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Checkley, DM, Dagg MJ, Uye S.  1992.  Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus. Journal of Plankton Research. 14:71-96.   10.1093/plankt/14.1.71   AbstractWebsite

Diel variations in vertical distribution, gut pigment content, ammonium excretion and egg production were investigated for adult females of Acartia erythraea and A. pacifica in the vertically mixed Inland Sea of Japan and Centropages furcatus in the stratified, neritic Gulf of Mexico. Gut pigment content and egg production rate were maximal at night and ammonium excretion was maximal during the daytime. Neither A. erythraea nor A. pacifica adult females showed an apparent diel migration, but the former were highly concentrated in the sur-face layer during the afternoon. In contrast, C. furcatus adult females showed a clear diel migration, residing immediately above the bottom during the daytime and being concentrated between 10 and 25 m depth during the night-time. Individual-based data on gut content and excretion and egg production rates were combined with vertical-distribution data to calculate population values. In the Inland Sea of Japan, the resultant pattern for Acartia spp. reflected the diel variation in physiological rates and even distribution of adult females, except for the afternoon, surface aggregation of A. erythraea. In the Gulf of Mexico, the pattern for C. furcatus reflected largely the diel variation in each rate process and the heterogeneous distribution of adult females in the water column. Elevated nocturnal feeding activity of these copepods may be due to an endogenous rhythm. The daytime maximum in ammonium excretion and night-time maximum in egg production rate indicated approximate half-day and day time lags, respectively, after the intake of food until its conversion into dissolved excreta and released eggs.

Checkley, DM.  1980.  Food limitation of egg production by a marine, planktonic copepod in the sea off Southern California. Limnology and Oceanography. 25:991-998. AbstractWebsite

The in situ rate of egg production (B) and female size of Paracalanus parvus, a particle-grazing copepod, water temperature, and the concentrations of chlorophyll a and particulate nitrogen were measured in 31 collections from the euphotic zone of the sea off southern California. B was correlated positively with chlorophyll a and female size and negatively with temperature. A multiple regression of log B on chlorophyll a, particulate nitrogen, female size, and temperature accounts for44% ofthe variationin log B. B was predicted best by an empirical function of food concentration when the food available in nature was considered to be phytoplankton rather than of all types of particulate matter >5 µ.An index of immediate food limitation was derived from laboratory data as a function of food concentration. When applied to extensive measurements of chlorophyll a in the euphotic zone, this index indicates that Paracalanus was often food limited and that food limitation increased along an onshore-offshore transect. Paracalanus was rarely food limited in Santa Monica Bay.

Takahashi, M, Checkley Jr. DM.  2008.  Growth and survival of Pacific sardine (Sardinops sagax) in the California Current region. Journal of the Northwest Atlantic Fisheries Society. 41:129-136. Abstract
Checkley Jr., DM, Ayon P, Baumgartner TR, Bernal M, Coetzee JC, Emmett R, Guevara R, Hutchings L, Ibaibariaga L, Nakata H, Oozeki Y, Planque B, Schweigert J, Stratoudakis Y, Van der Lingen CD.  2009.  Habitats. Climate change and small pelagic fish. ( Checkley Jr. DM, Alheit J, Oozeki Y, Roy C, Eds.).:12-44., Cambridge, UK; New York: Cambridge University Press Abstract
Tommasi, D, Stock CA, Pegion K, Vecchi GA, Methot RD, Alexander MA, Checkley DM.  2017.  Improved management of small pelagic fisheries through seasonal climate prediction. Ecological Applications. 27:378-388.   10.1002/eap.1458   AbstractWebsite

Populations of small pelagic fish are strongly influenced by climate. The inability of managers to anticipate environment-driven fluctuations in stock productivity or distribution can lead to overfishing and stock collapses, inflexible management regulations inducing shifts in the functional response to human predators, lost opportunities to harvest populations, bankruptcies in the fishing industry, and loss of resilience in the human food supply. Recent advances in dynamical global climate prediction systems allow for sea surface temperature (SST) anomaly predictions at a seasonal scale over many shelf ecosystems. Here we assess the utility of SST predictions at this fishery relevant scale to inform management, using Pacific sardine as a case study. The value of SST anomaly predictions to management was quantified under four harvest guidelines (HGs) differing in their level of integration of SST data and predictions. The HG that incorporated stock biomass forecasts informed by skillful SST predictions led to increases in stock biomass and yield, and reductions in the probability of yield and biomass falling below socioeconomic or ecologically acceptable levels. However, to mitigate the risk of collapse in the event of an erroneous forecast, it was important to combine such forecast-informed harvest controls with additional harvest restrictions at low biomass.

