Anchovy boom and bust linked to trophic shifts in larval diet.

Although massive biomass fluctuations of coastal-pelagic fishes are an iconic example of the impacts of climate variability on marine ecosystems, the mechanisms governing these dynamics are often elusive. We construct a 45-year record of nitrogen stable isotopes measured in larvae of Northern Anchovy (Engraulis mordax) in the California Current Ecosystem to assess patterns in food chain length. Larval trophic efficiency associated with a shortened food chain increased larval survival and produced boom periods of high adult biomass. In contrast, when larval food chain length increased, and energy transfer efficiency decreased, the population crashed. We propose the Trophic Efficiency in Early Life (TEEL) hypothesis, which states that larval fishes must consume prey that confer sufficient energy for survival, to help explain natural boom-bust dynamics of coastal pelagic fishes. Our findings illustrate a potential for trophic indicators to generally inform larval survival and adult population dynamics of coastal-pelagic fishes.

Carbon sequestration by multiple biological pump pathways in a coastal upwelling biome.

Multiple processes transport carbon into the deep ocean as part of the biological carbon pump, leading to long-term carbon sequestration. However, our ability to predict future changes in these processes is hampered by the absence of studies that have simultaneously quantified all carbon pump pathways. Here, we quantify carbon export and sequestration in the California Current Ecosystem resulting from (1) sinking particles, (2) active transport by diel vertical migration, and (3) the physical pump (subduction + vertical mixing of particles). We find that sinking particles are the most important and export 9.0 mmol C m-2 d-1 across 100-m depth while sequestering 3.9 Pg C. The physical pump exports more carbon from the shallow ocean than active transport (3.8 vs. 2.9 mmol C m-2 d-1), although active transport sequesters more carbon (1.0 vs. 0.8 Pg C) because of deeper remineralization depths. We discuss the implications of these results for understanding biological carbon pump responses to climate change.

Machine learning techniques to characterize functional traits of plankton from image data.

Plankton imaging systems supported by automated classification and analysis have improved ecologists' ability to observe aquatic ecosystems. Today, we are on the cusp of reliably tracking plankton populations with a suite of lab-based and in situ tools, collecting imaging data at unprecedentedly fine spatial and temporal scales. But these data have potential well beyond examining the abundances of different taxa; the individual images themselves contain a wealth of information on functional traits. Here, we outline traits that could be measured from image data, suggest machine learning and computer vision approaches to extract functional trait information from the images, and discuss promising avenues for novel studies. The approaches we discuss are data agnostic and are broadly applicable to imagery of other aquatic or terrestrial organisms.

On the sensitivity of plankton ecosystem models to the formulation of zooplankton grazing.

Model representations of plankton structure and dynamics have consequences for a broad spectrum of ocean processes. Here we focus on the representation of zooplankton and their grazing dynamics in such models. It remains unclear whether phytoplankton community composition, growth rates, and spatial patterns in plankton ecosystem models are especially sensitive to the specific means of representing zooplankton grazing. We conduct a series of numerical experiments that explicitly address this question. We focus our study on the form of the functional response to changes in prey density, including the formulation of a grazing refuge. We use a contemporary biogeochemical model based on continuum size-structured organization, including phytoplankton diversity, coupled to a physical model of the California Current System. This region is of particular interest because it exhibits strong spatial gradients. We find that small changes in grazing refuge formulation across a range of plausible functional forms drive fundamental differences in spatial patterns of plankton concentrations, species richness, pathways of grazing fluxes, and underlying seasonal cycles. An explicit grazing refuge, with refuge prey concentration dependent on grazers' body size, using allometric scaling, is likely to provide more coherent plankton ecosystem dynamics compared to classic formulations or size-independent threshold refugia. We recommend that future plankton ecosystem models pay particular attention to the grazing formulation and implement a threshold refuge incorporating size-dependence, and we call for a new suite of experimental grazing studies.

Timing is everything: Drivers of interannual variability in blue whale migration.

Blue whales need to time their migration from their breeding grounds to their feeding grounds to avoid missing peak prey abundances, but the cues they use for this are unknown. We examine migration timing (inferred from the local onset and cessation of blue whale calls recorded on seafloor-mounted hydrophones), environmental conditions (e.g., sea surface temperature anomalies and chlorophyll a), and prey (spring krill biomass from annual net tow surveys) during a 10 year period (2008-2017) in waters of the Southern California Region where blue whales feed in the summer. Colder sea surface temperature anomalies the previous season were correlated with greater krill biomass the following year, and earlier arrival by blue whales. Our results demonstrate a plastic response of blue whales to interannual variability and the importance of krill as a driving force behind migration timing. A decadal-scale increase in temperature due to climate change has led to blue whales extending their overall time in Southern California. By the end of our 10-year study, whales were arriving at the feeding grounds more than one month earlier, while their departure date did not change. Conservation strategies will need to account for increased anthropogenic threats resulting from longer times at the feeding grounds.

Multidecadal increase in plastic particles in coastal ocean sediments.

