A Comparison of Nutrient Characteristics and Primary Productivity
in the Sulu Sea and South China Sea1
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Maria Lourdes, San Diego-McGlone2, Gil S. Jacinto2, Vilma C. Dupra2, Ingrid S. Narcise2, Daisy O. Padayao2, and Imelda B. Velasquez2

(received 1998/9/24, revised 1998/10/27, accepted 1998/11/5)
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ABSTRACT

A comparison of vertical nutrient characteristics in the South China Sea (SCS) and Sulu Sea showed the typical nutrient depletion in surface water and enrichment with depth with greater maximum apparent in the SCS relative to the uniform behavior of nutrients below 400 m in the Sulu Sea. This may be explained by sill inflow of SCS intermediate waters that supplies intermediate and bottom waters of the Sulu Sea. Shallowing of the nutricline on the shelf may be due to geostrophic adjustment or may be pulses of upwelled water associated with shoaling of internal waves. The deep chlorophyll maximum was found above the nutricline at all stations and coincided with the maximum dissolved oxygen concentration. The depth-integrated primary production rates in the oceanic region of the Sulu Sea (195¡Ó 2 gC¡Ñ m-2¡Ñ yr-1) was determined to be the highest followed by the shoal area (166¡Ó 3 gC¡Ñ m-2¡Ñ yr-1) and the oceanic region in the SCS (147¡Ó 3 gC¡Ñ m-2¡Ñ yr-1). Plankton biomass estimates integrated with depth showed similar trends for both basins.

(Keywords: South China Sea, Sulu Sea, nutrients, primary productivity, deep chlorophyll maximum)

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1 Contribution no. 291, Marine Science Institute, University of the Philippines.
2 Marine Science Institute, University of the Philippines, Diliman, Quezon City 1101, Philippines.

INTRODUCTION
The South China Sea (SCS) is the largest marginal basin in the Asiatic Mediterranean Sea with its central part exceeding 4000 m in depth (Gong et al., 1992). The Sulu Sea is an adjacent smaller marginal basin that is interconnected with the SCS by a topographic sill (Mindoro Strait). Studies have shown that tidal flow over a sill (Pearl Bank) can be influenced by the sharp topographic slope resulting in the generation of internal waves (e.g., Apel et al., 1985; Liu et al., 1985). Shoaling of these internal waves may in turn induce upwelling events. It has also been reported that the inflow of the North Pacific Intermediate Water (from the SCS) across the sill (Mindoro Strait) is the source for intermediate and deep waters of the Sulu Sea (Frische and Quadfasel, 1990). Such hydrodynamic features could influence the behavior of chemical constituents (e.g., nutrients) and biota (e.g., plankton) in these marginal basins.

Oceanographic studies were conducted to describe gross features and variability within and across the SCS and Sulu Sea. Of particular interest are nutrient characteristics and productivity levels in representative shelf, shoal, and oceanic regimes in the basins. These features are important in inferring possible interactions between the shelf reef systems of western Philippines (e.g., Palawan and Sulu Sea) and the SCS. This could have implications on the recruitment of propagules advected from the centrally located shoal reefs in the SCS to the Philippine shelf reef system and the transport of materials (e.g., nutrients) across hydrographic regimes.
 

MATERIALS AND METHODS
Water sampling and productivity measurements in the Sulu Sea and SCS were conducted on board the RPS Explorer from 9-22 May 1998. A total of 14 hydrographic stations (Fig. 1) were occupied, with 6 stations in the Sulu Sea and 8 in the SCS. Representative shelf, shoal, and oceanic areas in the two basins were identified. For this study, the shelf is defined as the zone around an island extending from the low water line to less than 200 m depth. The shoal is a submerged ridge or bank, and oceanic refers to greater than 200 m depth. Stations 1, 2, 3, 4, and 6 are the oceanic stations in the Sulu Sea, while Stn. 5 is a shelf station. In the SCS, Stns. 9, 10, 11, 12, and 13 are the oceanic stations, Stns. 7 and 8 are the shelf stations, and Stn. 14 is a shoal station (Reed Bank). Samples were taken at 12 sampling depths using 1.7-L and 5-L Niskin samplers (General Oceanics Inc.) mounted on a SEABIRD Rosette equipped with an SBE-19-CTD profiler and an oxygen sensor. Sampling was limited to the top 1000 m of the water column. The standard water sampling depths were 5, 25, 50, 75, 100, 150, 200, 300, 400, 500, 600, and 700 m.

Samples collected were analyzed for dissolved oxygen, nutrients, and chlorophyll-a. Dissolved oxygen measurements were done on board the ship using the Winkler titration method (Parsons et al., 1984). Samples for nutrient determination were stored frozen in Nalgene HDPE (high density polyethylene) bottles and later analyzed spectrophotometrically (Spectronic Genesys, Milton Roy) following the methods described by Parsons et al. (1984), and using a nutrient autoanalyzer (San Plus System, Skalar). Samples for chlorophyll-a analysis were filtered through Whatman membrane filters (cellulose nitrate, 0.45£gm). The filters were then frozen and later analyzed following the method given by Parsons et al. (1984).
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Fig. 1. Map with sampling stations.