Rykaczewski, RR, Checkley DM.  2008.  Influence of ocean winds on the pelagic ecosystem in upwelling regions. Proceedings of the National Academy of Sciences of the United States of America. 105:1965-1970.   10.1073/pnas.0711777105   AbstractWebsite

Upwelling of nutrient-rich, subsurface water sustains high productivity in the ocean's eastern boundary currents. These ecosystems support a rate of fish harvest nearly 100 times the global mean and account for >20% of the world's marine fish catch. Environmental variability is thought to be the major cause of the decadal-scale biomass fluctuations characteristic of fish populations in these regions, but the mechanisms relating atmospheric physics to fish production remain unexplained. Two atmospheric conditions induce different types of, upwelling in these ecosystems: coastal, alongshore wind stress, resulting in rapid upwelling (with high vertical velocity, w); and wind-stress curl, resulting in slower upwelling (low w). We show that the level of wind-stress curl has increased and that production of Pacific sardine (Sardinops sagax) varies with wind-stress curl over the past six decades. The extent of isopycnal shoaling, nutricline depth, and chlorophyll concentration in the upper ocean also correlate positively with wind-stress curl. The size structure of plankton assemblages is related to the rate of wind-forced upwelling, and sardine feed efficiently on small plankters generated by slow upwelling. Upwelling rate is a fundamental determinant of the biological structure and production in coastal pelagic ecosystems, and future changes in the magnitude and spatial gradient of wind stress may have important and differing effects on these ecosystems. Understanding of the biological mechanisms relating fisheries production to environmental variability is essential for wise management of marine resources under a changing climate.

Bax, NJ, Appeltans W, Brainard R, Duffy JE, Dunstan P, Hanich Q, Davies HH, Hills J, Miloslavich P, Muller-Karger FE, Simmons S, Aburto-Oropeza O, Batten S, Benedetti-Cecchi L, Checkley D, Chiba S, Fischer A, Garcia MA, Gunn J, Klein E, Kudela RM, Marsac F, Obura D, Shin YJ, Sloyan B, Tanhua T, Wilkin J.  2018.  Linking capacity development to GOOS monitoring networks to achieve sustained ocean observation. Frontiers in Marine Science. 5   10.3389/fmars.2018.00346   AbstractWebsite

Developing enduring capacity to monitor ocean life requires investing in people and their institutions to build infrastructure, ownership, and long-term support networks. International initiatives can enhance access to scientific data, tools and methodologies, and develop local expertise to use them, but without ongoing engagement may fail to have lasting benefit. Linking capacity development and technology transfer to sustained ocean monitoring is a win-win proposition. Trained local experts will benefit from joining global communities of experts who are building the comprehensive Global Ocean Observing System (GOOS). This two-way exchange will benefit scientists and policy makers in developing and developed countries. The first step toward the GOOS is complete: identification of an initial set of biological Essential Ocean Variables (EOVs) that incorporate the Group on Earth Observations (GEO) Essential Biological Variables (EBVs), and link to the physical and biogeochemical EOVs. EOVs provide a globally consistent approach to monitoring where the costs of monitoring oceans can be shared and where capacity and expertise can be transferred globally. Integrating monitoring with existing international reporting and policy development connects ocean observations with agreements underlying many countries' commitments and obligations, including under SDG 14, thus catalyzing progress toward sustained use of the ocean. Combining scientific expertise with international capacity development initiatives can help meet the need of developing countries to engage in the agreed United Nations (UN) initiatives including new negotiations for the conservation and sustainable use of marine biological diversity of areas beyond national jurisdiction, and the needs of the global community to understand how the ocean is changing.

Checkley Jr., DM.  2000.  Michael M. Mullen: A biography. PICES Press. 8:13-17. Abstract
Cambalik, JJ, Checkley DM, Kamykowski D.  1998.  A new method to measure the terminal velocity of small particles: A demonstration using ascending eggs of the Atlantic menhaden (Brevoortia tyrannus). Limnology and Oceanography. 43:1722-1727. AbstractWebsite

A new method, incorporating video, motion analysis, and a novel experimental apparatus, was used to measure the terminal velocity of particles. The method facilitated the investigation of treatment effects and maximized the number of measurements for each replicate, thus improving the statistics for a population of particles. The eggs of the Atlantic menhaden (Brevoortia tyrannus) were used to demonstrate the method by investigating the effects of salinity and stage of development on their ascent rate. Egg ascent rate was greatest at intermediate salinity (36.5 parts per thousand) and decreased in the late stage of embryonic development. We estimate eggs at oceanic salinities (>35.5 parts per thousand) in nature to ascend at 0.19-0.25 cm s(-1).

