates across environmental gradients in the ocean.

Author:
Kyle Edwards (Univ. Hawaii at Manoa)

Impacts of atmospheric nitrogen deposition and coastal nitrogen fluxes on oxygen concentrations in Chesapeake Bay

Posted by mmaheigan

· Tuesday, April 30th, 2019

How do atmospheric and oceanic nutrients impact oxygen concentrations in the Chesapeake Bay? Generally, researchers focus on how terrestrial nutrients impact hypoxia. The relative importance of river, atmosphere, and ocean inputs have not been quantified, largely because estimates of nitrogen fluxes from the atmosphere and ocean are limited.

A recent study in Journal of Geophysical Research: Oceans quantified the relative impacts of atmospheric and oceanic nitrogen inputs on dissolved oxygen (DO) in the Chesapeake Bay. The authors combined a 3-D biogeochemical model and estimates of atmospheric deposition from the Community Multiscale Air Quality model and interpolations of nitrogen concentrations along the continental shelf from the Ocean Acidification Data Stewardship Project. Atmospheric nitrogen deposition and coastal nitrogen fluxes most impact Chesapeake Bay DO concentrations during the summer when surface waters are depleted in nitrogen. Overall, atmospheric nitrogen deposition has about the same gram-for-gram impact on Chesapeake Bay DO as riverine loading. Although all three nutrient sources vary spatially and temporally, in the central bay, where summer hypoxia is most prevalent, coastal nitrogen fluxes and atmospheric nitrogen fluxes have roughly the same impact on bottom oxygen as a ~10% change in riverine nitrogen loading (Figure 1).

Figure caption: (Left) Four-year (2002–2005) average increase in DO in the summer by removing the atmospheric nitrogen deposition (AtmN), reducing the riverine loading (ΔRiverN) by ~10% (roughly equivalent to turning off the atmospheric deposition), and removing the nitrogen fluxes from the continental shelf (CoastalN). (Right) Relative impacts of the three nitrogen modification scenarios on summertime bottom DO.

These results indicate that two often-neglected sources of nitrogen—direct atmospheric deposition and fluxes of nitrogen from the continental shelf—substantially impact Chesapeake Bay DO, especially in the summer. Future study is needed to investigate the long-term trend of these relative impacts by continued coordination between modeling and observational work, such as applying higher-resolution atmospheric deposition products and integrating more in situ data along the model ocean boundary when they are available. These efforts will improve our understanding of the impacts of different nutrient sources on biogeochemical cycles in coastal water bodies.

Authors:
Fei Da (VIMS, College of William & Mary)
Marjorie A. M. Friedrichs (VIMS, College of William & Mary)
Pierre St-Laurent (VIMS, College of William & Mary)

When it comes to carbon export, the mesoscale matters

Posted by hbenway

· Tuesday, September 11th, 2018

Figure 1. Difference in annual mean carbon export (ΔPOC flux) between a high resolution (0.1º, Hi-res) and standard resolution (1º, Analog) global climate model simulation using the CESM model. Highlighted regions show areas where vertical (purple boxes) and horizontal (red boxes) changes in nutrient transport drive increases or decreases in export, respectively.

Most Earth System models (ESMs) that are used to study global climate and the carbon cycle do not resolve the most energetic scales in the ocean, the mesoscale (10-100 km), encompassing eddies, coastal jets, and other dynamic features strongly affecting nutrient delivery, productivity, and carbon export. This prompts the question: What are we missing in climate models by not resolving the mesoscale?

Authors of a recent study published in Global Biogeochemical Cycles conducted a comparative analysis of the importance of mesoscale features in biological production and associated carbon export using standard resolution (1°) and mesoscale-resolving (0.1°) ESM simulations. The mesoscale-resolving ESM yielded only a ~2% reduction in globally integrated export production relative to the standard resolution ESM. However, a closer look at the local processes driving export in different basins revealed much larger, compensating differences (Fig. 1). For example, in regions where biological production is driven by natural iron fertilization from shelf sediment sources (Fig. 2), improved representation of coastal jets in the higher-resolution ESM reduces the cross-shelf iron delivery that fuels production (red boxes in Fig. 1). Resolving mesoscale turbulence further reduces the spatial extent of blooms and associated export, yielding a more patchy distribution than in the coarse resolution models. Together, these processes lead to a reduction in export in the Argentine Basin, one of the most productive regions on the planet, of locally up to 50%. In contrast, resolving the mesoscale results in enhanced export production in the Subantarctic (purple box in Fig. 1), where the mesoscale model resolves deeper, narrower mixed layer depths that support stronger nutrient entrainment, in turn enhancing local productivity and export.

