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106-128 Ruzicka 11/17/07 8:50 AM Page 106 106-128 Ruzicka 11/17/07 8:50 AM Page 106 RUZICKA ET AL.: SEASONAL FOOD WEB MODELS CalCOFI Rep., Vol. 48, 2007 SEASONAL FOOD WEB MODELS FOR THE OREGON INNER-SHELF ECOSYSTEM: INVESTIGATING THE ROLE OF LARGE JELLYFISH JAMES J. RUZICKA RICHARD D. BRODEUR AND THOMAS C. WAINWRIGHT Cooperative Institute for Marine Resources Studies National Oceanic and Atmospheric Administration Oregon State University Northwest Fisheries Science Center 2030 SE Marine Science Drive Fisheries Ecology Division Newport, Oregon 97365 2030 SE Marine Science Drive [email protected] Newport, Oregon 97365 ABSTRACT Jellyfish biomass has increased dramatically in many We developed two seasonal food-web models, spring ecosystems around the world in the past two decades and summer, within the Ecopath framework for the (Mills 2001; Brodeur et al. 2002; Kawahara et al. 2006; Oregon upwelling ecosystem to investigate the role of Attrill et al. 2007). Jellyfish have several characteristics large jellyfish as competitors for zooplankton prey. We that place them in a unique and influential position used information about fish and jellyfish biomass, dis- within an ecosystem, which can have negative affects tribution, and diet derived from pelagic trawl survey upon pelagic fish: high rates of reproduction and growth, data. Information about lower trophic-level production generally broad diets that can overlap with planktivo- was acquired from zooplankton survey data. The mod- rous fish, and few predators. Increases in jellyfish bio- els indicate that in spring, jellyfish are a modest con- mass are generally accompanied by decreases in fish sumer of zooplankton, and forage fishes dominate the biomass (e.g., Lynam et al. 2006), which suggests sub- system in terms of biomass and consumption. By late stantial fish-jellyfish interactions that may affect fish summer, jellyfish become the major zooplankton con- growth, survival, and distribution. Thus, there is a rec- sumers, and they consume 17% of the summer zoo- ognized need to understand the role of jellyfish in pelagic plankton production while forage fish consume 9%. ecosystems, the causes of jellyfish proliferation, and the Jellyfish appear to divert zooplankton production away potential consequences to ecosystem functioning and to from upper trophic levels. Only 2% of the energy con- fisheries when jellyfish biomass blooms. Jellyfish may sumed by jellyfish is passed to higher trophic levels. have a negative impact upon pelagic fishes as both preda- However, the role of jellyfish as competitors may be tors and competitors (Purcell and Arai 2001). Jellyfish, moderate; a large proportion of zooplankton produc- in particular, can obtain a high biomass and may become tion (40%–44%) is not consumed but lost to detritus. an important energy pathway diverting zooplankton pro- duction away from pelagic fishes (Mills 1995, 2001; INTRODUCTION Lynam et al. 2006). The northern California Current (NCC) off Oregon Here, we examine the role of jellyfish in the NCC and Washington supports a seasonally productive and upwelling ecosystem off the Oregon coast. Jellyfish rep- open ecosystem. Upwelling-favorable winds dominate resent a major portion of the pelagic biomass in the along the Oregon and Washington coasts after the spring NCC (Shenker 1984; Suchman and Brodeur 2005), al- transition during March or April, and continue through though neither their long-term trends in biomass nor October or November when downwelling conditions their trophic role in the ecosystem has been well stud- normally occur (Strub et al. 1987). During the upwelling ied. Suchman et al. (in press) examined the diet of sev- season, the NCC is home to a diverse pelagic fish com- eral dominant jellyfish in this region and compared their munity, including the juveniles of important salmon consumption to available zooplankton. They found that stocks, resident species such as anchovies, smelts, and these species can have a major impact on production of herring, and transient species migrating from the south several zooplankton taxa. Brodeur et al.1 compared the such as sardines, hake, and mackerels (Brodeur et al. diets and distribution of these jellyfish to those of co- 2005). Ecosystem productivity and food-web structure occurring pelagic fishes and found that the potential for vary on seasonal-to-decadal time scales due to the tim- competitive interactions can be substantial due to high ing and strength of seasonal alongshore winds and forc- dietary and spatial overlap. ing by basin-scale physical processes (e.g., El Niño, Pacific The goals of this study are to: (1) develop two mass- Decadal Oscillation) and longer-term climate trends (Batchelder et al. 2002). These variations affect the sur- 1Brodeur, R. D., C. L. Suchman, D. C. Reese, T. W. Miller, and E. A. Daly. vival and productivity of all members of the pelagic com- Submitted-a. Spatial overlap and trophic interactions between pelagic fish and large jellyfish in the northern California Current. Mar. Biol. munity in the NCC. NOAA/NMFS/NWFSC, Newport, Oregon. 