Southern California Current Ecosystem Research Articles

Guadalupe fur seal pup production predicted from annual variations of sea surface temperature in the southern California Current Ecosystem

AbstractWe evaluated the effects of sea surface temperature anomalies (SSTA) in the southern California Current Ecosystem on the annual Guadalupe fur seal (Arctocephalus townsendi) pup production, a species recovering from near extinction. Pup counts from 1991 to 1993 and from 2006 to 2019 were used to estimate deviations from a long-term trend as a proxy for the population's reproductive success. We estimated interannual SSTA as a subtraction from the linear trend spanning 1991–2019 for a 778,000 km2 area, which represents the primary foraging range of adult females. The long-term increase in pup production followed an exponential curve ( ${\rm{R}}_{\rm{B}}^2 = {\rm{\ }}1$), typical of species in a recovery phase. Pup production deviations from this trend responded to SSTA during the gestation period as a cubic polynomial function ( ${\rm{R}}_{\rm{B}}^2 = {\rm{\ }}0.837$), revealing that SSTA < −0.2°C and between ∼0.6 and 1.38°C increased pup production in the subsequent breeding season, whereas normal to slightly warm (−0.17 to 0.6°C) and extreme SSTA (>1.4°C) decreased pup counts, arguably resulting from low prey availability and quality. This model allowed us to estimate pup production for years without observations, needed to understand the environmental variability influence on the recovery process of this species, and therefore constitutes a practical tool for its conservation and management.

Microbial communities associated with sinking particles across an environmental gradient from coastal upwelling to the oligotrophic ocean

Complex interactions between microbial communities in the epipelagic and mesopelagic ocean drive variability in sinking carbon flux as part of the biological carbon pump. To investigate the relative contribution of dominant prokaryotic, protistan, and phytoplankton taxa to sinking particles, we used 16S/18S rRNA gene sequencing of particles collected in sediment traps and water column communities. Comparisons were done across a coastal upwelling-to-oligotrophic environmental gradient in the southern California Current Ecosystem to assess effects of natural environmental variability on taxon-specific contributions to sinking particle export. Particle-associated microbial assemblages differ from ambient mixed-layer communities. Gammaproteobacteria, dinoflagellates, and rhizarians were dominant microbes associated with sinking particles at all sampling locations, with diatoms increasing significantly at the inshore mesotrophic site. Parasitic groups, syndiniales and apicomplexans, were also major particle-associated taxa. Relative sequence abundance on exported particles from the mesotrophic inshore site more closely resembled the water-column dominants (especially diatoms) found above, while oligotrophic communities exhibited greater dissimilarity between the microbial communities on sinking particles and in the mixed layer. Protists generally showed greater similarity between mixed-layer and sinking particle communities than prokaryotes. Synechococcus was over-represented on sinking particles (relative to eukaryotic phytoplankton and other cyanobacteria) only in oligotrophic areas. Diatoms were the only phytoplankton group consistently over-represented on sinking particles across the region. Our results highlight important variability in microbial contributions to export that seem to align with differences in dominant grazing pathways. At the upwelling influenced site where export was primarily due to the fecal pellets of large copepods, diatoms were dominant contributors to export and protistan communities on sinking particles resembled those in the water column above. When doliolids were abundant, dinoflagellates contributed substantially to export, while the dominant picoeukaryote (Ostreococcus) was only a minor contributor. In oligotrophic areas, where grazing pathways were dominated by protists and sinking particles were primarily amorphous aged aggregates, there was little similarity between mixed layer and sinking particle communities.

Zooplankton biogeographic boundaries in the California Current System as determined from metabarcoding.

Within the southern California Current ecosystem there are two well-documented breaks in marine community structure at Point Conception and Punta Eugenia. We explored the presence of similar breaks in a diverse zooplankton community through metabarcoding of mixed net tow tissue samples collected during an expedition from Monterey to Baja California in February of 2012. We recovered a high diversity of species as well as patterns of species presence that align with their previously documented ranges in this region. We found a clear break at Punta Eugenia in overall zooplankton community structure, while Point Conception was weakly linked to changes in community structure. We analyzed this dataset through two parallel bioinformatic pipelines to examine the robustness of these results. Our overall conclusions were consistent across both pipelines, however there were differences in species detection. This study illustrates the utility of metabarcoding analysis on mixed tissue samples for recovering known patterns of diversity, as well as allowing elucidation of broad patterns of community differentiation across many groups of organisms.