Figure 2. An iron-driven plankton bloom structured by mesoscale features in the South Atlantic. Left is simulated dissolved iron (Fe), the limiting nutrient for this region, and right is iron in all phytoplankton classes, a proxy for biomass (phytoFe, shown in log10 scale), on January 11, the height of the bloom. Plankton blooms in the Subantarctic Atlantic are fueled by horizontal iron transport off coastal and island shelves and vertical injection from seamounts, whereas farther south in the Southern Ocean, winter vertical mixing is the primary driver of iron delivery. Mesoscale circulation, largely an unstructured mix of interacting jets and vortices, strongly affects the location and timing of carbon production and export. Click here for an animation.

In regions with very short productivity seasons like the North Pacific and Subantarctic, internally generated mesoscale variability (captured in the higher resolution ESM) yields significant interannual variation in local carbon export. In these regions, a few eddies, filaments or more amorphous mesoscale features can structure the entire production and export pattern for the short bloom season. These findings document the importance of resolving mesoscale features in ESMs to more accurately quantify carbon export, and the different roles mesoscale variability can play in different oceanographic settings.

Determining how to best sample these mesoscale turbulence-dominated blooms and scale up these measurements to regional and longer time means, is an outstanding joint challenge for modelers and observationalists. A key piece is obtaining the high temporal and spatial resolution data sets needed for validating modeled carbon export in bloom regions strongly impacted by mesoscale dynamics, which represent a large portion of the global carbon export.

Authors
Cheryl Harrison (NCAR, University of Colorado Boulder)
Matthew Long (NCAR)
Nicole Lovenduski (University of Colorado Boulder)
J. Keith Moore (University of California Irvine)

Shelf-wide pCO2 increase across the South Atlantic Bight

Posted by mmaheigan

· Thursday, August 2nd, 2018

Relative to their surface area, coastal regions represent some of the largest carbon fluxes in the global ocean, driven by numerous physical, chemical and biological processes. Coastal systems also experience human impacts that affect carbon cycling, which has large socioeconomic implications. The highly dynamic nature of these systems necessitates observing approaches and numerical methods that can both capture high-frequency variability and delineate long-term trends.

Figure 1: The South Atlantic Bight (SAB) was divided into four sections using isobaths: the coastal zone (0 to 15 m), the inner shelf (15 to 30 m), the middle shelf (30 to 60 m), and the outer shelf (60 m and beyond). The X’s indicate the locations of the Gray’s Reef mooring (southern X) and the Edisto mooring (northern X).

In two recent studies using mooring- and ship-based ocean CO₂ system data, authors observed that pCO₂ is increasing from the coastal zone to the outer shelf of the South Atlantic Bight at rates greater than the global average oceanic and atmospheric increase (~1.8 µatm y^-1). In recent publications in Continental Shelf Research and JGR-Oceans, the authors analyzed pCO₂ data from 46 cruises (1991-2016) using a novel linear regression technique to remove the seasonal signal, revealing an increase in pCO₂ of 3.0-3.7 µatm y^-1on the outer and inner shelf, respectively. Using a Generalized Additive Mixed Model (GAMM) approach for trend analysis, authors observed that the rates of increase were slightly higher than the deseasonalization technique, yielding pCO₂ increases of 3.3 to 4.5 µatm y^-1 on the outer and inner shelf, respectively. The reported pCO₂ increases result in potential pH decreases of -0.003 to -0.004 units y^-1.

Figure 2: The time series of fCO2 in the four regions of the SAB (cruise observations) and from the Gray’s Reef mooring on the inner shelf indicate an increase across the shelf. These data are the observed values, however, the trend lines for each time series are calculated using deseasonalized values using the reference year method.

Analysis of the pCO₂ time-series from the Gray’s Reef mooring (using a NOAA Moored Autonomous pCO₂ system from July 2006 -July 2015) yielded a rate of increase (3.5 ± 0.9 µatm y^-1) that was comparable to the cruise data on the inner shelf (3.7 ± 2.2 and 4.5 ± 0.6 µatm y^-1, linear and GAMM methods, respectively). Validation data collected at the mooring suggest that underway data from cruises and the moored data are comparable. Neither thermal processes nor atmospheric dissolution (the primary driver of oceanic acidification) can explain the observed pCO₂ increase and concurrent pH decrease across the shelf. Unlike the middle and outer shelves, where an increase in SST could account for up to 1.1 µatm y^-1 of the observed pCO₂ trend, there is no thermal influence in the coastal zone and inner shelf. While 1.8 µatm y^-1 could be attributed to the global average atmospheric increase, the remainder is likely due to transport from coastal marshes and in situ biological processes. As the authors have shown, the increasing coastal and oceanic trend in pCO₂ can lead to a decrease in pH, especially if there is no increase in buffering capacity. More acidic waters can have a long term affect on coastal ecosystem services and biota.

