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Abstract
Seasonal changes of uptake of nitrogenous nutrients (NH14 , NO23 , NO22 , and urea) and regeneration (NH14 and
NO 22 production) in the plankton fraction of a mangrove ecosystem on the west coast of India were investigated.
Nitrate was the major fraction of assimilable N (72%), followed by NH14 (16%), NO22 (6%), and urea (6%). Changes
of nutrient concentrations followed clear seasonal cycles and were mainly regulated by in situ biological processes.
The plankton took up NO 23 and NH 14 in more or less equal proportions (39 and 44% respectively), followed by
urea (11%) and NO 22 (6%). Seasonal patterns of uptake were distinct, with a dominance of NO 23 and NO 22 uptake
in the postmonsoon, followed by a dominance of NH14 and urea uptake in the premonsoon. The high and prolonged
use of NO 23 at the beginning of the productive season was due to a strong allochthonous supply of NO23 , dominance
of microplankton, and low NH14 regeneration rates. Heterotrophs may take up all four nutrients and could account
for half of the annual total N uptake. Ammonium and NO22 regeneration rates were among the highest known from
nearshore waters and showed clear seasonal patterns. Production and use of NH14 were closely coupled. Nitrite
production rates were related to NH14 production rates in a rectangular-hyperbolic fit. Nitrogen balance analyses
showed that proximity to mangrove vegetation enhanced the flux rates, noninclusion of nitrification may lead to an
overestimation of new production by 30%, and regeneration in the plankton fraction provided about 40% more N
than was assimilated.
Since the demonstration that 15N is a sensitive tracer for ecosystems such as coral reefs and mangroves. Both are
measuring rates of nitrogen transformations in biological structurally and functionally among the more complex coast-
systems (Neess et al. 1962) and the postulation of the con- al marine ecosystems (Qasim and Wafar 1990) and together
cepts of new and regenerated production (Dugdale and Goe- account for more than 10% of global coastal primary pro-
ring 1967), there has been an increased interest in quanti- duction (Longhurst et al. 1995); yet studies on N cycles are
fying N fluxes in the marine environment. In the last three few in coral reefs and nonexistent in mangroves.
decades, the uptake of NO 23 and NH 14 , representing the ma- In an earlier study on a mangrove ecosystem from the
jor fractions of new and regenerated N, has been investigated central west coast of India (Wafar et al. 1997), a model for
using 15N in a number of instances (see Dortch 1990). The prediction of litter fall was constructed and elemental flux,
ability to separate the new production fraction from the total as C, N, and P, from mangrove vegetation to the aquatic food
production finds application in assessment of export of C chain was quantified. Their study also showed that most of
and N to the deep sea (Eppley and Peterson 1979) and in the terrestrial production ends up in mangrove waters. While
models of biogeochemical cycles of N in the sea (Platt et al. this was important for sustaining the microbial food chain
1991). The other component, regenerated production, is a and nutrient regeneration in the water column, it was still
useful indicator of ecological efficiency—under conditions inadequate for the N budget of the latter. The nitrogenous
of low new N fluxes, the ability of an ecosystem to sustain nutrition of the phytoplankton in mangrove waters, in order
high levels of biological production depends entirely on the to agree with the high productivity generally ascribed to
rates of internal recycling of N. This would be particularly them, therefore, would have to depend on a strong supply
true of tropical waters that, as a general rule, have low levels of new N or rapid in situ N recycling. The present study
of ambient nitrogenous nutrients at any time of the year was designed to examine this, by measuring over a seasonal
(Qasim and Wafar 1990). cycle the uptake rates of the major assimilable N forms
A rough estimate from the data of Longhurst et al. (1995) (NH 14 , NO23 , NO22 , and urea) and in situ regeneration of
shows that tropical waters account for between 20 and 40% NH 14 and NO 22 .
of global marine primary production. Nevertheless, with a
few exceptions, almost all studies on N uptake carried out
so far are from high-latitude ecosystems (Wafar et al. 1995). Material and methods
The imbalance is still more marked with high-productivity
The study site was an undisturbed mangrove forest of
1
Corresponding author (wafar@darya.nio.org). about 300 ha on the central west coast of India (168129–
168149N; 738259–738309E), fed by a small river and con-
Acknowledgments nected with the Arabian Sea by a single channel (Fig. 1).
