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2019
Nickels, CF, Sala LM, Ohman MD.  2019.  The euphausiid prey field for blue whales around a steep bathymetric feature in the southern California current system. Limnology and Oceanography. 64:390-405.   10.1002/lno.11047   AbstractWebsite

Euphausiids are important prey for many marine organisms and often occur in patchy aggregations. Euphausiid predators, such as blue whales, may become concentrated in the vicinity of these aggregations. We investigated an area called Nine Mile Bank (NMB) near San Diego, California, defined by an area of steep bathymetry, to determine whether the frequent whale sightings in that locality can be explained by the distribution of euphausiids across the bank and by the vertical distribution of euphausiids in the water column. Thysanoessa spinifera, the strongly preferred blue whale prey euphausiid in this area, was consistently more abundant on the bank or inshore of it than offshore. In contrast, Euphausia pacifica, a minor blue whale prey item, was much more abundant and distributed across the study region. Adults of both species were concentrated in a stratum corresponding to the feeding depth of blue whales. Other euphausiids that form a negligible part of the blue whale diet also showed no association with NMB. Both blue whales and their primary prey species Thysanoessa spinifera were more abundant on or inshore of the bank than offshore, suggesting that the bank may serve as an offshore limit of high prey abundance that helps to concentrate blue whales.

2018
Ohman, MD, Davis RE, Sherman JT, Grindley KR, Whitmore BM, Nickels CF, Ellen JS.  2018.  Zooglider: An autonomous vehicle for optical and acoustic sensing of zooplankton. Limnology and Oceanography: Methods.   10.1002/lom3.10301   Abstract

We present the design and preliminary results from ocean deployments of Zooglider, a new autonomous zooplankton-sensing glider. Zooglider is a modified Spray glider that includes a low-power camera (Zoocam) with telecentric lens and a custom dual frequency Zonar (200 and 1000 kHz). The Zoocam quantifies zooplankton and marine snow as they flow through a defined volume inside a sampling tunnel. Images are acquired on average every 5 cm from a maximum operating depth of ~ 400 m to the sea surface. Biofouling is mitigated using a dual approach: an ultraviolet light-emitting diode and a mechanical wiper. The Zonar permits differentiation of large and small acoustic backscatterers in larger volumes than can be sampled optically. Other sensors include a pumped conductivity, temperature, and depth unit and chlorophyll a fluorometer. Zooglider enables fully autonomous in situ measurements of mesozooplankton distributions, together with the three-dimensional orientation of organisms and marine snow in relation to other biotic and physical properties of the ocean water column. It is well suited to resolve thin layers and microscale ocean patchiness. Battery capacity supports 50 d of operations. Zooglider includes two-way communications via Iridium, permitting near-real–time transmission of data from each dive profile, as well as interactive instrument control from remote locations for adaptive sampling.

Stukel, MR, Biard T, Krause J, Ohman MD.  2018.  Large Phaeodaria in the twilight zone: Their role in the carbon cycle. Limnology and Oceanography. 63:2579-2594.   10.1002/lno.10961   AbstractWebsite

Advances in in situ imaging allow enumeration of abundant populations of large Rhizarians that compose a substantial proportion of total mesozooplankton biovolume. Using a quasi-Lagrangian sampling scheme, we quantified the abundance, vertical distributions, and sinking-related mortality of Aulosphaeridae, an abundant family of Phaeodaria in the California Current Ecosystem. Inter-cruise variability was high, with average concentrations at the depth of maximum abundance ranging from < 10 to > 300 cells m(-3), with seasonal and interannual variability associated with temperature-preferences and regional shoaling of the 10 degrees C isotherm. Vertical profiles showed that these organisms were consistently most abundant at 100-150 m depth. Average turnover times with respect to sinking were 4.7-10.9 d, equating to minimum in situ population growth rates of similar to 0.1-0.2 d(-1). Using simultaneous measurements of sinking organic carbon, we find that these organisms could only meet their carbon demand if their carbon : volume ratio were similar to 1 mu g C mm(-3). This value is substantially lower than previously used in global estimates of rhizarian biomass, but is reasonable for organisms that use large siliceous tests to inflate their cross-sectional area without a concomitant increase in biomass. We found that Aulosphaeridae alone can intercept > 20% of sinking particles produced in the euphotic zone before these particles reach a depth of 300 m. Our results suggest that the local (and likely global) carbon biomass of Aulosphaeridae, and probably the large Rhizaria overall, needs to be revised downwards, but that these organisms nevertheless play a major role in carbon flux attenuation in the twilight zone.