Also see Eos Editor’s Vox on this research by Peter Brewer https://eos.org/editors-vox/coastal-ocean-warming-adds-to-co2-burden

Authors:

Multidecadal fCO2 Increase Along the United States Southeast Coastal Margin (JGR-Oceans)
Janet J. Reimer (University of Delaware)
Hongjie Wang (Texas A &M University – Corpus Christi)
Rodrigo Vargas (University of Delaware)
Wei-Jun Cai (University of Delaware)

And

Time series pCO2 at a coastal mooring: Internal consistency, seasonal cycles, and interannual variability (Continental Shelf Research)
Janet J. Reimer (University of Delaware)
Wei-Jun Cai (University of Delaware; University of Georgia)
Liang Xue (University of Delaware; First Institute of Oceanography, China)
Rodrigo Vargas (University of Delaware)
Scott Noakes (University of Georgia)
Xinping Hu (Texas A &M University – Corpus Christi)
Sergio R. Signorini (Science Applications International Corporation)
Jeremy T. Mathis (NOAA Arctic Research Program)
Richard A. Feely (NOAA Pacific Marine Environmental Laboratory)
Adrienne J. Sutton (NOAA Pacific Marine Environmental Laboratory; University of Washington)
Christopher Sabine (University of Hawaii Manoa)
Sylvia Musielewicz (NOAA Pacific Marine Environmental Laboratory; University of Washington)
Baoshan Chen (University of Delaware; University of Georgia)
Rik Wanninkhof (NOAA Atlantic Oceanographic and Meteorological Laboratory)

Long-term coastal data sets reveal unifying relationship between oxygen and pH fluctuations

Posted by mmaheigan

· Thursday, June 7th, 2018

Coastal habitats are critically important to humans, but without consistent and reliable observations we cannot understand the direction and magnitude of unfolding changes in these habitats. Environmental monitoring is therefore a prescient—yet still undervalued—societal service, and no effort better exemplifies this than the work conducted within the National Estuarine Research Reserve System (NERRS). NERRS is a network of 29 U.S. estuarine sites operated as a partnership between NOAA and the coastal states. NERRS has established a system-wide monitoring program with standardized instrumentation, protocols, and data reporting to guide consistent and comparable data collection across all NERRS sites. This has resulted in high-quality, comparable data on short- to long-term changes in water quality and biological systems to inform effective coastal zone management.

Figure 1: Using dissolved oxygen and salinity, monthly mean pH can be predicted within and across coastal systems due to the unifying metabolic coupling of oxygen and pH.

In a recent study published in Estuaries and Coasts, Baumann and Smith (2017) used a subset of this unique data set to analyze short- and long-term variability in pH and dissolved oxygen (DO) at 16 NERRS sites across the U.S. Atlantic, Caribbean, Gulf of Mexico, and Pacific coasts (> 5 million data points). They observed that large, metabolically driven fluctuations of pH and DO are indeed a unifying feature of nearshore habitats. Furthermore, mean pH or mean diel pH fluctuations can be predicted across habitats simply from salinity and oxygen levels/fluctuations (Fig.1). These results provide strong empirical evidence that common metabolic principles drive diel to seasonal pH and DO variations within and across a diversity of estuarine environments. As expected, the study did not yield interannual, monotonic trends in nearshore pH conditions; rather, interannual fluctuations were of similar magnitude to the pH decrease predicted for the average surface ocean over the next three centuries (Fig.2). Correlations of weekly anomalies of pH, oxygen, and temperature yielded strong empirical support for the hypothesis that coastal acidification—in addition to being driven by eutrophication and atmospheric CO₂ increases—is exacerbated by warming, likely via increased community respiration.

Figure 2: Interannual variations in temperature, pH, and dissolved oxygen (DO) anomalies in 16 NERRS sites across the US Atlantic, Gulf of Mexico, Caribbean, and Pacific coasts.

Analyses of these long-term data sets have provided important insights on biogeochemical variability and underlying drivers in nearshore environments, highlighting the value and utility of long-term monitoring efforts like NERRS. Sustained, high-quality data sets in these nearshore environments are essential for the study of environmental change and should be prioritized by funding agencies. The observed metabolically driven pH and DO fluctuations suggest that local measures to reduce nutrient pollution can be an effective management tool in support of healthy coastal environments, a boon for both the habitats and humans.

Authors:
Hannes Baumann (University of Connecticut)
Erik M. Smith (North Inlet-Winyah Bay National Estuarine Research Reserve, University of South Carolina)

Seagrass carbon dynamics: Gulf of Mexico

Posted by mmaheigan

· Thursday, March 1st, 2018

Seagrasses have died-off in great numbers, resulting in the release of stored carbon. Seagrasses represent a substantive and relatively unconstrained North American and Caribbean Sea blue carbon sink in the tropical Western Hemisphere. Fine-scale estimates of regional seagrass carbon stocks, as well as carbon fluxes from anthropogenic disturbances and natural processes and gains in sedimentary carbon from seagrass restoration are currently lacking for the bulk of tropical Western Hemisphere seagrass systems.