We are grateful to E. Desa, D. Chandramohan, and A. G. Unta-
The study area experiences a monsoon climate; thus, the
wale for their support to this study and to B. J. Peterson, S. V.
Smith, W. G. Harrison, S. L’Helguen, and P. Wangersky for their traditional seasonal breakup is into monsoon (June–Septem-
thoughtful comments on the manuscript. Financial assistance to this ber), postmonsoon (October–January), and premonsoon
work was provided by a grant from the Office of Naval Research, (February–May). Monsoon is characterized by heavy rainfall
USA. This is Contribution 3691 from the National Institute of (2,500–3,000 mm), postmonsoon by dry and relatively cool
Oceanography, Goa. climate and premonsoon by dry, hot, and humid conditions.
241
242 Dham et al.
Heterotrophic
Average uptake uptake as % of
N nutrient Intercept Slope (95% CI) (nmol N L21 h21) average uptake
2
NO (low uptake)
3 23.0 2.9 (1.2, 4.5) 44 52
2
NO (high uptake)
3 49.0 18.4 (6.3, 30.3) 223 22
1
NH (low uptake)
4 21.3 4.8 (0.5, 9.4) 59 36
1
NH (high uptake)
4 44.4 31.6 (16, 88) 356 12
2
NO (at Sta. 1)
2 4.6 1.6 (0.2, 3.2) 16 29
Urea (at Sta. 1) 4.6 1.3 (0.15, 2.45) 26 35
Urea (at Sta. 2) 3.3 1.3 (0.5, 1.85) 30 22
All nutrients (pooled data) 179.0 18.7 (0.6, 36.5) 339 53
N flux in mangrove waters 245
Fig. 16. Seasonally integrated rates of uptake of (a) nitrate, (b) nitrite, (c) ammonium, and (d)
urea; (e) ammonium production and (f) nitrite production at the three stations.
tality in summer (Figs. 5a and 9b). A regression analysis of BENDSCHNEIDER, K., AND N. J. ROBINSON. 1952. A new spectro-
the f-ratios calculated with and without allowance for in situ photometric determination of nitrite in seawater. J. Mar. Res.
nitrification gave a slope of 0.7 (95% CI 5 0.46, 0.93, p , 11: 87–96.
0.001). The range and the average extent of overestimate BEROUNSKY, V. M., AND S. W. NIXON. 1993. Rates of nitrification
along an estuarine gradient in Narragansett Bay. Estuaries. 16:
(30%) are substantially of the same order as in coastal waters 718–730.
(Priscu and Downes 1985) or oceanic waters (Dore and Karl BODE, A., AND Q. DORTCH. 1996. Uptake and regeneration of in-
1996) and suggest that even in nearshore waters, where up- organic nitrogen in coastal waters influenced by the Mississippi
take of oxidized forms of N is presumably driven by inputs River: Spatial and seasonal variations. J. Plankton Res. 18:
of allochthonous N, estimation of new production from mea- 2251–2268.
surements of NO23 uptake alone would be of little value if BOTO, K. G., AND J. T. WELLINGTON. 1988. Seasonal variations in
not accompanied by those on nitrification. concentrations and fluxes of dissolved organic and inorganic
The third feature relates to the balance between uptake materials in a tropical, tidally dominated mangrove waterway.
Mar. Ecol. Prog. Ser. 50: 151–160.
and production of N. Ammonium regeneration and nitrifi- CARPENTER, E. J., AND S. DUNHAM. 1985. Nitrogenous nutrient up-
cation provided on average 20% more N than assimilated by take, primary production, and species composition of phyto-
plankton. The sinks for this excess dissolved N would be plankton in the Carmans River Estuary, Long Island, New
the mangrove vegetation and the benthic algae. Partitioning York. Limnol. Oceanogr. 30: 513–526.
of dissolved N by these consumers would then explain the DORE, J. E., AND D. M. KARL. 1996. Nitrite distributions and dy-
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and primary production in permanently well-mixed temperate Received: 15 February 2001
coastal waters. Estuar. Coast. Shelf Sci. 17: 431–446. Accepted: 30 July 2001
, , AND S. L’HELGUEN. 1995. f-ratios calculated with Amended: 18 October 2001