Vertical niche definition of test‐bearing protists (Rhizaria) into the twilight zone revealed by in situ imaging

AbstractThe Rhizaria is a super‐group of amoeboid protists with ubiquitous distributions, from the euphotic zone to the twilight zone and beyond. While rhizarians have been recently described as important contributors to both biogenic silica and carbon fluxes, we lack the most basic information about their ecological habitats and preferences. Here, using in situ imaging (Underwater Vision Profiler 5), we characterize the vertical ecological niches of different test‐bearing pelagic rhizarian taxa in the southernCalifornia Current Ecosystem. We define three vertical layers between 0 and 500 m occupied, respectively, by (1) surface dwelling and mostly symbiont‐bearing rhizarians (Acantharia and Collodaria), (2) flux‐feeding phaeodarians in the lower epipelagic (100–200 m), and (3) Foraminifera and Phaeodaria populations adjacent to the oxygen minimum zone. We then use Generalized Additive Models to analyze the response of each rhizarian category to a suite of environmental variables. The models explain between 9% and 93% of the total variance observed for the different groups. While temperature and the depth of the deep chlorophyll maximum appear as the main abiotic factors influencing populations in the upper 200 m, dissolved silicon concentration is related to the abundance of mesopelagic phaeodarians, though it explains only a portion of the variance. The importance of biotic interactions (e.g., prey availability, predation, parasitism, symbiosis) is still to be considered, in order to fully incorporate the dynamics of test‐bearing pelagic rhizarians in ecological and biogeochemical models.

Influence of wind events on larval fish mortality rates in the southern California Current Ecosystem

Wind-induced mixing can affect the vertical distribution of plankton in the upper water column, influencing the prey available for larval fishes. The stable ocean hypothesis proposes that periods of calm winds facilitate the development of plankton layers at concentrations sufficient for successful larval foraging and increased survival. Conversely, storm events redistribute prey, leading to reduced foraging success. Here, we investigate this hypothesis by comparing larval fish mortality rates estimated from 37 years of ichthyoplankton data against metrics of wind events defined as storms and calm periods. Contrary to expectations, we found that mortality for Pacific hake (Merluccius productus) significantly decreased as storm events increased in the southern California Current Ecosystem. Mortality rates for northern anchovy (Engraulis mordax), Pacific sardine (Sardinops sagax), Pacific mackerel (Scomber japonicus), and jack mackerel (Trachurus symmetricus) had no relationship to storms, and no species’ mortality rates were related to the number of calm events. Our results highlight the differing sensitivities of larval survival among fishes in the region and indicate that responses to atmospheric processes are species-dependent.

Pervasive iron limitation at subsurface chlorophyll maxima of the California Current

Subsurface chlorophyll maximum layers (SCMLs) are nearly ubiquitous in stratified water columns and exist at horizontal scales ranging from the submesoscale to the extent of oligotrophic gyres. These layers of heightened chlorophyll and/or phytoplankton concentrations are generally thought to be a consequence of a balance between light energy from above and a limiting nutrient flux from below, typically nitrate (NO3). Here we present multiple lines of evidence demonstrating that iron (Fe) limits or with light colimits phytoplankton communities in SCMLs along a primary productivity gradient from coastal to oligotrophic offshore waters in the southern California Current ecosystem. SCML phytoplankton responded markedly to added Fe or Fe/light in experimental incubations and transcripts of diatom and picoeukaryote Fe stress genes were strikingly abundant in SCML metatranscriptomes. Using a biogeochemical proxy with data from a 40-y time series, we find that diatoms growing in California Current SCMLs are persistently Fe deficient during the spring and summer growing season. We also find that the spatial extent of Fe deficiency within California Current SCMLs has significantly increased over the last 25 y in line with a regional climate index. Finally, we show that diatom Fe deficiency may be common in the subsurface of major upwelling zones worldwide. Our results have important implications for our understanding of the biogeochemical consequences of marine SCML formation and maintenance.

Introduction to collection of papers on the response of the southern California Current Ecosystem to the Warm Anomaly and El Niño, 2014–16

This contribution provides an introduction to a sequence of five papers (CCE I- CCE V) that describe the impact of the Warm Anomaly of 2014–15 and El Niño 2015–16 on the pelagic food web of the southern California Current Ecosystem. These contributions analyze the influence of these two warm water perturbations on satellite-based measures of ocean fronts, export efficiency out of the euphotic zone, copepod egg production, mesozooplankton community structure, and a synthesis of primary production, mesozooplankton grazing, and gravitational fluxes of organic carbon.

