Hydrobiologia 436: 217–235, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

217

Nutrient fluxes in intertidal communities of a South European

lagoon (Ria Formosa) – similarities and differences with a
northern Wadden Sea bay (Sylt-Rømø Bay)

Ragnhild M. Asmus1 , Martin Sprung2 & Harald Asmus1

1 Alfred-Wegener-Institut für Polar-und Meeresforschung, Wadden Sea Station Sylt 25992 List/Sylt, Germany
2 CCMAR-UCTRA, Universidade do Algarve, Campus de Gambelas, P-8000810 Faro, Portugal

Received 28 May 1999; in revised form 3 July 2000; accepted 7 August 2000

Key words: nutrient flux, nitrogen, phosphate, tidal flats, temperature, geographical comparison

Abstract
During an annual cycle, flux rates of oxygen, nitrate, nitrite, ammonium, phosphate and silicate were measured
in light and dark bell jars at three sites in Ria Formosa (Algarve, Portugal) enclosing either a natural macrophytic
community (macroalgae on sand or mud, a seagrass bed of Zostera noltii) or bare sediments. The results are
compared with a preceeding study in which the same bell jar technique has been applied in the Sylt-Rømø Bay of
the northern Wadden Sea. Nitrate flux was mainly directed from the water column to the benthic communities in
Ria Formosa, as well as in the Sylt-Rømø Bay. However, nitrate uptake was higher in the northern, more eutrophic
study area. In Ria Formosa, nutrient concentrations were lower than in the Sylt-Rømø Bay possibly due to strong
water exchange with Atlantic waters. High temperatures and strong insolation had a greater impact on nitrate fluxes
in Ria Formosa than in the Sylt-Rømø Bay. Bioturbating macrofauna increased ammonium efflux in the Sylt- Rømø
Bay while this effect was not as pronounced in the Ria Formosa study sites. Benthic phosphate uptake dominated
in the Ria Formosa and was correlated to initial phosphate concentrations in incoming waters. At both study sites,
oxygen and nutrient fluxes were correlated with temperature. Additionally, flux rates were strongly influenced
by biotic components and levels of eutrophication. A literature survey showed that mainly in temperate regions,
material fluxes increase with temperature, whereas in warmer areas, ammonium and phosphate fluxes between
sediment and water were generally lower.

Introduction the state of oxidation of sediments is important for the

The nutrient and oxygen exchange between sediments
and water has been studied extensively in the tem-
perate regions of the United States and Europe since
the pioneering work of Pamatmat (1968), Rowe et al.
(1975) and Nixon et al. (1976). Despite differences
between ecosystems, some common features emerge.
Interactions of chemical, biological and physical pro-
cesses determine the extent to which nutrients are
released or taken up by benthic communities. Remin-
eralisation increases with the amount of labile organic
matter (Zeitzschel, 1980; Jensen et al., 1990, Sloth
et al., 1995), and benthic-pelagic fluxes are depend-
ent on the nutrient concentrations in the water column
(Boynton et al., 1980; Asmus, 1986). Additionally,
influx or efflux of nutrients (Rysgaard et al., 1996).
Hence, eutrophication is of great importance (Nowicki
& Oviatt, 1990). Benthic communities may alter ex-
change processes profoundly by bioturbation, excre-
tion (Kristensen, 1984; Hüttel, 1990), as well as sedi-
ment enrichment with organic material by macrofauna
(Baudinet et al., 1990). At least temporarily, retention
of nutrients in the benthic system is caused by benthic
microalgae (Granéli & Sundbäck, 1985; Rizzo, 1990),
seagrasses and macroalgae (Sand-Jensen & Borum,
1991; Duarte, 1995). Bacterial remineralisation may
be the source of nutrient release from sediments in-
cluding denitrification (Seitzinger, 1987), but bacteria
need inorganic nutrients as well and may act as nutri-
ent sink (Alongi, 1991; Van Duyl et al., 1993; Zweifel
218

