Hydrographic Control of the Marine Ecosystem in the South Shetland-Elephant Island and Bransfield Strait Region
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ARTICLE IN PRESS Deep-Sea Research II 57 (2010) 519–542 Contents lists available at ScienceDirect Deep-Sea Research II journal homepage: www.elsevier.com/locate/dsr2 Hydrographic control of the marine ecosystem in the South Shetland-Elephant Island and Bransfield Strait region Valerie Loeb a,n, Eileen E. Hofmann b, John M. Klinck b, Osmund Holm-Hansen c a Moss Landing Marine Laboratories, 8272 Moss Landing Road, Moss Landing, CA 95039, USA b Old Dominion University, Norfolk, VA 23529, USA c Scripps Institution of Oceanography, La Jolla, CA 92093, USA article info abstract Article history: The South Shetland-Elephant Island and Bransfield Strait region of the West Antarctic Peninsula is an Accepted 30 October 2009 important spawning and nursery ground of Antarctic krill (Euphausia superba) and is an important Available online 11 November 2009 source of krill to the Southern Ocean. Krill reproductive and recruitment success, hence supply of krill to predator populations locally and in downstream areas, are extremely variable on interannual and Topical issue on "Krill Biology and Ecology." longer time scales. Interannual ecosystem variability in this region has long been recognized and The issue is compiled and guest-edited by the North Pacific Marine Science thought related to El Nin˜o Southern Oscillation (ENSO) events, but understanding of how has been Organization (PICES), International Council limited by the hydrographic complexity of the region and lack of appropriate ocean-atmosphere for the Exploration of the Sea (ICES), and interaction models. Global Ocean Ecosystem Dynamics This study utilizes multidisciplinary data sets collected in the region from 1990 to 2004 by the U.S. (GLOBEC) project. Antarctic Living Marine Resources (AMLR) Program. We focus on hydrographic conditions associated with changes in the distribution, abundance and composition of salp- and copepod-dominated Keywords: zooplankton assemblages during 1998 and 1999, years characterized respectively by a strong El Nin˜o Southern Ocean event and La Nin˜a conditions. We provide detailed analyses of hydrographic, biological and ecological Euphausia superba conditions during these dichotomous years in order to identify previously elusive oceanographic Climate regime shifts processes underlying ecosystem variability. We found that fluctuations between salp-dominated Atmospheric-oceanic coupled processes Antarctic Dipole coastal zooplankton assemblages and copepod-dominated oceanic zooplankton assemblages result Antarctic Circumpolar Current from the relative influence of Weddell Sea and oceanic waters and that these fluctuations are associated with latitudinal movement of the Southern Antarctic Circumpolar Current Front (sACCf). Latitudinal movements of the sACCf can be explained by meridional atmosphere teleconnections instigated in the western tropical Pacific Ocean by ENSO variability and are consistent with out-of-phase forcing in the South Pacific and South Atlantic Oceans by the Antarctic Dipole high-latitude climate mode. During El Nin˜o decreased northwest winds, equatorward movement of the sACCf and an intensified Weddell Gyre allow Weddell Sea water to flow into eastern Bransfield Strait. During these periods mixing between oceanic and coastal waters is reduced, chlorophyll a concentrations are low, salps numerically dominate the zooplankton, and krill recruitment success is poor. During La Nin˜a increased and more frequent northwest winds and poleward movement of the sACCf allows increased influence of oceanic waters and mixing of these with cold coastal waters. These periods are characterized by numerical dominance of copepods, elevated concentrations of oceanic zooplankton taxa and phytoplankton blooms that promote krill reproduction and recruitment success. Hydrographic and ecological changes after the 1998 El Nin˜o are associated with a shift from frequent El Nin˜os to the prevalence of La Nin˜a and neutral conditions and conform to a decadal-scale climate regime shift in the Antarctic Peninsula region. & 2009 Elsevier Ltd. All rights reserved. 1. Introduction 2004) and therefore plays a vital role in the krill-based food web. In particular the South Shetland-Elephant Island and Bransfield The Southwest Atlantic, including the West Antarctic Peninsu- Strait region (Fig. 2) is an important krill spawning and nursery la (Fig. 1), is an important source of Antarctic krill (Euphausia ground as well as an important area for the commercial krill superba) to the Southern Ocean (Spiridonov, 1996; Atkinson et al., fishery (Siegel, 2005). Krill reproductive success, population size and supply to dependent predator populations here and in downstream areas are highly variable on interannual and longer n Corresponding author. Fax: +831 632 4403. time scales (Priddle et al., 1988; Siegel and Loeb, 1995; Loeb et al., E-mail address: [email protected] (V. Loeb). 