106 106-128 Ruzicka 11/17/07 8:50 AM Page 107 RUZICKA ET AL.: SEASONAL FOOD WEB MODELS CalCOFI Rep., Vol. 48, 2007 n balance food-web models of the northern California ͚ B • (Q/B) • DCji Current upwelling ecosystem with focus upon the pelagic j j sub-system using data from large-scale surveys for bio- j = 1 mass, distribution, seasonal patterns of biomass change, is the total predation mortality rate, and the term and local and contemporary diet information, (2) in- (P/B)i • Bi • EEi is the non-predation mortality rate vestigate change in trophic structure during the early (Christensen and Walters 2004). As input parameters for (spring) and late (summer) upwelling season, and (3) in- each trophic group, Ecopath requires the weight-spe- vestigate the importance of large jellyfish within the cific diet composition, the fishery harvest rate, and at northern California Current upwelling ecosystem, their least three of the following parameters: B, P/B, Q/B, impact on lower trophic levels, their importance as com- or EE. As an assumption of steady-state community petitors with planktivorous fishes, and their impact upon composition is not made in the seasonal models devel- higher trophic levels. oped here, biomass change rate (BA) from endemic growth and mortality and emigration (or immigration) MATERIALS AND METHODS rates (E) are also required. Ecopath also accounts for the energy flow within individual trophic groups: Model Overview consumption = production + respiration + egestion, (2) Two seasonal-scale food-web models have been de- veloped for the inner-shelf of the Oregon upwelling where egestion is assumed to be 20% for all groups in ecosystem within the Ecopath framework (Christensen the present models. and Walters 2004). The models represent the spring The two seasonal models presented here each consist (April-June) and summer (July-September) periods for of one producer group, 48 consumer groups, two egg a composite of the years 2000 and 2002; these are the groups, and three detritus groups. They are based upon most recent years during which pelagic fish surveys were the annual-scale northern California Current models conducted over the full North-South extent of the developed by Field and colleagues within the Ecopath Oregon shelf. The models’ domain extends from 46˚N framework (Field 2004; Field and Francis 2005; Field et to 41.8˚N (southern Oregon border) and excludes the al. 2006). The benthic food web (trophic groupings, mouth of the Columbia River which has its own dis- diet, physiological rate parameters) is modified from the tinct and important physical and ecological characteris- Field models as are the marine mammal and seabird tics (Hickey and Banas 2003). Offshore, the models groups. The information required to develop the pelagic extend to the 125 m isobath, encompassing an area of food web was obtained from a variety of sources: recent approximately 9,650 km2. pelagic fish and plankton survey studies off Oregon, local Ecopath is a software package for synthesizing diet, diet information, fishery records, the literature, and other production, and metabolic information into a mass-bal- northeast Pacific food-web models. anced system of interactions between all trophic groups that define an ecosystem (Christensen and Walters 2004). Community Composition The Ecopath master equation allocates the productivity BPA and GLOBEC pelagic trawl surveys: The of each trophic group to fishery harvest, transfer to higher composition of the pelagic nekton and jellyfish com- trophic level via predation, emigration out of the ecosys- munity on the Oregon shelf in spring and summer (2000 tem, growth, and other mortality (e.g., senescence): and 2002) was estimated from the Bonneville Power Administration (BPA) ocean salmon survey program and n the GLOBEC pelagic survey program. The BPA ocean Ϫ Ϫ Bi • (P/B)i ͚ Bj • (Q/B)j • DCji salmon survey sampled three transect lines in May, June, j = 1 and September from 45.7˚N to 44.6˚N and from the 30 m isobath onto the continental slope. The GLOBEC Ϫ Ϫ Ϫ (P/B)i • Bi • EEi Yi Ei BAi = 0 (1) survey consisted of four cruises (June 2000 and 2002, September 2000 and 2002) from 44.4˚N to 42˚N from where, for each trophic group (i), B is the biomass, P/B the 30 m isobath onto the continental slope. Both sur- is the mass-specific production rate, Q/B is the mass- vey programs quantitatively sampled the upper 20 m of × specific consumption rate, DCji is the fraction of prey the water-column using an 18 30 m Nordic Rope (i) in the diet of predator (j), Y is the fishery harvest trawl during daylight hours. Detailed trawl and sampling rate, E is the emigration rate, EE is the ecotrophic ef- protocol information for both programs are provided by ficiency (the fraction of production consumed within Emmett et al. (2005). The combined sampling area is the system), and BA is the biomass accumulation rate.
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