Lilly, LE, Ohman MD.  2018.  CCE IV: El Nino-related zooplankton variability in the southern California Current System. Deep-Sea Research Part I-Oceanographic Research Papers. 140:36-51.   10.1016/j.dsr.2018.07.015   AbstractWebsite

We analyzed seven El Nino events (springs 1958-59, 1983, 1992-93, 1998, 2003, 2010, and 2016) and the 2014-15 Pacific Warm Anomaly (spring 2015) for their impacts on zooplankton biomass and community composition in the southern sector of the California Current System (CCS). Although total mesozooplankton carbon biomass was only modestly affected during El Nino springs, community composition changed substantially. Carbon biomass of five major zooplankton taxa correlated negatively with San Diego sea level anomaly (SDSLA), a regional metric of El Nino physical impacts in the CCS. Additional taxa were negatively related to SDSLA via a time-lagged response reflected in an autoregressive-1 (AR-1) model. All five SDSLA-correlated taxa decreased in carbon biomass during most El Nino years compared to the surrounding years; the exception was the mild event of 2003. Principal Component Analysis revealed coherent species-level responses to El Nino within the euphausiids, copepods, and hyperiid amphipods. Percent similarity index (PSI) comparisons showed pronounced changes in the compositions of euphausiid and, to a lesser extent, calanoid copepod communities during El Nino. By grouping El Ninos into Eastern Pacific (EP) versus Central Pacific (CP) events based on their expressions along the equator, we found that CCS zooplankton assemblages showed a tendency toward greater changes in species composition during EP than CP El Ninos, although we had low statistical power for these comparisons. Several species showed consistent biomass changes across La Nina events as well, generally opposite in sign to El Nino responses, but overall community composition showed minimal change during La Nina. Carbon biomass and community composition returned to pre-Nino levels within 1-2 years following almost every event, suggesting high resilience of southern CCS zooplankton to El Nino perturbations to date.

Morrow, RM, Ohman MD, Goericke R, Kelly TB, Stephens BM, Stukel MR.  2018.  CCE V: Primary production, mesozooplankton grazing, and the biological pump in the California Current Ecosystem: Variability and response to El Nino. Deep-Sea Research Part I-Oceanographic Research Papers. 140:52-62.   10.1016/j.dsr.2018.07.012   AbstractWebsite

Predicting marine carbon sequestration in a changing climate requires mechanistic understanding of the processes controlling sinking particle flux under different climatic conditions. The recent occurrence of a warm anomaly (2014-2015) followed by an El Nino (2015-2016) in the southern sector of the California Current System presented an opportunity to analyze changes in the biological carbon pump in response to altered climate forcing. We compare primary production, mesozooplankton grazing, and carbon export from the euphotic zone during quasi-Lagrangian experiments conducted in contrasting conditions: two cruises during warm years - one during the warm anomaly in 2014 and one toward the end of El Nino 2016 - and three cruises during El Ninoneutral years. Results showed no substantial differences in the relationships between vertical carbon export and its presumed drivers (primary production, mesozooplankton grazing) between warm and neutral years. Mesozooplankton fecal pellet enumeration and phaeopigment measurements both showed that fecal pellets were the dominant contributor to export in productive upwelling regions. In more oligotrophic regions, fluxes were dominated by amorphous marine snow with negligible pigment content. We found no evidence for a significant shift in the relationship between mesozooplankton grazing rate and chlorophyll concentration. However, massspecific grazing rates were lower at low-to-moderate chlorophyll concentrations during warm years relative to neutral years. We also detected a significant difference in the relationship between phytoplankton primary production and photosynthetically active radiation between years: at similar irradiance and nutrient concentrations, productivity decreased during the warm events. Whether these changes resulted from species composition changes remains to be determined. Overall, our results suggest that the processes driving export remain similar during different climate conditions, but that species compositional changes or other structural changes require further attention.

