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The Unique Continental Shelf Dynamics Off Western Australia: Physical Controls on Phytoplankton Productivity

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The Unique Continental Shelf Dynamics Off Western Australia: Physical Controls on Phytoplankton Productivity NOT TO BE CITED WITHOUT PRIOR REFERENCE TO THE AUTHORS International Council for the Exploration of the Sea Annual Science Conference 2003 Theme Session P: Physical-Biological Interactions in Marginal and Shelf Seas CM 2003/P:16 THE UNIQUE CONTINENTAL SHELF DYNAMICS OFF WESTERN AUSTRALIA: PHYSICAL CONTROLS ON PHYTOPLANKTON PRODUCTIVITY C.E. Hanson*, C.B. Pattiaratchi and A.M. Waite Centre for Water Research, The University of Western Australia, 35 Stirling Hwy, Crawley, Western Australia 6009 Australia *Corresponding author Email: [email protected] Phone: +618 9380 1698 Fax: +618 9380 1015 Abstract Continental shelf waters off Western Australia are dominated by the Leeuwin Current, an anomalous eastern boundary current that transports tropical water poleward and generates large- scale downwelling. However, in summer the inner shelf dynamics are influenced by wind- generated countercurrents (the Ningaloo and Capes Currents) that flow towards the equator and have the potential to create localized upwelling. Prior to the current study, there had been no investigations into the links between physical dynamics and phytoplankton productivity within or between any of these currents. In a series of cross-shelf transects encompassing the Leeuwin, Ningaloo and Capes Currents, we found strong associations between the nitracline and phytoplankton biomass that lead to persistent deep chlorophyll maxima throughout the study area. The countercurrents were highly productive (~ 800-1300 mg C m-2 d-1), in notable contrast to the low productivity signature (< 200 mg C m-2 d-1) within the boundary current. Seaward offshoots and coastal upwelling associated with the Ningaloo and Capes Currents provided a mechanism to access high nutrient concentrations normally confined to the base of the Leeuwin. This may be an important process leading to seasonal peaks in shelf productivity on this otherwise oligotrophic coast. Keywords: primary production; coastal upwelling; oligotrophic; deep chlorophyll maximum; eastern Indian Ocean; Leeuwin Current; Ningaloo Current; Western Australia, Gascoyne, 21°S to 30°S and 111°E to 115°E 1 1.0 Introduction Eastern boundary currents are present in all of the major ocean basins, and generally consist of an equatorward surface flow accompanied by large-scale upwelling, high rates of primary production and abundant fisheries (Wooster and Reid, 1963; Mann and Lazier, 1996). Off the coast of Western Australia (WA), the unusual poleward-flowing Leeuwin Current (LC) restricts the eastern arm of the Indian Ocean gyre to offshore regions, generating large-scale downwelling as it travels along the continental shelf break (Pearce, 1991). The LC is known to reduce coastal nutrient levels (Johannes et al., 1994), influence biological species distributions (Morgan and Wells, 1991) and limit levels of secondary productivity (Lenanton et al., 1991; Caputi et al., 1996). As opposed to the dominance of pelagic finfish stocks in other eastern boundary regions, the major fishery off Western Australia (WA) is benthic rock lobster, with its life cycle and recruitment strongly tied to the dynamics of the LC (Phillips et al., 1991). Inshore of the Leeuwin Current, a system of equatorward coastal countercurrents are driven by southerly winds which prevail during the austral summer months (December to March). Part of that system, the Capes Current (CC) extends along the southwest coast of WA (Pearce and Pattiaratchi, 1999) and the Ningaloo Current (NC) extends along the northwest coast (Taylor and Pearce, 1999). The combination of these wind-forced shelf currents and localized Ekman-driven upwelling (Gersbach et al., 1999; Woo and Pattiaratchi, 2003) seasonally restricts the inshore extent of the LC (Pearce and Pattiaratchi, 1999). The Gascoyne continental shelf extends from North West Cape (21.3°S) to Shark Bay (26.5°S), Western Australia (WA), and encompasses the northern portion of LC waters (Pearce, 1991). Control of phytoplankton production in coastal regions such as the Gascoyne is often closely tied to ambient nutrient levels, which in turn are strongly influenced by the local oceanography (Mann and Lazier, 1996). Along the west coast of WA, oligotrophic conditions in the nutrient-poor LC are thought to support only a limited amount of phytoplankton biomass and productivity. Phytoplankton biomass data in the Gascoyne region is quite sparse, however a review of field studies throughout WA characterized coastal waters as low chlorophyll (< 1 mg chl a m-3) environments, with higher chlorophyll a values considered representative only of shelf waters or estuaries subjected to anthropogenic nutrient inputs (Pearce et al., 2000). The only available data indicate that primary productivity off Western Australia is 150-250 mg C m-2 d-1 (Koblentz-Mishke et al., 1970; FAO, 1981), but these data were sourced from the International Indian Ocean Expedition (Kabanova, 1968), the Australian portion of which only sampled open ocean waters along 110°E (~ 400 km offshore; Jitts, 1969). To date, we are lacking both regional estimates of phytoplankton biomass and productivity in the nearshore and coastal Gascoyne region, and an understanding of the intermediate links between the physical environment and secondary productivity. The aim of the present paper is to understand how the Leeuwin, Ningaloo and Capes Currents impact water column production along the Gascoyne shelf. The oligotrophic LC is known to dominate the region, and the commonly accepted paradigm is that conditions remain oligotrophic along this coast through suppression of upwelling-driven production. We tested this hypothesis using field data from the northern portion of the LC and associated coastal countercurrents. 