To address this knowledge gap, in the subtropics and tropics, a recent study yielded estimates of organic carbon (C_org) stocks, losses, and restoration gains from several seagrass beds around the Gulf of Mexico (GoM). GoM-wide seagrass natural C_org stocks were estimated to be ~37.2–37.5Tg C_org. A unique method involving quadruplicate sampling in naturally-occurring, restored, continually-historically barren, and previously-disturbed-now-barren sites provided the first available C_org loss measurements for subtropical-tropical seagrasses. GoM C_org losses were slow, occurring over multiple years, and differed between sites, depending on disturbance type. Mean restored seagrass bed C_org stocks exceeded those of natural seagrass beds, underscoring the importance of seagrass restoration as a viable carbon sequestration strategy. For restored seagrass areas, the older the restoration site, the greater the C_orgstock.

Organic carbon stocks for Gulf of Mexico sediments for the top 20 cm of sediment in always barren, impacted barren, natural seagrass, and restored seagrass sites. Natural and restored seagrass beds had significantly higher organic carbon stocks than impacted barren or always barren sediments.

Seagrass restoration appears to be an important tool for climate-change mitigation. In the USA and throughout the tropics and subtropics, restoration could reduce sedimentary carbon leakage and bolster total blue carbon stores, while facilitating increased fisheries and shoreline stability. Although well-planned and executed restoration of seagrass is more difficult than mangroves or marshes, there are >1 million hectares of degraded seagrass habitats that could be restored, which would greatly increase blue carbon sinks and support diverse marine species that rely on seagrass for habitat and food.

Authors:
Anitra Thorhaug (Yale School of Forestry)
Helen M. Poulos (Earth Sci., Wesleyan Univ.)
Jorge López-Portillo (Inecol, Mexico)
Timothy C.W. Ku (Earth Sci., Wesleyan Univ.)
Graeme P. Berlyn (Yale School of Forestry)

Untangling the mystery of domoic acid events: A climate-scale perspective

Posted by mmaheigan

· Thursday, August 3rd, 2017

The diatom Pseudo-nitzchia produces a neurotoxin called domoic acid, which in high concentrations affects wildlife ranging from mussels and crabs to seabirds and sea lions, as well as humans. In humans, the effects of domoic acid poisoning can range from gastrointestinal distress to memory loss, and even death. Despite being studied in laboratories since the late 1980s, there is no consensus on the environmental conditions that lead to domoic acid events. These events are most frequent and impactful in eastern boundary current regions such as the California Current System, which is bordered by Washington, Oregon, and California. In Oregon alone, there have been six major domoic acid events: 1996, 1998-1999, 2001, 2002-2006, 2010, 2014-2015. McKibben et al. (2017) investigated the regulation of domoic acid at a climate scale to develop and test an applied risk model for the US West Coast” to read “McKibben et al. (2017) investigated the regulation of domoic acid at regional and decadal scales in order to develop and test an applied risk model for the impact of climate on the US West Coast. They used the PDO and ONI climate variability indices, averages of monthly and 3 month running means of SST anomaly values and variability to look at basin-scale ocean conditions. At a local scale, data were from zooplankton sampling every two to four weeks between 1996 to 2015 at hydrographic station offshore of Newport, OR. Additionally, the NOAA NCDC product “Daily Optimum Interpolation, Advanced Very High Resolution Radiometer Only, Version 2, Final+Preliminary SST” was used to obtain the monthly SST anomaly metric, based on combined in situ and satellite data.

(A) Warm and cool ocean regimes, (B) local SST anomaly, and (C and D) biological response. (A) PDO (red or blue vertical bars) and ONI (black line) indices; strong (S) to moderate (M) El Nino (+1) and La Nina (−1) events are labeled. (B) SST anomaly 20 nm off central OR. (C) The CSR anomaly 5 nm off central OR. (D) Monthly OR coastal maximum DA levels in razor clams (vertical bars); horizontal black line is the 20-ppm closure threshold. Black line in D shows the spring biological transition date (right y axis). At the top of the figure, black boxes indicate the duration of upwelling season each year; red vertical bars indicate the timing of annual DA maxima in relationship to upwelling. Gray shaded regions are warm regimes based on the PDO. Dashed vertical lines indicate onset of the six major DA events. The September 2014 arrival of the NE Pacific Warm Anomaly (colloquially termed “The Blob”) to the OR coastal region is labeled on B. “X” symbols along the x axes indicate that no data were available for that month (B–D).

Their findings show that these events have occurred when there is advection of warmer water masses onto the continental shelf from southern or offshore areas. When the warm phase of the Pacific Decadal Oscillation (PDO) and El Niño coincide, the effect is additive. In the warm regime years, there is a later spring biological transition date, weaker alongshore currents, elevated water temperatures, and plankton communities are dominated by subtropical rather than subarctic species. The authors also note relative differences between the prevalence and phenology of domoic acid events in OR, CA and WA, which warrants further study via regional-scale modeling. Overall, this research shows a clear and enhanced risk of toxicity in shellfish during warm phases of natural climate oscillations. If predictions of more extreme warming come to bear, this would potentially lead to increased DA event intensity and frequency in coastal zones around the globe. This will not only affect wildlife, but may cause significant closures of economically important fisheries (e.g., Dungeness crab, anchovy, mussel, and razor clam), which would impact local communities and native populations.