Mesopelagic fish assemblages across oceanic fronts: A comparison of three frontal systems in the southern California Current Ecosystem

With strong horizontal gradients in physical properties, oceanic frontal regions can lead to disproportionately high biological productivity. We examined cross-frontal changes in mesopelagic fish assemblages at three separate frontal systems in the southern California Current Ecosystem (CCE) as part of the CCE Long Term Ecological Research program: the A-Front sampled in October 2008, the C-Front in June/July 2011, and the E-Front in July/August 2012. We analyzed the differential effects of front-associated regions on density and species composition of adult migratory and non-migratory fishes and larvae, and the larval to adult ratio (as a possible index of a population growth potential) for migratory and non-migratory species. The fronts did not have a strong effect on densities of any subset of the mesopelagic fish assemblage. The species composition of the vertical migratory fishes (and their larvae) was typically altered across fronts, with different assemblages present on either side of each front. The migratory assemblages at the fronts themselves were indistinguishable from those at the more productive side of the frontal system. In contrast, the assemblage composition of the non-migratory fishes was indistinguishable between regions across all three of the fronts. The differences between the Northern and Southern assemblages at the A-Front were primarily based on biogeographic provinces, while the assemblages at the E-Front were largely distinguishable by their oceanic or coastal-upwelling zone associations. These results generally confirm those of previous studies on frontal systems in the California Current Ecosystem and elsewhere. The ratio of larvae to adults, a potential index of population growth potential, was altered across two of the fronts for migratory species, elevated on the colder side of the A-Front and the warmer side of the E-Front. This finding suggests that fronts may be regions of enhanced reproduction. The larvae to adult ratio was indistinguishable for non-migratory species at all three frontal systems. The non-migratory component of the community was little influenced by the presence of a front, apparently because the regions of strongest horizontal spatial gradients were too shallow to be experienced directly. We speculate that there was no change in larval community composition and population growth index at the most dynamic frontal system (C-Front) compared to the other fronts surveyed because the frontal feature was short-lived relative to the time scale for population growth of the fish. However, the difference in results of the C-Front may also be due to a change in methodology used in this study. If mesoscale features such as fronts increase in frequency off the California coast in the future as predicted, they have the potential to alter population growth potential and restructure mesopelagic fish assemblages, which are dominated by migratory species.

Persistence of trophic hotspots and relation to human impacts within an upwelling marine ecosystem.

Human impacts (e.g., fishing, pollution, and shipping) on pelagic ecosystems are increasing, causing concerns about stresses on marine food webs. Maintaining predator-prey relationships through protection of pelagic hotspots is crucial for conservation and management of living marine resources. Biotic components of pelagic, plankton-based, ecosystems exhibit high variability in abundance in time and space (i.e., extreme patchiness), requiring investigation of persistence of abundance across trophic levels to resolve trophic hotspots. Using a 26-yr record of indicators for primary production, secondary (zooplankton and larval fish), and tertiary (seabirds) consumers, we show distributions of trophic hotspots in the southern California Current Ecosystem result from interactions between a strong upwelling center and a productive retention zone with enhanced nutrients, which concentrate prey and predators across multiple trophic levels. Trophic hotspots also overlap with human impacts, including fisheries extraction of coastal pelagic and groundfish species, as well as intense commercial shipping traffic. Spatial overlap of trophic hotspots with fisheries and shipping increases vulnerability of the ecosystem to localized depletion of forage fish, ship strikes on marine mammals, and pollution. This study represents a critical step toward resolving pelagic areas of high conservation interest for planktonic ecosystems and may serve as a model for other ocean regions where ecosystem-based management and marine spatial planning of pelagic ecosystems is warranted.