et al., 1993). The annual temperature regimes in study

areas are of great importance to nutrient fluxes (Nixon
et al., 1976). Hydrodynamics are acting via currents,
waves and turbulence (Rutgers Van der Loeff, 1981;
Wildish & Kristmanson, 1997; Asmus et al., 1998;
Widdows et al., 1998). This may cause different res-
ults from mesocosms and flumes showing higher rates
than in benthic chamber and core incubations (Asmus
et al., 1998).
In this study, data from Ria Formosa (Southern
Portugal) are compared to previously published data
from Sylt-Rømø Bay (Northern German Wadden Sea)
using the same bell jar technique. Both ecosystems
are quite similar under many aspects, but separated
by 18◦ latitude (or 2350 km on a SW–NE axis). Main
structural difference are extensive salt marshes in the
Ria, not present in the Sylt-Rømø Bay. The two tidal
basins have little freshwater input but a large water
exchange by tidal inlets with coastal waters either of
the Atlantic or of the North Sea. Hence, they are
well mixed and oxygenated marine systems. Will the
community metabolism be possibly higher caused by
elevated temperatures in the southern ecosystem or
will other processes dominate?

Study sites
Ria Formosa
Ria Formosa is a tidal lagoon covering an area of
100 km2 at the southern coast of Portugal (Algarve).
It is separated from the Atlantic Ocean by several
sandy islands. Lagunal waters are exchanged with
the Atlantic by 6 deep inlets. The tidal range var-
ies between 2.8 m during spring tide and 0.6 m at
neap tide. The residence time is extremely short vary-
ing between half a day and 2 days (Neves et al.,
1996). Approximately two thirds of the area is inter-
tidal during mean low tide, one third of the total lagoon
area is occupied by saltmarshes or salinas (Teixeira
& Alvin, 1978). Sediments are mainly muddy along
the mainland coast and become more sandy close to
the dune islands. Perennial seagrass beds extend sub-
tidally (mainly Cymodocea nodosa (Ucria) Aschers.,
but also Zostera marina L. and Z. noltii Hornem.) and
intertidally (Zostera noltii). In the inner part of the
lagoon, green algae, mainly Ulva spp. and Entero-
morpha spp., but also Fucus spp. and Gracilaria spp.
regularly form thick mats in winter (up to 2970 g DW
m−2 ; Reis & Sprung, 1995) which decay during sum-
mer (except Fucus). On the sand flats, seasonality is
Figure 1. Study sites in the Ria Formosa (Algarve, Portugal) in
a muddy area (M), a sand flat (S) and a seagrass bed of Zostera
noltii (SG) growing on muddy sand. The Königshafen Bay (arrow)
is situated 2350 km northeast in the Sylt-Rømø tidal basin, a part of
the Wadden Sea.
more erratic. Salinity is between 36 and 38 all year
round apart from an occasional decrease after heavy
rainfall in winter (Falcão et al., 1985). Nutrient flux
measurements were carried out at a muddy station
(M), a sandy site (S) and in a muddy Zostera noltii
bed (SG) in the western part of Ria Formosa (Figure
1). Organic matter content of dried sediments ranged
from 1 to 1.4% for the sandy sediment, for muddy
sediments of the seagrass bed and under macroalgal
covers from 7 to 8% with little seasonal variation (loss
on ignition at 450 ◦ C for 3 h).
Sylt-Rømø Bay
In the North Sea, the Wadden Sea extends along the
coast of the Netherlands up to southern Denmark. At
the German-Danish border, the Sylt-Rømø Bay of 404
km2 is enclosed by the islands Sylt and Rømø and