Authors:
Morgaine McKibben (Oregon State Univ., NOAA Northwest Fisheries Science Center)
William Peterson (NOAA Northwest Fisheries Science Center)
Michelle Wood (Univ. Oregon)
Vera L. Trainer (NOAA Northwest Fisheries Science Center)
Matthew Hunter (Oregon Dept. Fish & Wildlife)
Angelicque E. White (Oregon State Univ.)

Do rivers supply nutrients to the open ocean?

Posted by mmaheigan

· Wednesday, May 24th, 2017

Rivers carry large amounts of nutrients (e.g., nitrogen and phosphorus) to the sea, but we do not know how much of that riverine nutrient supply escapes biological and chemical processing in shallow coastal waters to reach the open ocean. Most global ocean biogeochemical models, which are typically unable to resolve coastal processes, assume that either all or none of the riverine nutrients entering coastal waters actually contribute to open ocean processes.

While we know a good deal about the dynamics of individual rivers entering the coastal ocean, studies to date have been limited to a few major river systems, mainly in in developed countries. Globally, there are over 6,000 rivers entering the coastal ocean. In a recent study, Sharples et al (2017) devised a simple approach to obtain a global-scale estimate of riverine nutrient inputs based on the knowledge that low-salinity waters entering the coastal ocean tend to form buoyant plumes that turn under the influence of Earth’s daily rotation to flow along the coastline. Using published data on such flows and incorporating the effect of Earth’s rotation, they obtained estimates of typical cross-shore plume width and compared them to the local width of the continental shelf. This was used to calculate the residence time of riverine nutrients on the shelf, which is the key to estimating how much of a given nutrient is consumed in shelf waters vs. how much is exported to the open ocean.

Global distribution of the amount of riverine dissolved inorganic nitrogen that escapes the continental shelf to reach the open ocean.

The results indicate that, on a global scale, 75% (80%) of the nitrogen (phosphorus) supplied by rivers reaches the open ocean, whereas 25% (20%) of the nitrogen (phosphorus) is consumed on the shelf (e.g., fueling coastal productivity). Limited knowledge of nutrient cycling and consumption in shelf waters represents the primary source of uncertainty in this study. However, well-defined global patterns related to human land use (e.g., agricultural fertilizer use in developed nations) emerged from this analysis, underscoring the need to understand how land-use changes and other human activities will alter nutrient delivery to the coastal ocean in the future.

Authors:
Jonathan Sharples (School of Environmental Sciences, University of Liverpool, UK)
Jack Middelburg (Department of Earth Sciences, Utrecht University, Netherlands)
Katja Fennel (Department of Oceanography, Dalhousie University, Canada)
Tim Jickells (School of Environmental Sciences, University of East Anglia, UK)

Scientists reveal major drivers of aragonite saturation state in the Gulf of Maine, a region vulnerable to acidification

Posted by mmaheigan

· Thursday, May 11th, 2017

The Gulf of Maine (GoME) is a shelf region that is especially vulnerable to ocean acidification (OA). GoME’s shelf waters display the lowest mean pH, aragonite saturation state (Ω-Ar), and buffering capacity of the entire U.S. East Coast. These conditions are a product of many unique characteristics and processes occurring in the GoME, including relatively low water temperatures that result in higher CO₂ solubility; inputs of fresher, low-alkalinity water that is traceable to the rivers discharging into the Labrador Sea to the north, as well as local inputs of low-pH river water; and its semi-enclosed nature (long residence time >1 year), which enables the accumulation of respiratory products, i.e. CO₂.

A recent study by Wang et al. (2017) is the first to assess the major oceanic processes controlling seasonal variability of aragonite saturation state and its linkages with pteropod abundance in the GoME. The results indicate that surface production was tightly coupled with remineralization in the benthic nepheloid layer during highly productive seasons, resulting in occasional aragonite undersaturation. Mean water column Ω-Ar and abundance of large thecosomatous pteropods show some correlation, although discrete cohort reproductive success likely also influences their abundance. Photosynthesis-respiration is the primary driving force controlling Ω-Ar variability over the seasonal cycle. However, calcium carbonate (CaCO₃) dissolution appears to occur at depth in fall and winter months when bottom water Ω-Ar is generally low but slightly above 1. This is accompanied by a decrease in pteropod abundance that is consistent with previous CaCO₃ flux trap measurements.