Measurement of dark, particle-generated superoxide and hydrogen peroxide production and decay in the subtropical and temperate North Pacific Ocean

Reactive oxygen species (ROS), which include the superoxide radical (O2-) and hydrogen peroxide (H2O2), are thought to be generated mostly through photochemical reactions and biological activity in seawater and can influence trace metal speciation in the ocean. This study reports the results of an intercomparison of two methods to measure particle-generated [O2-] in seawater samples, as well as measurements of particle-generated O2- and H2O2 concentrations, decay kinetics, and dark production rates in seawater samples at Station ALOHA and (O2- only) in the southern California Current Ecosystem. O2- was measured using two different methods relying on chemiluminescence detection. The first method measured the difference between steady-state [O2-] in filtered and unfiltered seawater, while the second method (standard method) measured O2- decay to baseline in freshly filtered seawater. Because both methods detected [O2-] relative to the background signal from filtered seawater, both should have measured [O2-] generated by particles (presumably biota). However, the O2- concentrations determined by the first method were always much smaller than those obtained from the second (standard) method. Follow-up laboratory and field experiments showed that the increased signal in the standard method was due to a filtration artifact that could neither be eliminated nor consistently accounted for under the tested conditions. We therefore recommend the first method for measuring particle-generated [O2-]. Measured by this method, Station ALOHA had particle-generated O2- concentrations that ranged from undetectable to 0.02nM, with production rates less than 0.6nMhr−1 and decay rate coefficients from 0.003 to 0.014s−1. The southern California Current Ecosystem had particle-generated O2- concentrations that ranged from undetectable to 0.05nM, with production rates up to 4.7nMhr−1 and decay rate coefficients from 0.006 to 0.017s−1. H2O2 concentrations were measured by chemiluminescence detection, using dark incubations of unfiltered water samples to simultaneously determine production and decay rates. H2O2 concentrations at Station ALOHA ranged from 7 to 88nM. Dark production rates and decay rate coefficients were low (mostly <1.5nMhr−1 and <0.03h−1, respectively); higher values were detected when biota were pre-concentrated with net tows. These rates of ROS production are lower than those reported by previous studies in other regions of the Pacific Ocean, but could still be significant compared to photochemical production.

Persistence of hotspots and variability of seabird species richness and abundance in the southern California Current

Aggregations of seabirds at sea may provide information on centers of enhanced trophic interactions and concentrating mechanisms, however, to date most studies lack quantification of persistence, a key hotspot characteristic. Persistence statistics may reduce uncertainty about seabird habitat use, improve understanding of the spatio‐temporal scales of pelagic food web dynamics, and inform conservation planning. Using 26 years (1987–2012, 47 surveys) of shipboard surveys from a 300K km2 study area within the southern California Current Ecosystem, we conduct a spatial assessment of the inter‐annual and seasonal dynamics of the persistence of seabird hotspots and identify recurring sites of elevated seabird species richness and abundance. Previous studies document declines in abundance, but were based on broad spatial standardizations to assess where declines may have occurred. Here, we refine the hypothesis that seabird populations have declined off southern California by focusing on persistently used habitats in nearshore or offshore domains. We demonstrate that spatio‐temporal variability of seabird distribution and abundance is characterized by anomalous events embedded within trends. In addition to identifying the locations of persistence of seabird aggregations, we found significant declines in species richness and the density of sooty shearwater (Puffinus griseus) and Leach's storm petrel (Oceanodroma leucorhoa); in contrast, black‐footed albatross (Phoebastria nigripes) abundance appear to be increasing. This assessment provides a spatially‐explicit framework for future evaluations of biophysical drivers of seabird hotspots and their associations and impacts on forage fish and zooplankton populations.

Dissolved oxygen as a constraint on daytime deep scattering layer depth in the southern California current ecosystem

Climate change-induced ocean deoxygenation is expected to exacerbate hypoxic conditions in mesopelagic waters off the coast of southern California, with potentially deleterious effects for the resident fauna. In order to understand the possible impacts that the oxygen minimum zone expansion will have on these animals, we investigated the response of the depth of the deep scattering layer (i.e., upper and lower boundaries) to natural variations in midwater oxygen concentrations, light levels, and temperature over time and space in the southern California Current Ecosystem. We found that the depth of the lower boundary of the deep scattering layer (DSL) is most strongly correlated with dissolved oxygen concentration, and irradiance and oxygen concentration are the key variables determining the upper boundary. Based on our correlations and published estimates of annual rates of change to irradiance level and hypoxic boundary, we estimated the corresponding annual rate of change of DSL depths. If past trends continue, the upper boundary is expected to shoal at a faster rate than the lower boundary, effectively widening the DSL under climate change scenarios. These results have important implications for the future of pelagic ecosystems, as a change to the distribution of mesopelagic animals could affect pelagic food webs as well as biogeochemical cycles.