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Phytoplankton production was determined using the 24-h 14C incubation procedure modified from the JGOFS Protocol (1994). A set of four depths from the surface to below the subsurface chlorophyll maximum was selected at each productivity station. The depths were 5, 30, 70, and 90 m and the productivity stations were Stns. 2, 7, 11, and 14 (Fig. 1). Samples were obtained at dawn using 5-L Niskin samplers. BOD bottles (300 ml) were filled directly from the Niskin samplers and 3 bottles were used for each sample depth. Under low light conditions, 1 ml of 5 £gCi Na214CO3 was added to each bottle. The bottles were then incubated on board the ship using nylon screens to simulate the light levels in the water column where the samples were collected. The light intensities mimicked were 45, 25, 17, and 9% of incident PAR. Incubation was from dawn of the first day to dawn of the following day inside a plastic bin (approx. 1x1x0.5 m3) filled with running seawater. After incubation, the contents of the bottles were passed through GF/C filters and the filters stored inside the freezer until further analysis in the laboratory. The radioactivity of 14C taken in by the plankton was measured using a liquid scintillation counter (Deckman LS 6500).

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RESULTS AND DISCUSSION
The nutrient (NO3, PO4, and SiO4) profiles for both the Sulu Sea and the SCS showed a characteristic increase with depth reflecting surface water depletion from biological uptake and deeper water enrichment from regeneration processes (Figs. 2a-2c). The vertical range in concentrations for stations in the Sulu Sea were 0-2.15 £gM for PO4, 0-28.4 £gM for NO3, and 0.36-56.3 £gM for SiO4. In the SCS, the vertical range in concentrations was 0-2.59 £gM for PO4, 0-34.2 £gM for NO3, and 0.73-93.0 £gM for SiO4. These results showed a continuous increase in concentration with depth in the SCS producing larger nutrient maxima and the relatively unchanging concentrations of nutrients below 400 m in the Sulu Sea. The uniform behavior of water signatures (temperature, salinity, nutrients) in the Sulu Sea may be attributed to the presence of a 400 m deep sill that separates the marginal basins. While the near-surface circulation is mainly governed by the seasonally reversing monsoon winds, the deep circulation in the Sulu Sea is forced by an inflow of intermediate water from the SCS through the Mindoro Strait that supplies its intermediate and bottom waters (Wyrtki, 1961). The erosion of vertical nutrient gradients below sill depth is associated with the tidally induced vertical mixing in the Mindoro Strait that transforms deeper water masses in the Sulu Sea by turbulent mixing of over- and underlying waters (Frische and Quadfasel, 1990; Villanoy and Udarbe, 1995).

Among the Sulu Sea stations, a shallowing of the nutricline by about 10-20 m was observed when moving from the oceanic (Stn. 1) to the shelf stations (Stn. 4) near Palawan (Figs. 3a-3c). This may be due to geostrophic adjustments as water moves to shallower shelf areas. It may also be attributed to the well-documented fortnightly occurrence of large-scale (5-16 km wavelengths) oceanic internal waves generated in the southeastern Sulu Sea by intense tidal flow over the Pearl Bank sill (e.g., Apel et al., 1985; Liu et al., 1985). It has been hypothesized that these internal waves could induce upwelling pulses resulting in higher nutrient concentrations at shallower depths (e.g., Pingree et al., 1981; Pingree et al., 1986; New, 1988). The shallowing of the nutricline translated to a shallowing of the chlorophyll maximum (Fig. 3d).

Vertical profiles of chlorophyll-a showed the presence of a deep chlorophyll maximum (DCM) occurring between 50-80 m in the Sulu Sea and between 40-75 m in the SCS (Fig. 2d). Although the depth of the chlorophyll maximum was similar for both basins, the maximum chlorophyll-a concentration in the Sulu Sea was higher at 1.11 £gg¡Ñ L-1 than the SCS (0.66 £gg¡Ñ L-1). In what Herbland and Voituriez (1979) refers to as the ¡§Typical Tropical Structure¡¨, the DCM is found near the nutricline and thermocline, and the peak in the primary production is coincident with or just above the chlorophyll maximum (Cullen, 1982). In this study the DCM was found above the nutricline (Fig. 4). This implies that DCM represents not only a physiological adaptation to lower irradiance but also to greater availability of nutrients (e.g., Banse, 1987; Cullen, 1982). The plankton may be utilizing the nutrients diffusing upward from the deeper parts of the pycnocline and thus trap this flux (e.g., Jamart et al., 1977). Appreciable biomass accumulation will only take place when sustained new production is possible (Harris, 1986). It was also determined that the maximum dissolved oxygen (DO) concentration was found at depths above or coinciding with the DCM in both the Sulu Sea and SCS (Fig. 4). Calculations showed that DO concentrations at these depths were oversaturated by as much as 10%. This value may represent part of the DO released from increased plankton production at the DCM. Similar results were reported by Huang (1992).
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Fig. 3¡@ of the nutricline and chlorophyll maxima from oceanic (Stn. 1) to closer to shelf areas (Stn.4) in the Sulu Sea.

