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The Official Magazine of The OceTHE OFFICIALa MAGAZINEnog OF THE OCEANOGRAPHYra SOCIETYphy CITATION Ruzicka, J.J., J.H. Steele, S.K. Gaichas, T. Ballerini, D.J. Gifford, R.D. Brodeur, and E.E. Hofmann. 2013. Analysis of energy flow in US GLOBEC ecosystems using end-to-end models. Oceanography 26(4):82–97, http://dx.doi.org/10.5670/oceanog.2013.77. DOI http://dx.doi.org/10.5670/oceanog.2013.77 COPYRIGHT This article has been published inOceanography , Volume 26, Number 4, a quarterly journal of The Oceanography Society. Copyright 2013 by The Oceanography Society. All rights reserved. USAGE Permission is granted to copy this article for use in teaching and research. Republication, systematic reproduction, or collective redistribution of any portion of this article by photocopy machine, reposting, or other means is permitted only with the approval of The Oceanography Society. Send all correspondence to: [email protected] or The Oceanography Society, PO Box 1931, Rockville, MD 20849-1931, USA. DOWNLOADED FROM HTTP://WWW.TOS.ORG/OCEANOGRAPHY SPECIAL ISSUE ON US GLOBEC: UNDERSTANDING CLIMATE IMPACTS ON MARINE ECOSYSTEMS Analysis of Energy Flow in US GLOBEC Ecosystems Using End-to-End Models BY JAMES J. RUZICKA, JOHN H. STEELE, SARAH K. GAICHAS, TOSCA BALLERINI, DIAN J. GIFFORD, RICHARD D. BRODEUR, AND EILEEN E. HOFMANN Krill and anchovies photos credit: NOAA-NWFSC, FE Division. Jellyfish photo credit: R. Brodeur. Albatross photo credit: J. Ruzicka. 82 Oceanography | Vol. 26, No. 4 ABSTRACT. End-to-end models were constructed to examine and compare the relationships within four US GLOBEC trophic structure and energy flow in coastal shelf ecosystems of four US Global Ocean ecosystems. An end-to-end model Ecosystem Dynamics (GLOBEC) study regions: the Northern California Current, describes the flow of energy (as biomass) the Central Gulf of Alaska, Georges Bank, and the Southwestern Antarctic Peninsula. through the ecosystem from the input High-quality data collected on system components and processes over the life of the of nutrients, through the production Analysis of Energy Flow in US GLOBEC Ecosystems program were used as input to the models. Although the US GLOBEC program was of plankton, fish, seabirds, mammals, species-centric, focused on the study of a selected set of target species of ecological and fisheries, to detritus and recycled or economic importance, we took a broader community-level approach to describe nutrients. Our primary goal is to identify Using End-to-End Models end-to-end energy flow, from nutrient input to fishery production. We built four end- the main attributes that regulate each to-end models that were structured similarly in terms of functional group composition system’s response to perturbations at and time scale. The models were used to identify the mid-trophic level groups that multiple trophic levels. We use the mod- place the greatest demand on lower trophic level production while providing the els to estimate the relative importance of greatest support to higher trophic level production. In general, euphausiids and the different functional groups as energy- planktivorous forage fishes were the critical energy-transfer nodes; however, some transfer nodes and to estimate the impact differences between ecosystems are apparent. For example, squid provide an important of changes at these nodes. In addition to alternative energy pathway to forage fish, moderating the effects of changes to forage understanding and comparing ecosystem fish abundance in scenario analyses in the Central Gulf of Alaska. In the Northern structure and dynamics, a major goal California Current, large scyphozoan jellyfish are important consumers of plankton of this study is to develop an end-to- production, but can divert energy from the rest of the food web when abundant. end model platform that can be applied broadly across diverse ecosystems. INTRODUCTION (e.g., Wiebe et al., 2003; Lough et al., The broad objective of the Global Ocean 2005). However, understanding tropho- Four US GLOBEC Ecosystems Ecosystem Dynamics (GLOBEC) pro- dynamic interactions among species There are striking differences among gram was to understand the processes has long been recognized as critical the GLOBEC ecosystems in bottom that control population variability. The to understanding the dynamics of depth and topography, circulation and GLOBEC approach was to study linkages the ecosystem as a whole (e.g., Frank stratification, seasonal cycles, and com- between the recruitment variability of et al., 2005). Multispecies ecosystem munity composition across all trophic target species (e.g., calanoid copepods, models of increasing sophistication are levels. These differences have prompted euphausiids, cod, haddock, salmon) and being developed to meet the need for a collection of different data sets and environmental processes operating across community-level approach to manage- application of different food web models broad temporal and spatial scales. The ment of marine resources and ecosystem for each ecosystem, making direct inability to conduct controlled experi- services subject to fishing pressures