Kahru, M, Jacox MG, Ohman MD.  2018.  CCE1: Decrease in the frequency of oceanic fronts and surface chlorophyll concentration in the California Current System during the 2014-2016 northeast Pacific warm anomalies. Deep-Sea Research Part I-Oceanographic Research Papers. 140:4-13.   10.1016/j.dsr.2018.04.007   AbstractWebsite

Oceanic fronts are sites of increased vertical exchange that are often associated with increased primary productivity, downward flux of organic carbon, and aggregation of plankton and higher trophic levels. Given the influence of fronts on the functioning of marine ecosystems, an improved understanding of the spatial and temporal variability of frontal activity is desirable. Here, we document changes in the frequency of sea-surface fronts and the surface concentration of chlorophyll-a (Chla) in the California Current System that occurred during the Northeast Pacific anomalous warming of 2014-2015 and El Nino of 2015-2016, and place those anomalies in the context of two decades of variability. Frontal frequency was detected with the automated histogram method using datasets of sea-surface temperature (SST) and Chla from multiple satellite sensors. During the anomalous 2014-2016 period, a drop in the frequency of fronts coincided with the largest negative Chla anomalies and positive SST anomalies in the whole period of satellite observations (1997-2017 for Chla and 1982-2017 for SST). These recent reductions in frontal frequency ran counter to a previously reported increasing trend, though it remains to be seen if they represent brief interruptions in that trend or a reversal that will persist going forward.

Nickels, CF, Ohman MD.  2018.  CCEIII: Persistent functional relationships between copepod egg production rates and food concentration through anomalously warm conditions in the California Current Ecosystem. Deep-Sea Research Part I-Oceanographic Research Papers. 140:26-35.   10.1016/j.dsr.2018.07.001   AbstractWebsite

We hypothesized that copepod egg production rates would be suppressed relative to their normal functional relationships with food availability during the NE Pacific warm anomalies of 2014-2015 and the El Nino event of 2015-2016. We also hypothesized that copepods would show increased rates of egg production and recruitment in association with ocean fronts. Three species of copepods (Calanus pacificus, Metridia pacifica, and Eucalanus caltfornicus) were incubated at sea in simulated in situ conditions to ascertain egg production rates and egg hatching success during a sequence of 3-5 California Current Ecosystem-Long Term Ecological Research (CCE-LTER) cruises across diverse ocean conditions. Several indices of food availability were measured. The warm events had no significant effect on egg production rates measured in the experiments conducted here. Egg production and net naupliar production rates varied markedly among copepod species and across spatial gradients, but relationships with food concentration during warm events did not consistently diverge from functional relationships during El Nino-neutral conditions. Egg production was most commonly elevated inshore compared to offshore, but was not elevated within fronts. The egg production of C. pacificus was best explained by total Chl-a, while E. californicus was best explained by the > 20 size fraction of Chl-a. M. pacifica showed no relationship between egg production and any food metric tested. C. pacificus showed higher EPR than the other two species in virtually all conditions encountered, which may contribute to its numerical dominance throughout the region.

Nickels, CF, Sala LM, Ohman MD.  2018.  The morphology of euphausiid mandibles used to assess selective predation by blue whales in the southern sector of the California Current System. Journal of Crustacean Biology. 38:563-573.   10.1093/jcbiol/ruy062   AbstractWebsite

We describe the mandibular morphology of the eight most abundant euphausiid species in the California Current and report regression relationships between mandible size and body total length. We applied these species-specific characters to the mandibles recovered from fecal samples of 18 blue whales (Balaenoptera musculus (Linnaeus, 1758)) collected between 1998 to 2015 off Southern California to test for selective feeding on the euphausiid assemblage. The diets of blue whales were consistently and overwhelmingly dominated by the large neritic euphausiid Thysanoessa spinifera Holmes, 1900, even when other species were present or dominant in closely collected net samples. More than 99% of the ingested euphausiids were longer than 10 mm, indicating that blue whales in this region are highly selective by prey species and size class, and dependent upon aggregations of juveniles or adults of a limited number of coastally associated euphausiid species.