We analyzed coastal sediments of the Santa Barbara Basin, California, for historical changes in microplastic deposition using a box core that spanned 1834-2009. The sediment was visually sorted for plastic, and a subset was confirmed as plastic polymers via FTIR (Fourier transform infrared) spectroscopy. After correcting for contamination introduced during sample processing, we found an exponential increase in plastic deposition from 1945 to 2009 with a doubling time of 15 years. This increase correlated closely with worldwide plastic production and southern California coastal population increases over the same period. Increased plastic loading in sediments has unknown consequences for deposit-feeding benthic organisms. This increase in plastic deposition in the post-World War II years can be used as a geological proxy for the Great Acceleration of the Anthropocene in the sedimentary record.

Effects of chemical preservation on bulk and amino acid isotope ratios of zooplankton, fish, and squid tissues.

RATIONALE: It is imperative to understand how chemical preservation alters tissue isotopic compositions before using historical samples in ecological studies. Specifically, although compound-specific isotope analysis of amino acids (CSIA-AA) is becoming a widely used tool, there is little information on how preservation techniques affect amino acid δ15 N values. METHODS: We evaluated the effects of chemical preservatives on bulk tissue δ13 C and δ15 N and amino acid δ15 N values, measured by gas chromatography/isotope ratio mass spectrometry (GC/IRMS), of (a) tuna (Thunnus albacares) and squid (Dosidicus gigas) muscle tissues that were fixed in formaldehyde and stored in ethanol for 2 years and (b) two copepod species, Calanus pacificus and Eucalanus californicus, which were preserved in formaldehyde for 24-25 years. RESULTS: Tissues in formaldehyde-ethanol had higher bulk δ15 N values (+1.4, D. gigas; +1.6‰, T. albacares), higher δ13 C values for D. gigas (+0.5‰), and lower δ13 C values for T. albacares (-0.8‰) than frozen samples. The bulk δ15 N values from copepods were not different those from frozen samples, although the δ13 C values from both species were lower (-1.0‰ for E. californicus and -2.2‰ for C. pacificus) than those from frozen samples. The mean amino acid δ15 N values from chemically preserved tissues were largely within 1‰ of those of frozen tissues, but the phenylalanine δ15 N values were altered to a larger extent (range: 0.5-4.5‰). CONCLUSIONS: The effects of preservation on bulk δ13 C values were variable, where the direction and magnitude of change varied among taxa. The changes in bulk δ15 N values associated with chemical preservation were mostly minimal, suggesting that storage in formaldehyde or ethanol will not affect the interpretation of δ15 N values used in ecological studies. The preservation effects on amino acid δ15 N values were also mostly minimal, mirroring bulk δ15 N trends, which is promising for future CSIA-AA studies of archived specimens. However, there were substantial differences in phenylalanine and valine δ15 N values, which we speculate resulted from interference in the chromatographic resolution of unknown compounds rather than alteration of tissue isotopic composition due to chemical preservation.

Climate-mediated changes in marine ecosystem regulation during El Niño.

The degree to which ecosystems are regulated through bottom-up, top-down, or direct physical processes represents a long-standing issue in ecology, with important consequences for resource management and conservation. In marine ecosystems, the role of bottom-up and top-down forcing has been shown to vary over spatio-temporal scales, often linked to highly variable and heterogeneously distributed environmental conditions. Ecosystem dynamics in the Northeast Pacific have been suggested to be predominately bottom-up regulated. However, it remains unknown to what extent top-down regulation occurs, or whether the relative importance of bottom-up and top-down forcing may shift in response to climate change. In this study, we investigate the effects and relative importance of bottom-up, top-down, and physical forcing during changing climate conditions on ecosystem regulation in the Southern California Current System (SCCS) using a generalized food web model. This statistical approach is based on nonlinear threshold models and a long-term data set (~60 years) covering multiple trophic levels from phytoplankton to predatory fish. We found bottom-up control to be the primary mode of ecosystem regulation. However, our results also demonstrate an alternative mode of regulation represented by interacting bottom-up and top-down forcing, analogous to wasp-waist dynamics, but occurring across multiple trophic levels and only during periods of reduced bottom-up forcing (i.e., weak upwelling, low nutrient concentrations, and primary production). The shifts in ecosystem regulation are caused by changes in ocean-atmosphere forcing and triggered by highly variable climate conditions associated with El Niño. Furthermore, we show that biota respond differently to major El Niño events during positive or negative phases of the Pacific Decadal Oscillation (PDO), as well as highlight potential concerns for marine and fisheries management by demonstrating increased sensitivity of pelagic fish to exploitation during El Niño.

Mesoscale ocean fronts enhance carbon export due to gravitational sinking and subduction.

Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg C⋅m-2⋅d-1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional ∼225 mg C⋅m-2⋅d-1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.

Long-term aging and degradation of microplastic particles: Comparing in situ oceanic and experimental weathering patterns.

