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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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.
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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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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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
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| Stn 2- Sulu oceanic |
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| Stn 7- SCS shelf |
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| Stn 11- SCS oceanic |
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| Stn 14- SCS shoal |
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| 5-70 m depth range | ¡@ | ¡@ | % production below 30 m |
| Stn 2- Sulu oceanic |
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| Stn 11- SCS oceanic |
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| Stn 14- SCS shoal |
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(oceanic, this study) |
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(oceanic, this study) |
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(shoal, this study) |
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(Huang, 1988) |
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(Mackey et al., 1995) |
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(Han et al., 1990) |
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(Han and Ma, 1988) |
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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.
Apel, J.R., J.R. Holbrook, A.K. Liu, and J.J. Tsai (1985) The Sulu Sea internal soliton experiment. J. Phys. Oceanogr. 15, 1625-1651.
Banse, K. (1987) Clouds, deep chlorophyll maxima and the nutrient supply to the mixed layer of stratified water bodies. J. Plankton Res. 9, 1031-1036.
Cullen, J.J. (1982) The deep chlorophyll maximum: comparing vertical profiles of chlorophyll-a. Can. J. Fish. Aquat. Sci. 39, 791-803.
Frische, A. and D. Quadfasel (1990) Hydrography of the Sulu Sea. Proc. of the Ocean Drilling Program, Initial Reports, edited by C. Rangin, E. Silver, and M.T. Breymann. 124, 101-103.
Gong, G.C., K.K. Liu, C.T. Liu, and S.C. Pai (1992) The chemical hydrography of the South China Sea West of Luzon and a comparison with the West Philippine Sea. Terr. Atmos. Ocean. Sci. 3, 587-602.
Han, W., M. Wang. and K. Ma (1990) On the lowest surface water temperature area of China Sea in summer ¡V the upwelling along the east coast of Hainan Island. Oceanol. Limnol. Sin. 21, 267-275.
Han, W. and K. Ma (1988) Study on the upwelling along the east coast of Guangdong. Acta Oceanol. Sin. 7, 552-561.
Harris, G.P. (1986) Phytoplankton Ecology ¡V Structure, Function, and Fluctuation. Chapman and Hall Ltd.
Herbland, A. and B. Voituriez (1979) Hydrological structure analysis for estimating the primary production in the tropical Atlantic Ocean. J. Mar. Res. 37, 87-101.
Huang, L. (1992) Vertical variations of chlorophyll a and fluorescence values of different areas in the South China Sea. Trop. Oceanol. Redai. Haiyang. 11, 89-95.
Huang, L. (1988) Distribution characteristics of chlorophyll a and estimation of primary productivity in the waters around Nansha Islands [Abstract]. Proc. Marine Biology of the South China Sea, edited by X. Li and B. Morton. 261-276.
Jamart, B., D.F. Winter, K. Banse, G.C. Anderson, and R.K. Lam (1977) A theoretical study of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern U.S. coast. Deep Sea Res. 24, 753-773.
Joint Global Ocean Flux Study (JGOFS) Protocol. June 1994. (http://ads.smr.uib.no/jgofs/jgofs.htm)
Liu, A.K., J.R. Holbrook, J.R. Apel (1985) Nonlinear wave evolution in the Sulu Sea. J. Phys. Oceanogr. 15, 1613-1624.
Mackey, D.J., J. Parslow, H.W. Higgins, and F.B. Griffiths (1995) Plankton productivity and biomass in the western Equatorial Pacific: Biological and physical controls. Deep Sea Res. 42, 499-533.
New, A.L. (1988) Internal tidal mixing in the Bay of Biscay. Deep Sea Res. 35, 691-697.
Parsons, T.R., Y. Maita, and C.M. Lalli (1984) A Manual of Chemical and Biological Methods for Seawater Analysis. Pergamon Press.
Pingree, R.D. and G.T. Mardell (1981) Slope turbulence, internal waves and phytoplankton growth at the Celtic Sea shelf-break. Phil. Trans. Roy. Soc. Lond. A 3012, 663-682.
Pingree, R.D., G.T. Mardell, and A.L. New (1986) Propagation of the internal tides from the upper slopes of the Bay of Biscay. Nature 321, 154-158.
Villanoy, C.L. and M.J.B. Udarbe (1995) The circulation in the Eastern South China Sea during May 1993. The Philippine Scientist: Special Issue. Proc. 3rd National Symposium in Marine Science, edited by F.B. Sotto, J. Young, and J. Baumgartner. 128-142.
Wyrtki, K. (1961) Scientific results of marine investigations of the South China Sea and the Gulf of Thailand, 1959-1961. NAGA Report 2. La Jolla, CA (Scripps Institution of Oceanography).