Biard, T, Krause JW, Stukel MR, Ohman MD.  2018.  The significance of giant phaeodarians (rhizaria) to biogenic silica export in the California Current ecosystem. Global Biogeochemical Cycles. 32:987-1004.   10.1029/2018gb005877   AbstractWebsite

In marine ecosystems, many planktonic organisms precipitate biogenic silica (bSiO(2)) to build silicified skeletons. Among them, giant siliceous rhizarians (>500 mu m), including Radiolaria and Phaeodaria, are important contributors to oceanic carbon pools but little is known about their contribution to the marine silica cycle. We report the first analyses of giant phaeodarians to bSiO(2) export in the California Current Ecosystem. We measured the silica content of single rhizarian cells ranging in size from 470 to 3,920 mu m and developed allometric equations to predict silica content (0.37-43.42 mu g Si/cell) from morphometric measurements. Using sediment traps to measure phaeodarian fluxes from the euphotic zone on four cruises, we calculated bSiO(2) export produced by two families, the Aulosphaeridae and Castanellidae. Biogenic silica export ranged from <0.01 to 0.63 mmol Si.m(-2).day(-1). These two families alone contributed on average 10% (range 0-80%) of total bSiO(2) export from the euphotic zone. Their proportional contributions increased substantially in more oligotrophic regions with lower bSiO(2) fluxes. Using the in situ Underwater Vision Profiler 5, we characterized vertical distributions of the giant phaeodarian family Aulosphaeridae to a depth of 500 m and inferred their contribution to bSiO(2) export in deeper waters. We found a significant increase of Aulosphaeridae export (<0.01 to 2.82 mmol Si.m(-2).day(-1)) when extended to mesopelagic depths. Using a global data set of in situ profiles, we estimated the significance of Aulosphaeridae to bSiO(2) export and revealed that they can act as major exporters of bSiO(2) to the mesopelagic zone in various regions.

Lindegren, M, Checkley DM, Koslow JA, Goericke R, Ohman MD.  2018.  Climate-mediated changes in marine ecosystem regulation during El Nino. Global Change Biology. 24:796-809.   10.1111/gcb.13993   AbstractWebsite

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 (similar to 60years) 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 Nino. Furthermore, we show that biota respond differently to major El Nino 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 Nino.

2017
Stukel, MR, Aluwihare LI, Barbeau KA, Chekalyuk AM, Goericke R, Miller AJ, Ohman MD, Ruacho A, Song H, Stephens BM, Landry MR.  2017.  Mesoscale ocean fronts enhance carbon export due to gravitational sinking and subduction. Proceedings of the National Academy of Sciences of the United States of America. 114:1252-1257.   10.1073/pnas.1609435114   AbstractWebsite

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 U-238:Th-234 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 wasmechanistically 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 similar to 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.

2016
Brandon, J, Goldstein M, Ohman MD.  2016.  Long-term aging and degradation of microplastic particles: Comparing in situ oceanic and experimental weathering patterns. Marine Pollution Bulletin. 110:299-308.   10.1016/j.marpolbul.2016.06.048   AbstractWebsite

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 > 18 months, with some >30 months. Inferred particle weathering times are consistent with ocean circulation models suggesting a long residence time in the open ocean. (C) 2016 Elsevier Ltd. All rights reserved.

Ohman, MD, Romagnan JB.  2016.  Nonlinear effects of body size and optical attenuation on Diel Vertical Migration by zooplankton. Limnology and Oceanography. 61:765-770.   10.1002/lno.10251   AbstractWebsite

We adopt a trait-based approach to explain Diel Vertical Migration (DVM) across a diverse assemblage of planktonic copepods, utilizing body size as a master trait. We find a reproducible pattern of body size-dependence of day and night depths occupied, and of DVM. Both the smallest surface-dwelling and the largest deeper-dwelling copepods refrain from migrations, while intermediate-sized individuals show pronounced DVM. This pattern apparently arises as a consequence of size-dependent predation risk. In the size classes of migratory copepods the amplitude of DVM is further modulated by optical attenuation in the ocean water column because increased turbidity decreases encounter rates with visually hunting predators. Long-term changes in the ocean optical environment are expected to alter the vertical distributions of many copepods and thus to affect predator-prey encounters as well as oceanic carbon export.