and end-to-end comparisons of energy ments is a major impediment to the climatic change (Travers et al., 2007; flow patterns challenging. scientific study of the mechanics of ocean Fogarty et al., 2013, in this issue). ecosystem dynamics. Ecosystem models Applying multispecies ecosystem Northern California Current provide the best proxy for controlled models within a comparative analysis of The Northern California Current (NCC; experiments (deYoung et al., 2010) and different ecosystems provides additional Figure 1a) is a highly productive sea- offer a way to study the integrated effects insight to ecosystem structure and func- sonal upwelling ecosystem (Huyer, of the critical processes that occur on dif- tion. Comparative studies can serve as 1983; Checkley and Barth, 2009). On ferent scales (Fogarty and Powell, 2002). proxies for controlled, manipulative short time scales, lower trophic level Species-centric models have proved studies but require that each ecosystem dynamics are strongly coupled to the to be valuable tools for studying the model be similarly structured in terms of timing, strength, and duration of upwell- effects of fishery management policies spatial and temporal scale and functional ing (Thomas and Strub, 2001; Thomas on individual fish stocks (Rothschild, group resolution. Here, we describe the and Brickley, 2006). On interannual to 1986) and the effects of ocean physics development and analysis of end-to-end interdecadal time scales, basin-scale on the dynamics of individual species ecosystem models of the trophodynamic climate processes (e.g., El Niño-Southern Oceanography | December 2013 83 b. Central Gulf of Alaska a. Northern California d. Southwestern Current Antarctic Peninsula 61°30'N 63°N 48°0'N 58°30'N 66°N 55°30'N 46°30'N 52°30'N 69°N 162°W 156°W 150°W 144°W 45°0'N c. Georges Bank 72°W 69°W 66°W 63°W 60°W 57°W 43°30'N Figure 1. US GLOBEC regions and end-to-end 43°30'N model domains (shaded in darker blue). The blue bathymetry lines mark 200 m and the black bathymetry lines 1,000 m. 42°0'N 42°0'N 126°0'W 124°30'W 123°0'W 40°30'N 70°30'W 69°0'W 67°30'W 66°0'W Oscillation, Pacific Decadal Oscillation model of the NCC must incorporate mammal populations have changed [PDO]) and interregional transport of both local physical processes (upwelling- dramatically over the past 40 years, large water masses strongly influence driven primary production) and impor- with some species shifts correlating well local ecosystem dynamics (Di Lorenzo tant nonlocal factors that affect commu- with the 1976–1977 PDO shift (Francis et al., 2013, in this issue), control the nity composition across all trophic levels. et al., 1998; Anderson and Piatt, 1999). composition of upwelling source waters Connecting these physical and lower (Huyer et al., 2002), and affect the com- Central Gulf of Alaska trophic level processes with what appear position of the local mesozooplankton The Central Gulf of Alaska (CGOA) to be strong shifts among mid and upper grazer community (Batchelder et al., system (Figure 1b) is a highly produc- trophic level interactions in this ecosys- 2002; Keister and Peterson, 2003). These tive downwelling system (Stabeno et al., tem (Gaichas et al., 2011) is an important physical and lower trophic level pro- 2004). Offshore surface waters that are challenge for end-to-end modeling. cesses directly affect the production of advected onto the shelf during down- pelagic fishes (Brodeur and Pearcy, 1992; welling events originate from the high- Georges Bank Ruzicka et al., 2011; Burke et al., 2013), nutrient, low-chlorophyll (HNLC), iron- Georges Bank (GB) is a shallow bank benthic invertebrates (Barth et al., 2007), limited region of the North Pacific gyre. offshore of Cape Cod (Figure 1c). It and local seabird and marine mammal Mixing of HNLC waters with iron-replete has long been the site of economi- populations (Ainley and Boekelheide, shelf waters drives the production cycle cally important fisheries, includ- 1990; Keiper et al., 2005). An end-to-end (Fiechter et al., 2009). Fish and marine ing cod (Gadus morhua), haddock 84 Oceanography | Vol. 26, No. 4 (Melanogrammus aeglefinus), flatfishes, forage throughout the greater Antarctic to transform a top-down linear expres- Atlantic lobster (Homarus americanus), Peninsula region (Catry et al., 2004; sion of predation pressure (Ecopath) into and scallops (Placopecten magellanicus). Croxall et al., 2005; Phillips et al., 2005; a bottom-up map of energy flow from Over GB’s shallow, central region, tur- Biuw et al., 2007). An end-to-end model lower trophic level producers to upper bulent tidal mixing is sufficiently strong of the sWAP ecosystem must incorporate trophic level consumers (Steele, 2009). to keep the water column well mixed important local physical processes and With the inclusion of external nutrient year-round. A permanent hydrographic must take into account intra-regional fluxes as input for uptake by phyto- front near the 60 m isobath separates the connectivity within the greater Antarctic plankton, nutrient recycling via bacterial central bank from stratified waters on Peninsula–Scotia Sea ecosystem.
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