Grazer and viral impacts on microbial growth and mortality in the southern California Current Ecosystem

Protistan grazers and viruses are major agents of mortality in marine microbial communities with substantially different implications for food-web dynamics, carbon cycling and diversity maintenance. While grazers and viruses are typically studied independently, their impacts on microbial communities may be complicated by direct and indirect interactions of their mortality effects. Using a modification of the seawater dilution approach, we quantified growth and mortality rates for total phytoplankton and picophytoplankton populations (Prochlorococcus, Synechococcus, picoeukaryotes) at four contrasting sites in the California Current Ecosystem. Grazing mortality was significant in 10 of 15 cases, while viral effects were significant for 2 cases. Nonetheless, mortality estimates for the entire phytoplankton community based on chlorophyll a were 38 ± 13% higher when viral effects were included, relative to grazing alone. Mortality estimates for picophytoplankton varied in space and among groups. We also explored a potential methodological constraint of this method and hypothesize that heterotrophic bacteria may be affected by the dilution of their growth-sustaining substrates. For all picophytoplankton, estimates of grazing and viral mortality were inversely related within and across experiments. Indirect interactions among grazers and viruses may be important to consider if there are tradeoffs in the grazing and virus resistance strategies of prey/host cells.

Variability in diatom contributions to biomass, organic matter production and export across a frontal gradient in the California Current Ecosystem

AbstractIn the offshore waters of Southern California, submesoscale processes associated with fronts may stimulate phytoplankton blooms and lead to biomass shifts at multiple trophic levels. Here we report the results of a study on the cycling of biogenic silica (bSiO2) with estimates of the contributions of diatoms to primary and new production in water masses adjacent to (i.e., coastal or oceanic) and within an offshore front in the Southern California Current Ecosystem (CCE). The coastal and oceanic water were sampled in cyclonic and anticyclonic eddies, respectively, with the frontal water being an interaction region between the eddy types. Concentrations of bSiO2 varied by 25‐fold across the front, with concentrations in frontal waters 20–25% of those in coastal waters. Rates of biogenic silica production spanned an equally large range, with rates within the frontal region that were half those in the coastal regions. Contributions of diatoms to primary and new production were disproportionately higher than their contribution to autotrophic biomass in all areas, ranging from 5–8%, 19–30%, and 32–43% for both processes in the oceanic, frontal and coastal waters, respectively. Across the frontal area, diatoms could account for &lt;1.0%, 6–8%, and 44–72% of organic matter export in the oceanic, frontal and coastal waters, respectively. The results suggest that the regions of frontal interactions between eddies in the southern CCE can account for variability in diatom biomass, productivity and export over very short spatial scales that is comparable to the variability observed across the Pacific basin.

Mesopelagic fish biomass in the southern California current ecosystem

Mesopelagic fishes are the most common vertebrates on Earth, forming an important link between lower trophic levels and higher predators, and also between surface production and the deep sea. The biomass of these fishes is a key parameter for ecological modeling of oceanic ecosystems, but it is poorly known. The two most common methods to estimate the biomass of these fishes, acoustic and trawl surveys, are both sensitive to the ability of fishes to avoid nets. We show that size-dependent changes in trawl capture efficiency can affect acoustic estimates of biomass estimates 5-fold. We used both acoustic and trawl-based methods (informed by morphological data and acoustic modeling of individual backscattering) to estimate the biomass of mesopelagic fishes of southern California to be 25–37gm−2 of ocean surface, a comparable density to that of inshore epipelagic zooplanktivorous fishes. Our results indicate that mesopelagic fishes are likely to play a major role in regional food webs.