Lindegren, M, Checkley DM, Ohman MD, Koslow JA, Goericke R.  2016.  Resilience and stability of a pelagic marine ecosystem. Proceedings of the Royal Society B-Biological Sciences. 283   10.1098/rspb.2015.1931   AbstractWebsite

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.

2015
Stukel, MR, Kahru M, Benitez-Nelson CR, Decima M, Goericke R, Landry MR, Ohman MD.  2015.  Using Lagrangian-based process studies to test satellite algorithms of vertical carbon flux in the eastern North Pacific Ocean. Journal of Geophysical Research-Oceans. 120:7208-7222.   10.1002/2015jc011264   AbstractWebsite

The biological carbon pump is responsible for the transport of similar to 5-20 Pg C yr(-1) from the surface into the deep ocean but its variability is poorly understood due to an incomplete mechanistic understanding of the complex underlying planktonic processes. In fact, algorithms designed to estimate carbon export from satellite products incorporate fundamentally different assumptions about the relationships between plankton biomass, productivity, and export efficiency. To test the alternate formulations of export efficiency in remote-sensing algorithms formulated by Dunne et al. (2005), Laws et al. (2011), Henson et al. (2011), and Siegel et al. (2014), we have compiled in situ measurements (temperature, chlorophyll, primary production, phytoplankton biomass and size structure, grazing rates, net chlorophyll change, and carbon export) made during Lagrangian process studies on seven cruises in the California Current Ecosystem and Costa Rica Dome. A food-web based approach formulated by Siegel et al. (2014) performs as well or better than other empirical formulations, while simultaneously providing reasonable estimates of protozoan and mesozooplankton grazing rates. By tuning the Siegel et al. (2014) algorithm to match in situ grazing rates more accurately, we also obtain better in situ carbon export measurements. Adequate representations of food-web relationships and grazing dynamics are therefore crucial to improving the accuracy of export predictions made from satellite-derived products. Nevertheless, considerable unexplained variance in export remains and must be explored before we can reliably use remote sensing products to assess the impact of climate change on biologically mediated carbon sequestration.

Powell, JR, Ohman MD.  2015.  Changes in zooplankton habitat, behavior, and acoustic scattering characteristics across glider-resolved fronts in the Southern California Current System. Progress in Oceanography. 134:77-92.   10.1016/j.pocean.2014.12.011   AbstractWebsite

We report cross-frontal changes in the characteristics of plankton proxy variables measured by autonomous Spray ocean gliders operating within the Southern California Current System (SCCS). A comparison of conditions across the 154 positive frontal gradients (i.e., where density of the surface layer decreased in the offshore direction) identified from six years of continuous measurements showed that waters on the denser side of the fronts typically showed higher Chl-a fluorescence, shallower euphotic zones, and higher acoustic backscatter than waters on the less dense side. Transitions between these regions were relatively abrupt. For positive fronts the amplitude of Diel Vertical Migration (DVM), inferred from a 3-beam 750 kHz acoustic Doppler profiler, increased offshore of fronts and covaried with optical transparency of the water column. Average interbeam variability in acoustic backscatter also changed across many positive fronts within 3 depth strata (0-150 m, 150-400 m, and 400-500 m), revealing a front related change in the acoustic scattering characteristics of the assemblages. The extent of vertical stratification of distinct scattering assemblages was also more pronounced offshore of positive fronts. Depth-stratified zooplankton samples collected by Mocness nets corroborated the autonomous measurements, showing copepod-dominated assemblages and decreased zooplankton body sizes offshore and euphausiid-dominated assemblages with larger median body sizes inshore of major frontal features. (C) 2014 Elsevier Ltd. All rights reserved.