Figure. Changes of aragonite saturation states (ΔΩ) between three consecutive cruises from April – July 2015 as a function of changes in salinity-normalized DIC (ΔenDIC, including correction of freshwater inputs) (a) and changes in salinity-normalized TA (ΔenTA, including correction of freshwater inputs) (b). The data points circled in (b) represent potential alkalinity sources from CaCO3 dissolution and/or anaerobic respiration. Solid lines are theoretical lines of ΔΩ vs. ΔenDIC and ΔΩ vs. ΔenTA expected if only photosynthesis and respiration/remineralization occur. Dashed lines are theoretical lines if only calcification and dissolution of CaCO3 occur.

Under the current rate of OA, the mean Ω-Ar of the subsurface and bottom waters of the GoME will approach undersaturation (Ω-Ar < 1) in 30-40 years. As photosynthesis and respiration are the major driving mechanisms of Ω-Ar variability in the water column, any biological regime changes may significantly impact carbonate chemistry and the GoME ecosystem, including the CaCO₃ shell-building capacity of organisms that are critical to the GoME food web.

Author:

Zhaohui Aleck Wang (Woods Hole Oceanographic Institution)

Impact of ENSO on biogeochemistry and lower trophic level response in the California Current System

Posted by mmaheigan

· Thursday, February 16th, 2017

El Niño events are one of the “most spectacular instances of interannual variability in the ocean” with “profound consequences for climate and the ocean ecosystem” (Cane 1986). Perturbations in the atmosphere directly influence the ocean with long-term effects on environmental variability in the tropical Pacific Ocean as the El Niño-Southern Oscillation (ENSO) shifts between El Niño, neutral, and La Niña states on a timescale of two to seven years. On longer timescales, teleconnections from the tropics to extratropical regions drive Pacific decadal variability, and these can be both oceanic and atmospheric in nature. Mid-latitude variability of the Pacific Decadal Oscillation (PDO) has been associated with ENSO (e.g., Newman et al. 2003) and is distinguished from ENSO in part by its multidecadal timescale (20-30 years; 50 years). The PDO is dependent upon ENSO as a response to the combined effects of atmospheric noise (Newman et al. 2003), as well as the asymmetry of the ENSO cycle (Rodgers et al. 2004). Therefore, when discussing decadal variability of the northeast Pacific, we are referring to the delayed impacts of ENSO.

Ecosystem impacts of northeast Pacific variability

Given the complex influence of tropical climate on northeast Pacific ecosystems, there is significant overlap between ENSO signals and higher frequency modes of the PDO Index. It is widely recognized that interactions between these two climate modes drive substantial ecosystem variability on a range of time and space scales. Large regime shifts in the North Pacific that have reverberated throughout the ecosystem, from physics to fish, are recurring patterns now associated with low-frequency changes in sea surface temperature (SST) that characterize the PDO (e.g., Mantua et al. 1997). Some of the higher-frequency fluctuations in ecosystem variables of the northeast Pacific that have not been successfully attributed to PDO or ENSO are now thought to be driven by an intermediate mode of variability called the North Pacific Gyre Oscillation (NPGO). While the PDO is characterized as the first empirical orthogonal function (EOF) of SST, NPGO is defined as the first EOF of both SST and sea surface height (SSH) anomalies (Di Lorenzo et al. 2008). Compared to PDO and ENSO, NPGO is more closely tied to variations in salinity, nutrient upwelling, and chlorophyll a (chl-a) in the long-running California Cooperative Oceanic Fisheries Investigations (CalCOFI) time-series. DiLorenzo et al. (2008) suggest that major ecosystem regime shifts require a simultaneous and opposite sign reversal of the NPGO and PDO, as was seen shortly after the massive ENSO event of 1997/98, and all three indices relate back to dynamics in the tropical Pacific. The struggle is to understand how these low- to high(er)-frequency modes of variability in the climate and physics drive fluctuations in biogeochemistry and coastal ecology.

As discussed by Jacox et al. (this post), there is an expected or canonical set of physical conditions associated with ENSO in the California Current System (CCS). This physical response to ENSO generally includes: 1) changes in surface wind stress that alter the strength of coastal upwelling and downwelling; 2) remote oceanic forcing by coastally trapped waves that propagate poleward along the US West Coast and modify thermocline depth and coastal stratification; and 3) changes to alongshore advection (Jacox et al., this post). The ecological response of the coastal marine environment includes changes in primary production and the community composition of plankton and higher trophic levels that can be directly or more subtly related to these physical factors. Primary production is driven by vertical nutrient flux to well-lit surface waters; nutrient supply is related to upwelling magnitude, upwelling source depth, and nutrient concentrations at the source depth. ENSO-related processes are also important for interannual and seasonal variability of oxygen concentrations and carbonate biogeochemistry on the Washington and Oregon Shelves (Siedlecki et al. 2015). In this article, we highlight some of the modeling and observational studies that have successfully attributed ENSO-like variability to specific impacts on the biogeochemistry and lower-trophic level organisms of the northeast Pacific and CCS.