Temporal and spatial patterns of microbial community biomass and composition in the Southern California Current Ecosystem

As part of the California Current Ecosystem Long Term Ecological Research (CCE-LTER) Program, samples for epifluorescence microscopy and flow cytometry (FCM) were collected at ten ‘cardinal’ stations on the California Cooperative Oceanic Fisheries Investigations (CalCOFI) grid during 25 quarterly cruises from 2004 to 2010 to investigate the biomass, composition and size–structure of microbial communities within the southern CCE. Based on our results, we divided the region into offshore, and inshore northern and southern zones. Mixed-layer phytoplankton communities in the offshore had lower biomass (16±2μgCL−1; all errors represent the 95% confidence interval), smaller size-class cells and biomass was more stable over seasonal cycles. Offshore phytoplankton biomass peaked during the winter months. Mixed-layer phytoplankton communities in the northern and southern inshore zones had higher biomass (78±22 and 32±9μgCL−1, respectively), larger size-class cells and stronger seasonal biomass patterns. Inshore communities were often dominated by micro-size (20–200μm) diatoms; however, autotrophic dinoflagellates dominated during late 2005 to early 2006, corresponding to a year of delayed upwelling in the northern CCE. Biomass trends in mid and deep euphotic zone samples were similar to those seen in the mixed-layer, but with declining biomass with depth, especially for larger size classes in the inshore regions. Mixed-layer ratios of autotrophic carbon to chlorophyll a (AC:Chl a) had a mean value of 51.5±5.3. Variability of nitracline depth, bin-averaged AC:Chl a in the mixed-layer ranged from 40 to 80 and from 22 to 35 for the deep euphotic zone, both with significant positive relationships to nitracline depth. Total living microbial carbon, including auto- and heterotrophs, consistently comprised about half of particulate organic carbon (POC).

Modeling Physical-Biological Responses to Climate Change in the California Current System

Understanding the effects of climate change on planktonic ecosystems requires the synthesis of large, diverse data sets of variables that often interact in nonlinear ways. One fruitful approach to this synthesis is the use of numerical models. Here, we describe how models have been used to gain understanding of the physical-biological couplings leading to decadal changes in the southern California Current ecosystem. Moving from basin scales to local scales, we show how atmospheric, physical oceanographic, and biological dynamics interact to create long-term fluctuations in the dynamics of the California Current ecosystem.

Ecological Transitions in a Coastal Upwelling Ecosystem

The southern California Current Ecosystem (CCE) is a dynamic eastern boundary current ecosystem that is forced by ocean-atmosphere variability on interannual, multidecadal, and long-term secular time scales. Recent evidence suggests that apparent abrupt transitions in ecosystem conditions reflect linear tracking of the physical environment rather than oscillations between alternative preferred states. A space-for-time exchange is one approach that permits use of natural spatial variability in the CCE to develop a mechanistic understanding needed to project future temporal changes. The role of (sub)mesoscale frontal systems in altering rates of nutrient transport, primary and secondary production, export fluxes, and the rates of encounters between predators and prey is an issue central to this pelagic ecosystem and its future trajectory because the occurrence of such frontal features is increasing. © 2013 by The Oceanography Society. All rights reserved.

Stable Isotope Analysis Challenges Wasp-Waist Food Web Assumptions in an Upwelling Pelagic Ecosystem

Eastern boundary currents are often described as ‘wasp-waist’ ecosystems in which one or few mid-level forage species support a high diversity of larger predators that are highly susceptible to fluctuations in prey biomass. The assumption of wasp-waist control has not been empirically tested in all such ecosystems. This study used stable isotope analysis to test the hypothesis of wasp-waist control in the southern California Current large marine ecosystem (CCLME). We analyzed prey and predator tissue for δ13C and δ15N and used Bayesian mixing models to provide estimates of CCLME trophic dynamics from 2007–2010. Our results show high omnivory, planktivory by some predators, and a higher degree of trophic connectivity than that suggested by the wasp-waist model. Based on this study period, wasp-waist models oversimplify trophic dynamics within the CCLME and potentially other upwelling, pelagic ecosystems. Higher trophic connectivity in the CCLME likely increases ecosystem stability and resilience to perturbations.

Sharp gradients in phytoplankton community structure across a frontal zone in the California Current Ecosystem

Spatial variability of plankton biomass, community composition and size structurewas investigated across a strong frontal transition (A-Front) in the southernCalifornia Current Ecosystem in October 2008. Depth profiles were taken across a25-km transect of nine stations sampled semi-synoptically during one night andfor 3 days following drifter arrays in the adjacent water masses. Community com-positions are compared based on analyses by digital epifluorescence microscopy,flow cytometry and pigment composition by high-pressure liquid chromatography.Our results show three assemblages sharply delineated in space, with plankton atthe front being compositionally distinct and biomass elevated relative to either ofthe adjacent water masses. Depth-averaged chlorophyll a (Chl a) varied by a factorof 2.3 (0.35–0.81 mg Chl a L