Bednarsek, N, Ohman MD.  2015.  Changes in pteropod distributions and shell dissolution across a frontal system in the California Current System. Marine Ecology Progress Series. 523:93-103.   10.3354/meps11199   AbstractWebsite

We tested the sensitivity of the vertical distributions and shell dissolution patterns of thecosome pteropods to spatial gradients associated with an eddy-associated front in the southern California Current System. The aragonite saturation horizon (Omega(arag) = 1.0) shoaled from > 200 to <75 m depth across the front. The vertical distribution of thecosome pteropods tracked these changes, with all 5 species showing reduced occurrence at depths below 100 m where waters were less saturated with respect to aragonite. Shell dissolution patterns of the numerically dominant thecosome Limacina helicina corresponded to the cross-frontal changes in Omega(arag) saturation state. Severe shell dissolution ( categorized here as Type II and Type III) was low in near-surface waters where Omega(arag) > 1.4, while peak dissolution occurred in depths where Omega(arag) = 1.0 to 1.4. Vertical habitat compression and increased shell dissolution may be expected to accompany future shoaling of waters that are undersaturated with respect to aragonite.

Sydeman, WJ, Thompson SA, Santora JA, Koslow JA, Goericke R, Ohman MD.  2015.  Climate-ecosystem change off southern California: Time-dependent seabird predator-prey numerical responses. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 112:158-170.   10.1016/j.dsr2.2014.03.008   AbstractWebsite

Climate change may increase both stratification and upwelling in marine ecosystems, but these processes may affect productivity in opposing or complementary ways. For the Southern California region of the California Current Ecosystem (CCE), we hypothesized that changes in stratification and upwelling have affected marine bird populations indirectly through changes in prey availability. To test this hypothesis, we derived trends and associations between stratification and upwelling, the relative abundance of potential prey including krill and forage fish, and seabirds based on the long-term, multi-disciplinary CalCOFI/CCE-LTER program. Over the period 1987 through 2011, spring and summer seabird density (all species combined) declined by similar to 2% per year, mostly in the northern sector of the study region. Krill showed variable trends with two species increasing and one deceasing, resulting in community reorganization. Nearshore forage fish, dominated by northern anchovy (Engraulis mordax) as well as offshore mesopelagic species, show declines in relative abundance over this period. The unidirectional decline in springtime seabird density is largely explained by declining nearshore fish abundance in the previous season (winter). Interannual variability in seabird density, especially in the 2000s, is explained by variability in krill abundance. Changes in the numerical responses of seabirds to prey abundance correspond to a putative ecosystem shift in 1998-1999 and support aspects of optimal foraging (diet) theory. Predator-prey interactions and numerical responses clearly explain aspects of the upper trophic level patterns of change in the pelagic ecosystem off southern California. (C) 2014 Elsevier Ltd. All rights reserved.

Powell, JR, Ohman MD.  2015.  Covariability of zooplankton gradients with glider-detected density fronts in the Southern California Current System. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 112:79-90.   10.1016/j.dsr2.2014.04.002   AbstractWebsite

Fronts represent sharp boundaries between water masses, but seasonal and interannual variation in their occurrence and effects on the distributions of pelagic organisms are poorly understood. This study reports results from six years of ocean front observations (2006-2011) along two transect lines across the Southern California Current System (SCCS) using autonomous Spray ocean gliders. During this time, 154 positive near-surface density fronts were identified within 124 completed transects consisting of nearly 23,000 vertical profiles. The incidence of surface density fronts showed distinct seasonality along line 80 off Pt. Conception, with fewer fronts occurring during winter months and more numerous fronts in the nearshore and during spring, summer and fall. On line 90, fronts were the least common nearshore and most frequent in a transitional region offshore. Horizontal density gradients in the surface layer (0-50 m) were significantly correlated with horizontal gradients in surface layer Chlorophyll-a (Chl-a) fluorescence, as well as with mean volume backscatter (MVBS) recorded by a 750 kHz acoustic Doppler profiler. Density fronts were not only zones of rapidly changing phytoplankton and zooplankton biomass concentrations, but also more likely to be zones of enhanced acoustic backscatter and Chl-a fluorescence than regions flanking the fronts. MVBS and Chl-a gradients were significantly correlated with gradients in other hydrographic variables such as temperature, salinity, and spiciness, and weakly with cross-track current velocity, though density gradients remained the single best predictor of strong MVBS and fluorescence gradients. Large mobile predators foraging in the vicinity of such features could locate habitat with higher zooplankton biomass concentrations up to 85% of the time by traveling up local density gradients (i.e., toward rather than away from denser surface waters). We discuss implications of these results in the context of long-term trends in ocean fronts in the SCCS. (C) 2014 Elsevier Ltd. All rights reserved.