Carbon dioxide

Numerical models are widely employed to diagnose climatic forcing of the physical and biogeochemical conditions of the northeast Pacific. For instance, using a fully coupled ocean and biogeochemical model, Xiu and Chai (2014) found that after accounting for atmospheric effects, the air-sea flux and resulting pCO₂ of sea water in the Pacific was significantly correlated (0.6) to the Multivariate ENSO Index (MEI) with a lag of ten months. Similarly, Wong et al. (2010) found that sea surface pCO₂ was significantly correlated with the MEI in the northeast Pacific. Biogeochemical models of different complexity have also highlighted the connection between PDO and the interannual variability of air-sea CO₂ fluxes (e.g., McKinley et al. 2006). These studies also demonstrate that the individual components controlling surface ocean pCO₂ in the northeast Pacific respond to PDO with significant amplitudes, but that their combined influence has a relatively small effect on the CO₂ fluxes in this region. Xiu and Chai (2014) showed that the dominant driver of North Pacific pCO₂ variability is anthropogenic CO_2, whereas air-sea CO₂ flux is more closely correlated with the PDO and the NPGO.

Nutrients and chlorophyll

In the coastal regions of the northeast Pacific, such as the CCS, ENSO significantly impacts the nutrient supply due to modifications of upwelling and source waters mentioned above (Jacox et al., this post). At the peak of the El Niño season in December-January, Frischknecht et al. (2016) found a pattern in the development of chlorophyll events through a modeling study focused on the CCS. Around the onset of the El Niño year, chlorophyll anomalies were consistently low. This pattern was even more pronounced during the spring of the following year. In spring of the second year (i.e. with the onset of the upwelling season), all events shared the development of a strong negative chlorophyll anomaly. Frischknecht et al. (2016) attributed this phenomenon to a persistent lack of nutrients to support production driven by a combination of physical mechanisms impacting the thermocline (Jacox et al. 2015; 2016; this post) and light limitation at the onset of the upwelling season. Consequently, El Niño events disrupt the biogeochemical cycling in these systems for months, even years, after the event is over. The observations in Oregon, from the Newport line in Fisher et al. (2015), detail the nitrate anomalies from 1995 to 2015, and the nitrate anomalies remain negative long after the Niño-3.4 SST anomaly suggested that the event was over. This may contribute to the success surrounding seasonal forecast systems like J-SCOPE, in which forecasts of biogeochemical parameters (e.g., bottom oxygen) outperformed those of physical variables (e.g., SST) in terms of predictive skill (Siedlecki et al. 2016).

Oxygen and carbon

The relationship between ENSO and nutrient availability from source waters can serve as an analog for oxygen and carbon content. We would expect from observed stoichiometry that when nutrients are low, oxygen is relatively high and carbon is low. In California, this has been documented: El Niño events correlate to higher oxygen and pH, while La Niña events are correlated with lower oxygen and pH (e.g., Nam et al. 2011). In the northern CCS along the Washington and Oregon coasts, the interannual variability in oxygen content of source waters has been correlated to NPGO more than ENSO (Peterson et al. 2013). Consistent with these findings, oxygen has been increasing since 2010 and aragonite saturation state (a measure of the availability of carbonate ion to calcifying organisms) has been elevated in 2015-2016 relative to the year prior in both Oregon and California (McClatchie et al. 2016).

Primary production & particle export

As an eastern boundary upwelling region, the CCS is among the most productive in the world in terms of primary production and fisheries. The suppression of nutrient availability described above can be thought of as reduced “upwelling efficacy” that leads to reduced primary production in the CCS, while La Niña often has the opposite effect due to associated increases in the upwelling efficacy (Jacox et al. 2015). In the southern CCS, the 1997/1998 El Niño led to a significant deepening of the nutricline, with the strongest effects along CalCOFI Line 80, and a pronounced regional reduction of primary production (Bograd and Lynn 2001). The uptake of silicon increased in central California (Santa Barbara Channel) during the onset of the 1997 event, suggesting that diatoms were major drivers of the primary productivity prior to the 1998 spring season when overall productivity was reduced in response to density surface adjustments (Shipe and Brzezinski 2003). Despite reductions in surface layer primary productivity in response to the El Niño, export ratios of particulate organic carbon and particulate organic nitrogen increased during the spring of 1998 relative to the 1994-1997 period, while biogenic Si flux decreased in response to the El Niño (Shipe et al. 2002). This counterintuitive result appears to be due to an increase in particulate material exported to depth. By 1999, ratios of Si/N and Si/C had not recovered to pre-El Nino conditions.