Checkley, DM, Miller CA.  1989.  Nitrogen isotope fractionation by oceanic zooplankton. Deep-Sea Research Part a-Oceanographic Research Papers. 36:1449-1456.   10.1016/0198-0149(89)90050-2   AbstractWebsite

The ratio of 15N:14N for particulate matter suspended in oceanic, surface waters is high after recent nitrate depletion and low in the stable, oligotrophic ocean. We hypothesize that zooplankters and other pelagic heterotrophs produce 15N-depleted ammonium and 15N-enriched particulate matter that are, respectively, recycled in and exported from the euphotic zone and thus cause the low values of 15N:14N in oligotrophic seas. Heretofore, this pattern was attributed to nitrogen-fixation by the phytoplankton.We measured the ratio of 15N:14N in the bodies and excreted ammonium of zooplankters from the northwest Pacific Ocean and compared these values to the ratio of 15N:14N for subeuphotic, dissolved nitrate. We report that oceanic zooplankton excrete ammonium that is isotopically light relative to their bodies and subeuphotic nitrate. These results are consistent with our hypothesis and the view that the phytoplankton of oligotrophic seas is nourished primarily by nitrogen recycled within the euphotic zone. Nitrate injected into the euphotic zone may be manifest and hence detected by an increase of the ratio 15N:14N for the particulate matter suspended therein.

Checkley Jr., DM.  1985.  Nitrogen limitation of zooplankton production and its effect on the marine nitrogen cycle. Archiv fuer Hydrobiologie - Beiheft Ergebnisse der Limnologie. 21:103-113. Abstract
Osgood, KE, Checkley DM.  1997.  Observations of a deep aggregation of Calanus pacificus in the Santa Barbara Basin. Limnology and Oceanography. 42:997-1001. AbstractWebsite

An optical plankton counter/CTD package was used with zooplankton net samples to map the distribution of fifth copepodid (C5) Calanus pacificus in the Santa Barbara Basin region during two autumn cruises. Diapausing C5 C. pacificus were aggregated in a layer just above the basin's oxygen-deficient bottom waters and below its sill depth. The maximal concentration measured was 6,900 ind. m(-3) from a net sample spanning a depth range double the thickness of the C5 layer Although the C5 concentration varied, the layer was found at all stations of sufficient bottom depth within the basin. During November 1994, C5 C. pacificus accounted for 95-97% of all zooplankton caught in net samples from the layer. Relatively low concentrations of deep-dwelling C5 C. pacificus were observed at nearby stations outside the basin. We hypothesize that C5 C. pacificus descend into the Santa Barbara Basin at diapause, are trapped, and accumulate in a region of relatively low predator abundance. The resultant aggregation is estimated to contain a significant fraction of the regional C. pacificus population and thus assumes an important role in its dynamics.

Gonzalez-Quiros, R, Checkley DM.  2006.  Occurrence of fragile particles inferred from optical plankton counters used in situ and to analyze net samples collected simultaneously. Journal of Geophysical Research-Oceans. 111   10.1029/2005jc003084   AbstractWebsite

[1] We hypothesized that the optical plankton counter (OPC) senses particles in situ that are not collected by nets and analyzed in the laboratory. An OPC was deployed in situ between 1998 and 2004 in the mouth of a bongo net with 505-mu m-mesh nets in the upper 210 m at stations in the California Current Region. Here we compare paired data sets from the OPC in situ and the OPC analysis in the laboratory of the simultaneously collected net samples for four seasons of 2 years. We restricted our analysis to particle sizes 1.26 - 6.35 mm equivalent spherical diameter (ESD), a size class shown from the lab OPC data to be retained efficiently by the net. On average, 4 (3) times more particles by number (volume) were sensed in situ by the OPC than sensed in net collections by the OPC in the lab. These values varied an order of magnitude among the eight cruises examined. Time of day, distance offshore, season, year, chlorophyll a concentration, and Brunt-Vaisala frequency each explained significant variation in these differences. The excess of particles sensed in situ over that measured in the net samples was due primarily to smaller particles in the 1.26 - 6.35 mm ESD range. We infer that the particles measured by the OPC in situ but not in the lab were fragile and thus not collected by the net. We hypothesize that these fragile particles are primarily aggregates and abandoned houses of larvaceans.