2014
Kang, YS, Ohman MD.  2014.  Comparison of long-term trends of zooplankton from two marine ecosystems across the North Pacific: Northeastern Asian marginal sea and Southern California current system. California Cooperative Oceanic Fisheries Investigations Reports. 55:169-182. AbstractWebsite

Long-term trends of zooplankton biomass (1968-2009) and major zooplankton taxa (1978-2009) were examined across the North Pacific in the Northeastern Asian Marginal Sea (NeAMS) and the Southern California Current System (SCC) to test for evidence of basin-scale synchrony. Zooplankton biomass showed contrasting long-term patterns in the two regions: an increasing trend (as wet mass) in the NeAMS, but a decreasing trend (as displacement volume) in the SCC. Zooplankton biomass covaried with the Pacific Decadal Oscillation in the NeAMS, but with the North Pacific Gyre Oscillation in the SCC. In the NeAMS, increasing zooplankton biomass was closely associated with increases of all major zooplankton groups (copepods, chaetognaths, euphausiids, and hyperiid amphipods). In the SCC, decreasing zooplankton biomass was caused by declining tunicates and chaetognaths. Seasonal cycles and responses to El Nino also differed between the two regions. In this cross-basin comparison, zooplankton showed differing patterns that reflect region-specific physical and biotic processes rather than synchronous responses to large-scale atmosphere-ocean forcing.

Koslow, AJ, Davison P, Lara-Lopez A, Ohman MD.  2014.  Epipelagic and mesopelagic fishes in the southern California Current System: Ecological interactions and oceanographic influences on their abundance. Journal of Marine Systems. 138:20-28.   10.1016/j.jmarsys.2013.09.007   Abstract

We use zooplankton and ichthyoplankton data from the ~60-year CalCOFI time series to examine relationships of mesopelagic (i.e. midwater) fishes in the California Current System with midwater predators, potential competitors (epipelagic planktivorous fishes) and zooplankton prey, within the context of local and basin-scale oceanography. Equilibrium-based near-steady state models and the “wasp-waist” paradigm for eastern boundary currents predict tightly-coupled trophic interactions, with negative correlations between the abundance of planktivorous competitors and between dominant planktivores and their prey. Testing these hypotheses with the CalCOFI time series, we found them to be generally invalid. Potential competitors within the mesopelagic community (planktivorous vertical migrators (VMs) and non-migrators (NMs)) were highly positively correlated, as were these groups with the mesopelagic piscivores (e.g. dragonfishes) that prey on them. In addition, the abundance of VMs was mostly positively correlated with that of epipelagic planktivores, such as anchovy, mackerels and hake. The VMs and epipelagic planktivores were negatively correlated with key potential planktonic prey groups, indicating a lack of bottom-up forcing. However, neither do these negative correlations appear to signify top-down forcing, since they seem to be mediated through correlations with key environmental drivers, such as the Pacific Decadal Oscillation (PDO), sea surface temperature, and the relative strength of the California Current. We suggest that the web of correlations linking key meso- and epipelagic planktivores, their predators and prey is mediated through common links with basin-scale oceanographic drivers, such as the PDO and ENSO cycles. Thus, the abundance of mesopelagic fishes in the California Current is closely tied to variation in the oxygen minimum zone, whose dynamics have been linked to the PDO. The PDO and other drivers are also linked to the transport of the California Current System, which influences the abundance of many dominant taxa off southern California that have broad biogeographic distributions linked to water masses that extend to the north (Transition Zone/sub-Arctic faunas) or the south (tropical/subtropical faunas).

