David M. Checkley

Influence of ocean winds on the pelagic ecosystem in upwelling regions

Upwelling of nutrient-rich, subsurface water sustains high productivity in the oceans eastern boundary currents. These ecosystems support a rate of fish harvest nearly 100 times the global mean and account for >20% of the worlds 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.

Nitrogen isotope fractionation by oceanic zooplankton

Abstract The ratio of 15 N: 14 N 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 15 N-depleted ammonium and 15 N-enriched particulate matter that are, respectively, recycled in and exported from the euphotic zone and thus cause the low values of 15 N: 14 N in oligotrophic seas. Heretofore, this pattern was attributed to nitrogen-fixation by the phytoplankton. We measured the ratio of 15 N: 14 N in the bodies and excreted ammonium of zooplankters from the northwest Pacific Ocean and compared these values to the ratio of 15 N: 14 N 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 15 N: 14 N for the particulate matter suspended therein.

Elevated CO2 Enhances Otolith Growth in Young Fish

Acidification of the oceans may have unexpected effects on the development of bony structures in fish larvae. A large fraction of the carbon dioxide added to the atmosphere by human activity enters the sea, causing ocean acidification. We show that otoliths (aragonite ear bones) of young fish grown under high CO2 (low pH) conditions are larger than normal, contrary to expectation. We hypothesize that CO2 moves freely through the epithelium around the otoliths in young fish, accelerating otolith growth while the local pH is controlled. This is the converse of the effect commonly reported for structural biominerals.

Climate, fishing, and fluctuations of sardine and anchovy in the California Current

Since the days of Elton, population cycles have challenged ecologists and resource managers. Although the underlying mechanisms remain debated, theory holds that both density-dependent and density-independent processes shape the dynamics. One striking example is the large-scale fluctuations of sardine and anchovy observed across the major upwelling areas of the world. Despite a long history of research, the causes of these fluctuations remain unresolved and heavily debated, with significant implications for fisheries management. We here model the underlying causes of these fluctuations, using the California Current Ecosystem as a case study, and show that the dynamics, accurately reproduced since A.D. 1661 onward, are explained by interacting density-dependent processes (i.e., through species-specific life-history traits) and climate forcing. Furthermore, we demonstrate how fishing modifies the dynamics and show that the sardine collapse of the 1950s was largely unavoidable given poor recruitment conditions. Our approach provides unique insight into the origin of sardine–anchovy fluctuations and a knowledge base for sustainable fisheries management in the California Current Ecosystem and beyond.

Resilience and stability of a pelagic marine ecosystem

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

REFLICS: real-time flow imaging and classification system

Abstract. 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.

Essential ocean variables for global sustained observations of biodiversity and ecosystem changes

Sustained observations of marine biodiversity and ecosystems focused on specific conservation and management problems are needed around the world to effectively mitigate or manage changes resulting from anthropogenic pressures. These observations, while complex and expensive, are required by the international scientific, governance and policy communities to provide baselines against which the effects of human pressures and climate change may be measured and reported, and resources allocated to implement solutions. To identify biological and ecological essential ocean variables (EOVs) for implementation within a global ocean observing system that is relevant for science, informs society, and technologically feasible, we used a driver-pressure-state-impact-response (DPSIR) model. We (1) examined relevant international agreements to identify societal drivers and pressures on marine resources and ecosystems, (2) evaluated the temporal and spatial scales of variables measured by 100+ observing programs, and (3) analysed the impact and scalability of these variables and how they contribute to address societal and scientific issues. EOVs were related to the status of ecosystem components (phytoplankton and zooplankton biomass and diversity, and abundance and distribution of fish, marine turtles, birds and mammals), and to the extent and health of ecosystems (cover and composition of hard coral, seagrass, mangrove and macroalgal canopy). Benthic invertebrate abundance and distribution and microbe diversity and biomass were identified as emerging EOVs to be developed based on emerging requirements and new technologies. The temporal scale at which any shifts in biological systems will be detected will vary across the EOVs, the properties being monitored and the length of the existing time-series. Global implementation to deliver useful products will require collaboration of the scientific and policy sectors and a significant commitment to improve human and infrastructure capacity across the globe, including the development of new, more automated observing technologies, and encouraging the application of international standards and best practices.

Real-time detection and classification of objects in flowing water

This paper describes the design of a PC-based real-time machine vision system for detecting and classifying small marine organisms like fish eggs and planktons in flowing water. The system is called the Real-time FLow Imaging and Classification System, or ReFLICS for short, and it will automate the task of visually counting and classifying fish egg samples which is currently performed by trained humans. ReFLICS uses a line-scan image sensor to eliminate double counting and boundary effects. Using a combination of flowmeter and image-based flow error correction algorithm, ReFLICSs line-scan camera can work with changing flow. Design of the complete system from the camera and illumination housing to the machine vision software allows ReFLICS to work in the harsh environments of a ship at sea. Using an industry-standard multi-processor PC with PCI card pipeline image processor and running Microsoft Windows NT, ReFLICS can achieve the high performance required meanwhile maintaining relative low equipment, development, and maintenance costs. This paper provides the ReFLICS system design and presents initial results of the system.

Spatial patterns of Anchoveta (Engraulis ringens) eggs and larvae in relation to p CO 2 in the Peruvian upwelling system

Large and productive fisheries occur in regions experiencing or projected to experience ocean acidification. Anchoveta (Engraulis ringens) constitute the worlds largest single-species fishery and live in one of the oceans highest pCO2 regions. We investigated the relationship of the distribution and abundance of Anchoveta eggs and larvae to natural gradients in pCO2 in the Peruvian upwelling system. Eggs and larvae, zooplankton, and data on temperature, salinity, chlorophyll a and pCO2 were collected during a cruise off Peru in 2013. pCO2 ranged from 167–1392 µatm and explained variability in egg presence, an index of spawning habitat. Zooplankton abundance explained variability in the abundance of small larvae. Within the main spawning and larva habitats (6–10°S), eggs were found in cool, low-salinity, and both extremely low (less than 200 µatm) and high (more than 900 µatm) pCO2 waters, and larvae were collected in warmer, higher salinity, and moderate (400–600 µatm) pCO2 waters. Our data support the hypothesis that Anchoveta preferentially spawned at high pCO2 and these eggs had lower survival. Enhanced understanding of the influence of pCO2 on Anchoveta spawning and larva mortality, together with pCO2 measurements, may enable predictions of ocean acidification effects on Anchoveta and inform adaptive fisheries management.