Phytoplankton community composition

Warmer waters and changes in nutrient supply associated with ENSO can lead to phytoplankton community shifts such as an influx of coccolithophores or an increase in harmful algal blooms (HABs). The most common harmful algal bloom organism in the CCS is the diatom genus Pseudo-nitzschia. McCabe et al. (2016) recently observed a link between the Oceanic Niño Index (ONI), the Pseudo-nitzschia growth rate anomaly determined from temperature-growth relationships, and domoic acid levels in razor clams over a 16-year period (Figure 1), implicating El Niño-driven warming in the unprecedented 2015 HAB along the US West Coast. Similarly, McKibben et al. (2017) linked warm phases of the PDO and ONI to domoic acid in shellfish in the northern CCS. The toxic blooms off Newport in 2015 were the most prolonged (late-April through October 2015) and among the most toxic ever observed off Oregon (Du et al. 2015; McKibben et al. 2017). Conversely, Santa Barbara Basin sediment trap data showed no significant correlation between a 15-year record of domoic acid levels and PDO, NPGO, or ENSO indices; however, there was a strong change point in the frequency and toxicity of these blooms following the 1997/1998 ENSO (Sekula-Wood et al. 2011).

Figure 1. Records of razor clam toxicity and the potential growth rate anomaly for Pseudo-nitzschia spp. are plotted below the Oceanic Niño Index (gold = El Niño, blue = La Niña, gray = neutral) to illustrate the association between ENSO events and harmful algal blooms in the northern CCS. The potential for Pseudo–nitzschia growth does not always coincide with records of high domoic acid in shellfish, e.g., 1997 El Niño. Figure adapted from McCabe et al. (2016).

Zooplankton community composition

Off the Oregon coast, a 21-year time-series of fortnightly hydrography and plankton sampling of shelf and slope waters showed that the water masses (and thus the plankton) that dominate shelf and slope waters vary seasonally, interannually, and on decadal scales. Thus, it is a simple matter to track the timing of summer or winter arrival, ENSO events, and changes in sign of the PDO (Figure 2). During summer months, northerly winds drive surface waters offshore (Ekman transport), which are replaced by the upwelling of cold nutrient-rich waters that penetrate the continental shelf and fuel high primary production. Northerly winds also enhance the southward transport of water (and plankton) from the coastal Gulf of Alaska into the coastal northern CCS, and these species are referred to as ‘cold water’ or ‘northern species.’ During winter, the winds reverse and the poleward Davidson current transports warm coastal water from southern California to the northern CCS, bringing with it ‘southern species’ of plankton. On longer timescales (5-10 years), cold-water, northern copepods are largely replaced by warm-water, southern copepods during El Niño events (Fisher et al. 2015) and during the positive phase of the PDO (Keister et al. 2011). Incorporating the physiological response of these zooplankton groups into biogeochemical-ecosystem models (in addition to the effects of physical transport) will be essential for advancing our predictive capacity of plankton communities in the CCS.

Figure 2. Monthly time series of the Pacific Decadal Oscillation and Oceanic Niño Index (upper) and monthly-averaged biomass anomalies of northern copepods (middle) and southern copepods (lower). Note the high coherence between the PDO and ONI with the copepod time series – positive anomalies of northern copepods are correlated with negative PDO and ONI; positive anomalies of southern copepods are correlated with positive PDO and ONI.

Predicting ecosystem response to ENSO: Now and in the future

State-of-the-art models, in situ measurements, and available satellite observations are all required to adequately characterize short- and long-term physical dynamics associated with ENSO and Pacific decadal variability. Seasonal-to-interannual forecasting of the ecosystem response to ENSO in the CCS and throughout the northeast Pacific will depend on our understanding of how interannual climate variability alters ocean biogeochemistry and productivity at the base of the food web, and therefore how predictive models should be modified to capture the dynamic range introduced by these anomalous events. Unusual warm water anomalies, as observed during large ENSO events, may serve as important analogs for assessing the impacts of long-term warming on the pelagic ecosystem of the CCS. Regional simulations suggest similarities between the physical drivers leading to biogeochemical variability from ENSO and those in projected future upwelling systems (Rykaczewski and Dunne 2010). Further exploration of the mechanisms and predictive skill of forecasts on seasonal timescales will enhance our understanding and improve our projections further into the future. Global climate models are unable to anticipate anomalous warming events such as major ENSO events. As such, they are unable to detect large-scale events related to shifts in the distribution of pelagic species or track ecological changes associated with such events. Furthermore, the evaluation of model-based forecasts and projections of ecosystem variations and changes across timescales requires that long-term physical, biogeochemical, and ecological observation programs are maintained and others initiated. High-resolution modeling approaches for forecasts and projections should also be prioritized, so that ecosystem impacts of future climate anomalies can be anticipated and understood in greater detail.

Authors

Clarissa Anderson (Scripps Institution of Oceanography)
Samantha Siedlecki (University of Washington)
Cecile Rousseaux (NASA Goddard Space Flight Center, Universities Space Research Association)
Brian Powell (University of Hawaii, Manoa)
Bill Peterson (NOAA Northwest Fisheries Science Center)
Chris Edwards (University of California, Santa Cruz)

References

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Funding for the Ocean Carbon & Biogeochemistry Project Office is provided by the National Science Foundation (NSF) and the National Aeronautics and Space Administration (NASA). The OCB Project Office is housed at the Woods Hole Oceanographic Institution.

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