Smith, KL, Sherman AD, Huffard CL, McGill PR, Henthorn R, Von Thun S, Ruhl HA, Kahru M, Ohman MD.  2014.  Large salp bloom export from the upper ocean and benthic community response in the abyssal northeast Pacific: Day to week resolution. Limnology and Oceanography. 59:745-757.   10.4319/lo.2014.59.3.0745   AbstractWebsite

A large bloom of Salpa spp. in the northeastern Pacific during the spring of 2012 resulted in a major deposition of tunics and fecal pellets on the seafloor at similar to 4000 m depth (Sta. M) over a period of 6 months. Continuous monitoring of this food pulse was recorded using autonomous instruments: sequencing sediment traps, a time-lapse camera on the seafloor, and a bottom-transiting vehicle measuring sediment community oxygen consumption (SCOC). These deep-sea measurements were complemented by sampling of salps in the epipelagic zone by California Cooperative Ocean Fisheries Investigations. The particulate organic carbon (POC) flux increased sharply beginning in early March, reaching a peak of 38 mg C m(-2) d(-1) in mid-April at 3400 m depth. Salp detritus started appearing in images of the seafloor taken in March and covered a daily maximum of 98% of the seafloor from late June to early July. Concurrently, the SCOC rose with increased salp deposition, reaching a high of 31 mg C m(-2) d(-1) in late June. A dominant megafauna species, Peniagone sp. A, increased 7-fold in density beginning 7 weeks after the peak in salp deposition. Estimated food supply from salp detritus was 97-327% of the SCOC demand integrated over the 6-month period starting in March 2012. Such large episodic pulses of food sustain abyssal communities over extended periods of time.

Martz, T, Send U, Ohman MD, Takeshita Y, Bresnahan P, Kim HJ, Nam S.  2014.  Dynamic variability of biogeochemical ratios in the Southern California Current System. Geophysical Research Letters. 41:2496-2501.   10.1002/2014gl059332   AbstractWebsite

We use autonomous nitrate (NO3-), oxygen (O-2), and dissolved inorganic carbon (DIC) observations to examine the relationship between ratios of C:N:O at an upwelling site in the Southern California Current System. Mean ratios and 95% confidence intervals observed by sensors over 8 months were NO3-:O-2=-0.110.002, NO3-:DIC=0.140.001, and DIC:O-2=-0.830.01, in good agreement with Redfield ratios. Variability in the ratios on the weekly time scale is attributable to shifts in biological demand and nutrient availability and shown to exhibit a spectrum of values ranging from near 100% New Production to 100% Regenerated Production.

2013
Di Lorenzo, E, Ohman MD.  2013.  A double-integration hypothesis to explain ocean ecosystem response to climate forcing. Proceedings of the National Academy of Sciences. 110:2496-2499.   10.1073/pnas.1218022110   AbstractWebsite

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.

Combes, V, Chenillat F, Di Lorenzo E, Riviere P, Ohman MD, Bograd SJ.  2013.  Cross-shore transport variability in the California Current: Ekman upwelling vs. eddy dynamics. Progress in Oceanography. 109:78-89.   10.1016/j.pocean.2012.10.001   AbstractWebsite

The low-frequency dynamics of coastal upwelling and cross-shelf transport in the Central and Southern California Current System (CCS) are investigated using the Regional Ocean Modeling System (ROMS) over the period 1965-2008. An ensemble of passive tracers released in the numerical model is used to characterize the effects of linear (Ekman upwelling) and non-linear (mesoscale eddies) circulation dynamics on the statistics of advection of coastal waters. The statistics of passive tracers released in the subsurface show that the low-frequency variability of coastal upwelling and cross-shelf transport of the upwelled water mass are strongly correlated with the alongshore wind stress, and are coherent between the central and southern CCS. However, the offshore transport of tracers released at the surface is not coherent between the two regions, and is modulated by intrinsic mesoscale eddy activity, in particular cyclonic eddies. The transport of cyclonic eddies extends with depth and entrains water masses of southern origin, advected by the poleward California Undercurrent (CUC). The CUC water masses are not only entrained by eddies but also constitute a source for the central California upwelling system. The interplay between intrinsic (eddy activity) and deterministic (Ekman upwelling) dynamics in controlling the cross-shelf exchanges in the CCS may provide an improved framework to understand and interpret nutrients and ecosystem variability. Published by Elsevier Ltd.