2 dams connecting these islands with the mainland.
Water is exchanged between the North Sea and the
Sylt-Rømø Bay through one large tidal gully. The tidal
range is 1.80 m, and salinity varies seasonally between
28 and 32. The residence time in the Sylt-Rømø Bay is
about 51 days (Müller, unpublished). One third of the
Sylt-Rømø area is intertidal, only small salt marshes
exist. The study site ‘Königshafen’ is a small part (5.5
km2) of the total bay at the northern tip of the island of
Sylt. Main types of benthic communities of this Wad-
den Sea area are sand flats, mussel beds, small mud
flats and seagrass beds (Zostera noltii, Z. marina). Or-
ganic content of sandy sediments ranges from 0.5 to
1.0% and from 2 to 6% for muddy sediments (loss on
ignition) (Kristensen et al., 1997). Macroalgae, mainly
green algae, only thrive during particular summers to
a significant extent (Reise et al., 1989).
Materials and methods
Bell jar technique
Light (plexiglas) and dark bell jars (PVC, area 650
cm2 ) were pushed into the sediment just before the
tide reached the study site. During rising tide, the
bell jars were filled with water (volume 17.7 l). Wave
action was transmitted into the bell jars by the flex-
ible plastic foils covering their top. In the Sylt-Rømø
Bay, wave action was strong enough to secure water
mixing inside the bell jars, in Ria Formosa, an ad-
ditional hand driven stirrer was inserted. The study
sites in Ria Formosa were monitored every second
month during an annual cycle (1990–1991). On the
sand (S) and mud flats (M), one pair of bell jars was
placed over an existing algal patch, another pair was
installed at an adjacent site with no algal cover. In
the Zostera noltii bed, it was attempted to select sites
with maximum and minimum shoot density. Approx-
imately every hour, samples were taken for oxygen
determination by a syringe and analysed by Winkler
titration. Variation in oxygen content was calculated
by a linear regression of the oxygen content over time.
In the beginning and the end of incubation lasting 3–5
hrs additional samples were taken by syringes for nu-
trient analysis (nitrate, nitrite, dissolved phosphate and
silicate) following the methods described in Graßhoff
et al. (1983). The enclosed macrophytic and macro-
faunal biomass (dry weight and ash free dry weight,
respectively) was determined at the end of measure-
ments. Parallel to the measurements, the incident light
219
intensity was monitored by a discoidal sensor in air by
a Li-Cor recorder.
In the Sylt-Rømø Bay, flux rates of oxygen, ni-
trate, nitrite, ammonium and silicate were monitored
monthly during an annual cycle in 1980. A pair of light
and dark bell jars each were incubated on an Arenic-
ola sand flat, a Nereis sand flat and a sandy Zostera
noltii seagrass bed. The results from the bell jar meas-
urements in the Sylt-Rømø Bay have already been
published (Asmus, 1986) and serve as comparison to
the results obtained in Ria Formosa. In addition, nu-
trient fluxes and primary production were measured
in core incubations in 1993/94 in Königshafen at the
same sites showing similar nitrogen fluxes as some 10
years ago, but significantly higher oxygen production
in the 1990s than in the 1980s by microphytobenthos
(Kristensen et al., 1997; Asmus et al., 1998a, b).
Statistics
Because the data sets are comparably small (n<100),
the data have been treated statistically by nonpara-
metric tests. They are included in the STATISTICA
package (1994) (Mann–Whitney U -test, Spearman
R correlation, Wilcoxon matched pairs test; Sachs,
1984). Additionally multiple and nonlinear regression
analyses were applied.
Results
Temperature and light
Temperature and light conditions are distinctly differ-
ent in the northern and the southern European study
area. During the bell jar incubation, water temperature
at the sites in Ria Formosa varied from 12 to 13 ◦ C
in January and December and 24 ◦ C in August. The
lowest temperatures in the Ria are as high as those
in the Sylt-Rømø Bay in spring (May) and autumn
(September) (Figure 2). During winter, tidal flats of
the northern Wadden Sea are frozen regularly.
Mean incident radiation recorded during the exper-
iments around noon are distinctly higher in Ria For-
mosa than in the Sylt-Rømø Bay. The lowest winter
light intensity in Ria around noon is as high as in Sylt-
Rømø Bay in spring or autumn. In summer, highest
light intensity in Sylt-Rømø Bay reaches only values
of cloudy days in Ria Formosa (Figure 2).
220
Figure 2. Annual water temperature and light curves measured dur-
ing the bell jar incubations in Ria Formosa and Sylt-Rømø Bay.
Light intensity indicates the incident radiation above the water
surface around noon.
Seagrass, macroalgae and macrofauna
In Ria Formosa, seagrass beds are perennial. At the
intertidal study site, the biomass of Zostera noltii was
very high, showing little variation during the year
and approximately equal above and below ground
biomass (annual mean above ground biomass: 869
± 163 g DW m−2 , mean below ground biomass:
793 ± 202 g DW m−2 ) (Figure 3). In the northern
Wadden Sea, seagrass beds are losing their leaves in
winter, but the rhizomes of Zostera noltii are persist-
ent. Growing season for seagrasses begins in May in
the Sylt-Rømø Bay, they reach their highest biomass
in August/September and lose their leaves in autumn
(Asmus & Asmus, 1985, Asmus et al., 1998). The
maximum biomass of Zostera noltii in the Sylt-Rømø
Bay varies from year to year between 80 and 140
g DW m−2 (Asmus et al., 1998) and, therefore, it
is much lower than the annual mean biomass of the
Zostera noltii bed in Ria Formosa.
Macroalgae were present at the sandy study site of
the Ria Formosa from January until July (207 ± 53 g
Figure 3. Annual cycle of above ground biomass (g dry weight
m−2 ) of Zostera noltii at the study sites in the Ria Formosa and
in the Sylt-Rømø Bay.
DW m−2 ) and on the mud flat all year round (115 ± 50
g DW m−2 ). Green algae predominated (Ulva spp. and
Enteromorpha spp.) with the exception of bell jars at
the muddy site in summer. At this time, Fucus vesicu-
losus L. and Gracilaria spp. were abundant. Biomass
values given here originate from the bell jar measure-
ments only, and do not give large scale mean values
for Ria Formosa.
Data on the macrofauna extracted from the bell jars
have been published by Sprung (1994): their mean an-
nual biomass was estimated as 15.4 (sand flat), 17.2
(Zostera noltii bed) and 25.1 (mud flat) g ash free dry
weight m−2 . Biomass of the sand flat community was
characterised by suspension feeders dominated by the
gastropod Mesalia brevialis (Lamarck). Main species
of Zostera beds were polychaetes Melinna palmata
Grube, a suspension feeder, and Nereis diversicolor
O. F. Müller, an omnivorous species. On the mud flat,
only two species dominated biomass, the suspension
and deposit feeding bivalve Scrobicularia plana (Da
Costa) and the polychaete N. diversicolor.