Polypropylene, low-density polyethylene, and high-density polyethylene pre-production plastic pellets were weathered for three years in three experimental treatments: dry/sunlight, seawater/sunlight, and seawater/darkness. Changes in chemical bond structures (hydroxyl, carbonyl groups and carbon-oxygen) with weathering were measured via Fourier Transform Infrared (FTIR) spectroscopy. These indices from experimentally weathered particles were compared to microplastic particles collected from oceanic surface waters in the California Current, the North Pacific Subtropical Gyre, and the transition region between the two, in order to estimate the exposure time of the oceanic plastics. Although chemical bonds exhibited some nonlinear changes with environmental exposure, they can potentially approximate the weathering time of some plastics, especially high-density polyethylene. The majority of the North Pacific Subtropical Gyre polyethylene particles we measured have inferred exposure times>18months, with some >30months. Inferred particle weathering times are consistent with ocean circulation models suggesting a long residence time in the open ocean.

Resilience and stability of a pelagic marine ecosystem.

The accelerating loss of biodiversity and ecosystem services worldwide has accentuated a long-standing debate on the role of diversity in stabilizing ecological communities and has given rise to a field of research on biodiversity and ecosystem functioning (BEF). Although broad consensus has been reached regarding the positive BEF relationship, a number of important challenges remain unanswered. These primarily concern the underlying mechanisms by which diversity increases resilience and community stability, particularly the relative importance of statistical averaging and functional complementarity. Our understanding of these mechanisms relies heavily on theoretical and experimental studies, yet the degree to which theory adequately explains the dynamics and stability of natural ecosystems is largely unknown, especially in marine ecosystems. Using modelling and a unique 60-year dataset covering multiple trophic levels, we show that the pronounced multi-decadal variability of the Southern California Current System (SCCS) does not represent fundamental changes in ecosystem functioning, but a linear response to key environmental drivers channelled through bottom-up and physical control. Furthermore, we show strong temporal asynchrony between key species or functional groups within multiple trophic levels caused by opposite responses to these drivers. We argue that functional complementarity is the primary mechanism reducing community variability and promoting resilience and stability in the SCCS.

A double-integration hypothesis to explain ocean ecosystem response to climate forcing.

Long-term time series of marine ecological indicators often are characterized by large-amplitude state transitions that can persist for decades. Understanding the significance of these variations depends critically on the underlying hypotheses characterizing expected natural variability. Using a linear autoregressive model in combination with long-term zooplankton observations off the California coast, we show that cumulative integrations of white-noise atmospheric forcing can generate marine population responses that are characterized by strong transitions and prolonged apparent state changes. This model provides a baseline hypothesis for explaining ecosystem variability and for interpreting the significance of abrupt responses and climate change signatures in marine ecosystems.

Biological responses to environmental forcing: the linear tracking window hypothesis.

Determining the relative contributions of intrinsic and extrinsic processes to the regulation of biological populations has been a recurrent ecological issue. Recent discussions concerning ecosystem "regime shifts" again raise the question of whether population fluctuations are mainly controlled by external forcing. Results of nonlinear time series analyses indicate that pelagic populations typically do not passively track stochastic environmental variables. Rather, population dynamics are better described as nonlinear amplification of physical forcing by biological interactions. However, we illustrate that in some cases populations do show linear tracking of the physical environment. To explain why population dynamics can sometimes be linear, we propose the linear tracking window hypothesis: populations are most likely to track the stochastic environmental forcing when their generation time matches the characteristic time scale of the environmental signal. While our observations follow this hypothesis well, our results indicate that the linear tracking window is a necessary but not a sufficient condition.

Planktonic foraminifera of the California Current reflect 20th-century warming.

It is currently unclear whether observed pelagic ecosystem responses to ocean warming, such as a mid-1970s change in the eastern North Pacific, depart from typical ocean variability. We report variations in planktonic foraminifera from varved sediments off southern California spanning the past 1400 years. Increasing abundances of tropical/subtropical species throughout the 20th century reflect a warming trend superimposed on decadal-scale fluctuations. Decreasing abundances of temperate/subpolar species in the late 20th century indicate a deep, penetrative warming not observed in previous centuries. These results imply that 20th-century warming, apparently anthropogenic, has already affected lower trophic levels of the California Current.

Egg-brooding, body size and predation risk in planktonic marine copepods.

We investigated the interacting effects of copepod body size and the presence or absence of egg masses on the risk of predation by a visual predator. We conducted selection experiments involving three-spined sticklebacks (Gasterosteus aculeatus) and copepods ranging in body mass from 0.5 to 740 µg C: Oithona similis, Corycaeus anglicus, Pseudocalanus newmani, P. moultoni, Pseudodiaptomus marinus, and Paraeuchaeta elongata. We found that sticklebacks selected ovigerous females of the two smallest-bodied species of copepods (Oithona similis and Corycaeus anglicus). In contrast, the fish showed no significant selection for ovigerous females of the remaining, larger-bodied species. Unexpectedly, egg mass position (i.e., in a ventral, dorsal or lateral location on the urosome) appeared to influence predation risk more than did body size, resulting in higher predation risk for the cyclopoid and poecilostomatoid species than for the calanoid species we tested. Although the sticklebacks showed no statistically significant preference for ovigerous females of any of the four calanoid species, for each species the overall proportion of ovigerous females ingested was slightly greater than 0.50. Thus, whether body size influences the susceptibility of egg-brooding calanoid copepods to predation remains an open question.