Shen, SG, Chen FY, Schoppik DE, Checkley DM.  2016.  Otolith size and the vestibulo-ocular reflex of larvae of white seabass Atractoscion nobilis at high pCO(2). Marine Ecology Progress Series. 553:173-183.   10.3354/meps11791   AbstractWebsite

We investigated vestibular function and otolith size (OS) in larvae of white seabass Atractoscion nobilis exposed to high partial pressure of CO2 (pCO(2)) The context for our study is the increasing concentration of CO2 in seawater that is causing ocean acidification (OA). The utricular otoliths are aragonitic structures in the inner ear of fish that act to detect orientation and acceleration. Stimulation of the utricular otoliths during head movement results in a behavioral response called the vestibulo-ocular reflex (VOR). The VOR is a compensatory eye rotation that serves to maintain a stable image during movement. VOR is characterized by gain (ratio of eye amplitude to head amplitude) and phase shift (temporal synchrony). We hypothesized that elevated pCO(2) would increase OS and affect the VOR. We found that the sagittae and lapilli of young larvae reared at 2500 mu atm pCO(2) (treatment) were 14 to 20% and 37 to 39% larger in area, respectively, than those of larvae reared at 400 mu atm pCO(2) (control). The mean gain of treatment larvae (0.39 +/- 0.05, n = 28) was not statistically different from that of control larvae (0.30 +/- 0.03, n = 20), although there was a tendency for treatment larvae to have a larger gain. Phase shift was unchanged. Our lack of detection of a significant effect of elevated pCO(2) on the VOR may be a result of the low turbulence conditions of the experiments, large natural variation in otolith size, calibration of the VOR or mechanism of acid-base regulation of white seabass larvae.

Jackson, GA, Checkley DM.  2011.  Particle size distributions in the upper 100 m water column and their implications for animal feeding in the plankton. Deep-Sea Research Part I-Oceanographic Research Papers. 58:283-297.   10.1016/j.dsr.2010.12.008   AbstractWebsite

We deployed autonomous particle-sensing SOLOPC floats more than eight times during five cruises, amassing almost 400 profiles of particle size (d > 90 mu m) and abundance between the ocean surface and 100 m. The profiles consistently had subsurface maxima in particle volume. The median (by volume) equivalent spherical diameter for the particle distribution was 0.4-0.8 mm and increased with depth in a manner similar to that observed in coagulation simulations. There was a sharp cutoff at the bottom of the high particle concentration region. Estimation of particle fluxes made using the size distributions show an increasing downward movement through the particle field above the sharp particle cutoff. The increase of particle flux with depth through the euphotic zone implies a partial spatial separation of production and consumption. The sharp drop in particle volume and flux implies that the base of the particle-rich zone is a region of active particle consumption, possibly by zooplankton flux feeding. Our data show greater concentrations of zooplankton-type particles relative to marine snow-type particles below the particle maximum. Such behavior could explain why zooplankton are frequently observed at and immediately below the particle maximum rather than the productivity maximum and suggests an important role for flux feeding in carbon and nutrient cycling at the base of the particle maximum. This implies that zooplankton act as gatekeepers for the movement of organic matter to the mesopelagic. The ability of the SOLOPC to sample hourly with high resolution in the upper 100 m of the ocean provides a powerful complement for the study of particles where it has been difficult to use sediment traps. (C) 2011 Elsevier Ltd. All rights reserved.

Checkley, DM, Barth JA.  2009.  Patterns and processes in the California Current System. Progress in Oceanography. 83:49-64.   10.1016/j.pocean.2009.07.028   AbstractWebsite

The California Current System (CCS) is forced by the distribution of atmospheric pressure and associated winds in relation to the west coast of North America. In this paper, we begin with a simplified case of winds and a linear coast, then consider variability characteristic of the CCS, and conclude by considering future change. The CCS extends from the North Pacific Current (similar to 50 degrees N) to off Baja California, Mexico (similar to 15-25 degrees N) with a major discontinuity at Point Conception (34.5 degrees N). Variation in atmospheric pressure affects winds and thus upwelling. Coastal, wind-driven upwelling results in nutrification and biological production and a southward coastal jet. Offshore, curl-driven upwelling results in a spatially large, productive habitat. The California Current flows equatorward and derives from the North Pacific Current and the coastal jet. Dominant modes of spatial and temporal variability in physical processes and biological responses are discussed. High surface production results in deep and bottom waters depleted in oxygen and enriched in carbon dioxide. Fishing has depleted demersal stocks more than pelagic stocks, and marine mammals, including whales, are recovering. Krill, squid, and micronekton are poorly known and merit study. Future climate change will differ from past change and thus prediction of the CCS requires an understanding of its dynamics. Of particular concern are changes in winds, stratification, and ocean chemistry. (C) 2009 Elsevier Ltd. All rights reserved.