In Königshafen of the Sylt-Rømø Bay, biomass of
macrofauna was nearly doubled of comparable sites
in the Ria Formosa (30.2 g AFDW m−2 on a Zostera
noltii flat, 27.6 g AFDW m−2 on an Arenicola flat;
Asmus & Asmus, 1985). In Zostera noltii beds, the
grazer Hydrobia ulvae (Pennant) dominated the mac-
rofauna community, whereas in the Arenicola flat, the
lugworm Arenicola marina (L.) had the highest share
in macrozoobenthic biomass.
Oxygen flux
At all three Ria Formosa sites, net oxygen production
was significantly higher in bell jars enclosing mac-
rophytes than in those with bare sediments (Mann–
Whitney U -test, two sided; sand with algae versus
sand without algae, p<0.050, n=11; mud with algae
vs mud without algae, p<0.020, n=12; seagrass vs
control, p<0.020, n=12). Looking at all sites, net
oxygen production in bell jars enclosing macroalgae
or seagrass was correlated with macrophytic biomass
(Spearman R, p=0.055, n=18). Therefore, photosyn-
thesis in terms of net oxygen production was not in
general correlated with light intensity. A significant
increase in net oxygen production with light intensity
was found in the control bell jars bare of macrophytes
at the sandy and at the seagrass site with abundant mi-
crophytobenthos (Spearman R, p=0.085, n=9). The
amount of macroalgal or seagrass biomass displayed
also a pronounced impact on the oxygen consumption
in the dark bell jars since it was significantly higher
in those enclosing macrophytes (Mann–Whitney U -
test, all sites p<0.02, n (with macrophytes)=18, m
(without macrophytes)=15).
Oxygen was most intensely produced by green
algae mats on sand, whereas the corresponding en-
closure of bare sand showed smaller rates of oxygen
production by microphytobenthos throughout the year
(Table 1) (Figure 4). Net oxygen production at the
muddy site with comparable algae cover was much
lower than at the sandy site with algae cover. In June
and August, the dense macroalga were hardly pro-
ductive in the muddy area (Figure 4). Seagrass beds
showed a distinct and regular net oxygen production
throughout the year, while in the control bell jars net
oxygen production was much smaller (Table 1).
Considering all sites, the oxygen production in
light bell jars was more than two times higher in
the Ria Formosa than in the Sylt-Rømø Bay (U -test,
p=0.010, n(Sylt)=25, n(Ria)=35) (Table 1)). This was
in the first instance due to missing macroalgae in the
221
northern area, but it also applies to seagrass beds
(p=0.03, n (Ria)=7, n (Sylt)=9, same test). Next to
the higher seagrass biomass in the Ria Formosa, this
difference is due to the short vegetational period in
the Sylt-Rømø Bay between May and October. Dur-
ing this time, primary production may be at times
as high as in Ria Formosa. However, considering
the entire year, seagrasses are more productive in the
Ria Formosa than in the Sylt-Rømø Bay. Bare sands
at comparable sites in the Sylt-Rømø Bay showed
a lower oxygen production in 1980 than in the Ria
Formosa in 1990/91 (U -test, p=0.03, n(Ria)=7, n
(Sylt)=9).
The oxygen consumption of comparable benthic
communities was similar in the Ria Formosa and in the
Sylt-Rømø Bay (seagrass beds in Ria vs Sylt, sandy
areas, no significant difference). In the Sylt-Rømø
Bay, oxygen consumption was mainly influenced by
the macrozoobenthic biomass and temperature (As-
mus, 1982; Asmus & Asmus, 1985), while in the
Ria Formosa community, respiration was mainly in-
fluenced by macrophytes.
Nitrate and nitrite fluxes
At all three study sites in the Ria Formosa benthic
communities, both with and without macrophytes,
took up nitrate intensively from the water column (sign
test Dixon & Mood, p=0.010, n=64) (Figures 5 and
6). In the seagrass bed and at the muddy site, ni-
trate uptake was higher than at the sandy site (U -test,
p=0.050). Sometimes, nitrate release was registered
in the bell jars without macroalgae on the sand flat
(Figure 5a). In the muddy area, nitrate was taken up
at high rates in light incubations of mud covered with
macroalgae from December until June, while a distinct
nitrate production dominated especially in dark incub-
ations of mud with macroalgae in the second half of
the year probably due to decaying processes (Figure
6a). Nitrite flux rates were distinctly lower than nitrate
fluxes and they were not correlated with one another.
At the sandy site, nitrite was mainly released while
uptake of nitrite dominated in the muddy and in the
seagrass area (Table 1).
In the northern and the southern study areas, the
nitrate fluxes were mainly directed from the water
column to the benthic communities. In the Sylt-
Rømø Bay, the uptake rates were significantly higher
than in Ria Formosa (U -test, p=0.004, n(Sylt)=48,
n(Ria)=64) (Figure 5). Uptake was highest in the Sylt-
Rømø Bay in winter, whereas in the Ria, maximum
 222