2.0 Materials and Methods 2.1 Study area An oceanographic cruise was undertaken off Western Australia from 13 to 27 November 2000 (early austral summer) aboard the R.V. Franklin (FR10/00), incorporating eleven onshore/offshore transects and a total of 118 stations (Fig. 1). The study region was located between the Abrolhos 2 Islands and North West Cape (21°S to 30 °S; Fig. 1), and encompassed the entirety of the Gascoyne continental shelf (0 – 200 m isobath), shelf break (200 – 300 m isobath) and offshore (300 – 4000 m isobath) waters. 2.2 Oceanographic sampling and laboratory analyses Water samples were obtained using 24 General Oceanics 5 L Niskin bottles mounted on a rosette equipped with Seabird CTD, dissolved oxygen sensor, fluorometer and Li-Cor LI-192SA underwater quantum sensor. Between three and thirteen discrete depths were sampled at each station (dependent on bottom depth), including surface (roughly 2 m), and above, below and within the fluorescence maximum (as determined by the downcast fluorometer trace). Dissolved inorganic nutrients (nitrate + nitrite, phosphate and silicate) were analyzed for all depths using a shipboard Autoanalyzer. Detection limits were 0.1 µM for nitrate + nitrite (hereafter nitrate) and 0.01 µM for phosphate. Two litre water samples were filtered for chlorophyll (chl) a and pheopigments onto Whatman GF/F filters, stored at -20°C and returned to the laboratory for analysis. Pigments were extracted in 90% acetone with grinding, and measured using a Turner Designs Fluorometer following the acidification technique of Parsons et al. (1989). At 18 ‘production stations’, primary productivity versus irradiance (P vs. I) experiments were performed using the small-volume, short-incubation-time 14C incorporation technique (Lewis and Smith, 1983), with photosynthetron equipment and modifications as per Mackey et al. (1995; 1997). Due to the opportunistic nature of the biological sampling program, water from some stations was collected at night and held in the dark at ambient seawater temperature until processing at dawn the following morning. For the first six experiments, irradiance levels within the photosynthetron (maximum of 400 µE m-2 s-1) were insufficient to reach the maximum photosynthetic rate for shallow-water (< 50 m) samples (UNSAT). From the seventh production station onwards, additional incubations of the surface and next-deepest sample were conducted in natural sunlight (SAT) at two light levels (100% and 30% of incident irradiance). All samples were counted on-board the ship using a LKB Rackbeta liquid scintillation counter. Additional water samples were collected at the surface and DCM of production stations for particulate organic carbon (POC) and particulate nitrogen (PN) analysis (4 L filtered on pre- combusted Whatman GF/F filters and stored at -20°C until analysis by mass spectrometer, following the preparation techniques of Knap et al. (1996). 2.3 Data processing and production calculations Attenuation coefficients (Kd) were calculated from a linear regression of the natural log of PAR vs. depth, according to the relation: ln Ed(0) = -Kdz + ln Ed(z) where Ed(0) and Ed(z) are the values of downwelling PAR at the surface and at z m, respectively (Kirk, 1994). A smoothed irradiance profile was then plotted and used to determine the depth of the euphotic zone, as defined by the depth of the 0.1% light level. In-situ fluorescence was calibrated with extracted chl a data using linear regression, and used as a proxy for phytoplankton biomass. Where data permitted (minimum of 5 data points), a separate regression was performed for each station; otherwise, stations were calibrated using pooled data for that transect (r2 = 0.76-0.92). The deep chlorophyll maximum (DCM) was taken as the depth of maximum subsurface chl a concentration. The top of the nitracline was defined as the depth where the nitrate concentration equaled 0.2 µM, as linearly interpolated between Niskin sampling depths. In these extremely oligotrophic waters, this value was considered more 3 representative of the nitracline than the criteria of 1.0 µM commonly used for other regions (e.g. Maranon and Holligan, 1999; Moran and Estrada, 2001). For the purposes of estimating integrated production, Pm for the UNSAT samples was taken as the maximum photosynthetic rate achieved in the photosynthetron (at ~ 400 µE m-2 s-1), providing a conservative estimate of this parameter. UNSAT samples were also necessarily without a measure of the photoinhibition parameter (β).
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