1997). Understanding of factors driving krill population variability 0967-0645/$ - see front matter & 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.dsr2.2009.10.004 ARTICLE IN PRESS 520 V. Loeb et al. / Deep-Sea Research II 57 (2010) 519–542 Salpa thompsoni (Siegel and Loeb,1995; Loeb et al., 1997; Atkinson et al., 2004) and therefore influences two important components of the nekton/zooplankton sampled during summer months. Zooplankton in the South Shetland-Elephant Island region includes mixtures of species characteristic of coastal and oceanic environments that show significant seasonal and interannual fluctuations in total and relative abundance of dominant taxa ( Siegel and Piatkowski, 1990; Park and Wormuth, 1993; Schnack- Schiel and Mujica, 1994). Notable among these are marked interannual shifts between an abundant copepod-dominated zooplankton assemblage characteristic of warm oceanic water in Drake Passage (the ‘‘West Wind Drift’’) and a less-abundant salp- dominated assemblage characteristic of cold neritic waters (the ‘‘East Wind Drift’’; Jaz˙ dz˙ ewski et al., 1982; Witek et al., 1985; Schnack-Schiel and Mujica, 1994). During surveys associated with the Biological Investigations of Marine Antarctic Systems and Stocks (BIOMASS) Program in the early 1980s, variable represen- tation by the two assemblages was associated with latitudinal shifts in their distributions. These in turn were attributed to interannual variations in atmospheric, climatic and hydrographic conditions (Jaz˙ dz˙ ewski et al., 1982; Witek et al., 1985). As with Fig. 1. Major regions of high krill concentrations (black areas) in the Southern sea-ice extent, ENSO was believed to be a major factor driving Ocean relative to general hydrographic circulation (arrows), the Polar Front (PF), these variations (Witek et al., 1985; Priddle et al., 1988). However, Southern Antarctic Circumpolar Front (sACCf, long dashes) and Boundary (Bndy, short dashes) modified from Spiridonov (1996). establishing the linkage between ENSO and ecosystem variability in the region since that time has been severely limited by (a) the inability to identify specific hydrographic conditions associated with distribution shifts of the oceanic and coastal zooplankton has been limited due to the variable distribution and abundance assemblages and (b) lack of appropriate ocean-atmosphere patterns of relatively long-lived krill across multiple spatial and interaction models that can explain the interrelationships. temporal scales (Siegel, 2005) compounded by environmental The primary objectives of this work are to: complexity of their source regions (Fach et al., 2006) and lack of sufficiently long-term, internally consistent, multidisciplinary identify specific hydrographic conditions associated with data sets. variable representation of oceanic and coastal zooplankton The focus of the current study is the South Shetland-Elephant assemblages in the South Shetland-Elephant Island region ; Island region and adjacent Bransfield Strait and Drake Passage establish how these linked hydrographic-biological conditions (Fig. 1, Fig. 2A,B) that is annually surveyed by the U.S. Antarctic are related to the two opposite extreme ENSO phases, the Marine Living Resources (AMLR) Program. The hydrography and ‘‘warm’’ El Nin˜o and ‘‘cool’’ La Nin˜a; and circulation of this region is complex and variable (Amos, 1984, put these linked hydrographic-biological conditions in the 2001; Priddle et al., 1994). The southern Antarctic Circumpolar context of basin-scale coupled atmosphere-oceanic processes Current front (sACCf) and southern ACC boundary (Bndy) are manifested by ENSO variability extending over decadal time present in southern Drake Passage (Orsi et al., 1995). The western scales. portion of the Weddell Sea gyre influences circulation in the Bransfield Strait and Elephant Island region (Gordon and Nowlin, 1978; Whitworth et al., 1994) and inputs from upstream regions To achieve these objectives we employ the long-term data sets along the western Antarctic Peninsula enter the region through obtained by the US AMLR Program. Gerlache Strait and western Bransfield Strait (Stein, 1986, 1988, The AMLR Program has maintained a field effort in the South 1989; Niiler et al., 1991; Capella et al., 1992; Garcia et al., 1994; Shetland-Elephant Island region since the late 1980s, and since Hofmann et al., 1996). The rugged bathymetry of the region, 1993 has focused on a relatively fixed grid of sampling stations which includes the continental shelf around the islands, deep (Fig. 2B) across which hydrographic, chlorophyll a, krill and basins of Bransfield Strait, and the South Shetland Trench