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  Fig. 4 PO4 and DO profiles vis a vis the deep (subsurface) chlorophyll maxima for the Sulu Sea and South China Sea. ¡@
Primary production in the shelf, shoal, and oceanic areas of the Sulu Sea and SCS were estimated using various light conditions. The light regime chosen was based on vertical light readings and these were 47%, 25%, 17%, and 9% of incident PAR. A comparison of the depth integrated primary production on the top 30 m of the water column showed that rates in the oceanic part of the Sulu Sea (130¡Ó 2 gC¡Ñ m-2¡Ñ yr-1) were higher than that in the SCS (105¡Ó 1 gC¡Ñ m-2¡Ñ yr-1) (Table 1). Within the SCS, primary production in the shelf (147¡Ó 2 gC¡Ñ m-2¡Ñ yr-1) was highest followed by the oceanic region (105¡Ó 2 gC¡Ñ m-2¡Ñ yr-1) and lowest in the shoals (50¡Ó 3 gC¡Ñ m-2¡Ñ yr-1). The depth integrated (0-30 m) chlorophyll values followed similar trends with a high value of 9.2 mg¡Ñ m-2 Chl a in the Sulu Sea and a low value of 4.2 mg¡Ñ m-2 Chl a in the shoals of the SCS. The DCM was located within the top 80 m of the water column for both basins. When integrated over this depth (top 70 m), the productivity in the oceanic region of the Sulu Sea was still the highest at 195¡Ó 2 gC¡Ñ m-2¡Ñ yr-1 followed by the shoals of the SCS (166¡Ó 3 gC¡Ñ m-2¡Ñ yr-1) and then the oceanic region in the SCS (147¡Ó 3 gC¡Ñ m-2¡Ñ yr-1). The depth integrated chlorophyll value for this depth range was 24 mg¡Ñ m-2 Chl a in the Sulu Sea and 21 mg¡Ñ m2 Chl a in the SCS. These estimates compare well with the 152¡Ó 32 gC¡Ñ m-2¡Ñ yr-1 productivity estimate and the 17.8¡Ó 3.8 mg¡Ñ m-2 Chl a estimate reported by Huang (1988) for the Nansha Islands (Spratlys) (Table 2). They are also higher than Equatorial Pacific rates but lower than rates in coastal upwelling areas such as Guangdong (Table 2). Based on depth-integrated estimates, primary

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Table 1. Depth integrated primary production and chlorophyll estimates.
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Production
(gC¡P m-2¡P yr-1)
Chl-a
(mg¡P m-2)
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5-30 m depth range ¡@ ¡@ ¡@
Stn 2- Sulu oceanic
130 ¡Ó 2 (n=3)
5.2
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Stn 7- SCS shelf
147 ¡Ó 2 (n=3)
9.2
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Stn 11- SCS oceanic
105 ¡Ó 2 (n=3)
5.7
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Stn 14- SCS shoal
50 ¡Ó 3 (n=3)
4.2
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5-70 m depth range ¡@ ¡@ % production below 30 m
Stn 2- Sulu oceanic
195 ¡Ó 2 (n=3)
24
33%
Stn 11- SCS oceanic
147 ¡Ó 3 (n=3)
21
29%
Stn 14- SCS shoal
166 ¡Ó 3 (n=3)
-
70%
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Table 2. Comparison of depth integrated production and chlorophyll¡@
estimates from different sites.
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Sites
Production
(gC¡P m-2¡P yr-1)
Chl-a
(mg¡P m-2)
Sulu Sea¡@¡@¡@¡@
(oceanic, this study)
195 ¡Ó 2
24
South China Sea¡@¡@¡@¡@
(oceanic, this study)
147 ¡Ó 3
21
South China Sea¡@¡@¡@¡@
(shoal, this study)
166 ¡Ó 3
-
Nansha Islands¡@¡@¡@¡@
(Huang, 1988)
152 ¡Ó 32
17.8 ¡Ó 3.8
Western Equatorial Pacific¡@¡@¡@¡@
(Mackey et al., 1995)
35-88
23-32
Hainan¡@¡@¡@¡@
(Han et al., 1990)
889
-
Guangdong¡@¡@¡@¡@
(Han and Ma, 1988)
569
-
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production between depths of 30 and 70 m contributed 30-70% of water column production (Table 1). Higher production in the Sulu Sea may be attributed to the physical transport processes (internal wave induced upwelling and turbulent mixing) that provide the euphotic zone of the Sulu Sea with nutrients from the more enriched lower depths.

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CONCLUSION
Nutrient characteristics in marginal basins are influenced by sill induced hydrodynamic processes. Variability in nutrients for two marginal basins separated by a sill could, in turn, result in differences in production and plankton biomass estimates. Oceanographic studies done along a transect that includes oceanic, shelf, and shoal regions of the SCS and Sulu Sea showed the role of physical transport processes on the chemistry and biology in the basins.
acknowledgment
The authors gratefully acknowledge the support given by the Department of Science and Technology and the Mines and Geosciences Bureau (Department of Environment and Natural Resources) for this project.



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