Devries, AL, Checkley DM, Raymond JA.  1972.  Physiology and biochemistry of freezing resistance in Antarctic fishes. Antarctic Journal of the United States. 7:78-79. AbstractWebsite
Checkley Jr., DM, Cooper T, Lennert C.  1996.  Plankton pattern within and below the surface mixed layer. EOS Trans. AGU. 76:198. Abstract
Curtis, KA, Checkley DM, Pepin P.  2007.  Predicting the vertical profiles of anchovy (Engraulis mordax) and sardine (Sardinops sagax) eggs in the California Current System. Fisheries Oceanography. 16:68-84.   10.1111/j.1365-2419.2006.00414.x   AbstractWebsite

Several published models exist for simulating vertical profiles of pelagic fish eggs, but no one has rigorously assessed their capacity to explain observed variability. In this study, we applied a steady-state model, with four different formulations for vertical diffusivity, to northern anchovy (Engraulis mordax) and Pacific sardine (Sardinops sagax) eggs in the California Current region. Vertical mixing profiles, based on wind speed and hydrography, were combined with estimated terminal ascent velocities of the eggs based on measurements of egg buoyancy and size, to simulate the vertical profiles of the eggs. We evaluated model performance with two data sets: (1) vertically stratified tows for both species and (2) paired samples for sardine eggs from 3-m depth and in vertically integrated tows. We used two criteria: whether the model predicted individual observed vertical profiles (1) as well as the observed mean and (2) better than the observed mean. Model predictions made with the formulation producing the most gradual profile of vertical diffusivity provided the best match to observations from both data sets and for both species. Addition of a random error term to the terminal ascent velocity further improved prediction for anchovy eggs, but not sardine. For the paired data, model prediction of integrated abundance from abundance at 3-m depth had significantly lower mean square error than prediction based on a linear regression of 3 m on integrated abundance. Our results support the feasibility of using data from the Continuous Underway Fish Egg Sampler quantitatively as well as qualitatively in stock assessments.

Iwamoto, I, Trivedi MM, Checkley Jr. DM.  1998.  Real-time detection and classification of objects in flowing water. Machine vision systems for inspection and metrology VII : 4-5 November, 1998, Boston, Massachusetts . 3521( Batchelor BG, Miller JWV, Solomon S, Eds.).:214-220., Bellingham, Wash.: SPIE (International Society for Optical Engineering) Abstract
Iwamoto, S, Checkley DM, Trivedi MM.  2001.  REFLICS: Real-time flow imaging and classification system. Machine Vision and Applications. 13:1-13.   10.1007/pl00013270   AbstractWebsite

An accurate analysis of a large dynamic system like our oceans requires spatially fine and temporally matched data collection methods. Current methods to estimate fish stock size from pelagic (marine) fish egg abundance by using ships to take point samples of fish eggs have large margins of error due to spatial and temporal undersampling. The real-time flow imaging and classification system (REFLICS) enhances fish egg sampling by obtaining continuous, accurate information on fish egg abundance as the ship cruises along in the area of interest. REFLICS images the dynamic flow with a progressive-scan area camera (60 frames/s) and a synchronized strobe in backlighting configuration. Digitization and processing occur on a dual-processor Pentium II PC and a pipeline-based image-processing board. REFLICS uses a segmentation algorithm to locate fish-egg-like objects in the image and then a classifier to determine fish egg, species, and development stage (age). We present an integrated system design of REFLICS and performance results. REFLICS can perform in real time (60 Hz), classify fish eggs with low false negative rates on real data collected from a cruise, and work in harsh conditions aboard ships at sea. REFLICS enables cost-effective, real-time assessment of pelagic fish eggs for research and management.

Checkley, DM.  1984.  Relation of growth to ingestion for larvae of Atlantic herring Clupea harengus and other fish. Marine Ecology-Progress Series. 18:215-224.   10.3354/meps018215   AbstractWebsite

Larvae of Atlantic herring Clupea harengus were reared on wild plankton and Artemia salina nauplii in the laboratory at 7 to 9°C for 95 d. Between ages of 20 and 38 d, larvae were fed only Artemia nauplii and the specific rates of ingestion and growth were measured and compared. Relations of rate and efficiency of growth to ingestion were similar in terms of carbon and nitrogen. Growth was linearly related to ingestion (r2= 0.89, n = 9). Starved larvae lost mass at a specific rate of 0.03 d^-1 (3% d^-1) until death at 14 d. A specific ingestion rate of 0.04 d^-1 was required to balance defecation and metabolism. Gross growth efficiency (growth rate/ingestion rate) rose from -1.2 at a low ingestion rate (0.015 d^-1) to 0.4 at the greatest observed ingestion rate (0.11 d^-1) . Condition factor (dry weight length^-3) was significantly related to both ingestion rate and length (r2 = 0.69, n = 20).These results, combined with those for other fish larvae, indicate an asymptotic relation between rates of growth and ingestion such that gross growth efficiency is maximal (0.4) at intermediate ingestion rate. Fish larvae surviving in the sea appear to maximize their ingestion rate and thus grow rapidly but with a reduced efficiency.

Reiss, CS, Checkley DM, Bograd SJ.  2008.  Remotely sensed spawning habitat of Pacific sardine (Sardinops sagax) and Northern anchovy (Engraulis mordax) within the California Current. Fisheries Oceanography. 17:126-136.   10.1111/j.1365-2419.2008.00469.x   AbstractWebsite

We use trivariate kernel density estimation to define spawning habitat of northern anchovy (Engraulis mordax) and Pacific sardine (Sardinops sagax) in the California Current using satellite data and in situ egg samples from the Continuous Underway Fish Egg Sampler (CUFES) deployed during surveys in April by the California Cooperative Oceanic Fisheries Investigations (CalCOFI). Observed egg distributions were compared with monthly composite satellite sea surface temperature (SST) and surface chlorophyll a (chl a) data. Based on the preferred spawning habitat, as defined in SST and chl a space, the satellite data were used to predict potential spawning habitat along two areas of the west coast of North America. Data from the southern area (21.5 to 39 degrees N) were compared to observations from the CUFES data for the period 1998-2005. Northern anchovy and Pacific sardine exhibited distinctly different spawning habitat distributions. A significant relationship was found between satellite-based spawning area and that measured during surveys for sardine. CUFES area estimated for sardine was similar in magnitude to that estimated from satellite data (similar to 60 000 km(2)). In contrast, spawning habitat of anchovy averaged between 1000 and 200 000 km(2) for the period 1998-2005, for CUFES and satellite estimates, respectively. Interannual variability in the area (km(2)) and duration (months) of estimates of suitable habitat varied between species and between the northern (39 to 50.5 degrees N) and southern portions of the California Current. Long-term monitoring of habitat variability using remote sensing data is possible in the southern portion of the California Current, and could be improved upon in the northern area with the addition of surveys better timed to describe relationships between observed and estimated spawning habitats.

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.

Takahashi, M, Checkley DM, Litz MNC, Brodeur RD, Peterson WT.  2012.  Responses in growth rate of larval northern anchovy (Engraulis mordax) to anomalous upwelling in the northern California Current. Fisheries Oceanography. 21:393-404.   10.1111/j.1365-2419.2012.00633.x   AbstractWebsite

We examined variability in growth rate during the larval stage of northern anchovy (Engraulis mordax) in response to physical and biological environmental factors in 2005 and 2006. The onset of spring upwelling was anomalously delayed by 23 months until mid-July in 2005; in contrast, spring upwelling in 2006 began as a normal year in the northern California Current. Larval and early juvenile E. mordax were collected in August, September, and October off the coast of Oregon and Washington. Hatch dates ranged from May to September, with peaks in June and August in 2005 and a peak in July in 2006, based on the number of otolith daily increments. Back-calculated body length-at-age in the June 2005 hatch cohort was significantly smaller than in the August 2005 cohort, which had comparable growth to the July 2006 cohort. Standardized otolith daily increment widths as a proxy for seasonal variability in somatic growth rates in 2005 were negative until late July and then changed to positive with intensification of upwelling. The standardized increment width was a positive function of biomass of chlorophyll a concentration, and neritic cold-water and oceanic subarctic copepod species sampled biweekly off Newport, Oregon. Our results suggest that delayed upwelling in 2005 resulted in low food availability and, consequently, reduced E. mordax larval growth rate in early summer, but once upwelling began in July, high food availability enhanced larval growth rate to that typical of a normal upwelling year (e.g., 2006) in the northern California Current.