Table 1. Mean annual flux rates of oxygen (mg m−2 h−1 ) and nutrients (µmol m−2 h−1 ) given with (standard error) and (number of observations) measured in bell jars in different benthic
communities of Ria Formosa and Sylt-Rømø Bay, n.m.=not measured

O2 Prod. O2 Cons. Nitrate Nitrite Ammonium Silicate Phosphate

Light Dark Light Dark Light Dark Light Dark Light Dark

Ria Formosa
Sand + Alg. 715 (303) [4] −167 (39) [4] −3 (4) [4] −19 (16) [4] 2 (2) [4] 4 (3) [4] n.m. n.m. −206 (46) [4] −94 (53) [4] −6 (3) [4] −8 (1) [4]
Sand 130 (23) [7] −27 (7) [7] −2 (14) [6] 11 (18) [3] 4 (3) [5] 14 (10) [4] n.m. n.m. −53 (79) [6] 71 (81) [4] 3 (9) [6] 19 (15) [4]
Mud + Alg. 111 (27) [7] −32 (13) [7] −63 (40) [7] 33 (27) [7] −1 (2) [6] −3 (2) [7] 53 (60) [4] 253 (186) [5] 72 (75) [6] 36 (55) [6] −6 (4) [6] −7 (12) [6]
Mud 5 (18) [5] −7 (6) [4] −12 (6) [5] 16 (27) [6] −3 (3) [6] 1 (1) [4] 33 (53) [4] 206 (76) [4] 175 (96) [5] 124 (69) [6] −21 (17) [5] −13 (15) [6]
Seagrass 294 (55) [7] −68 (9) [7] −63 (28) [7] −30 (16) [7] −5 (2) [7] −2 (2) [7] −92 [1] 27 [1] 80 (60) [8] 60 (62) [8] −9 (5) [7] −8 (4) [7]
Control 77 (10) [5] −56 (18) [4] −40 (41) [4] −50 (25) [4] −3 (4) [4] 0 (1) [4] 215 [1] 223 [1] −11 (110) [4] −99 (39) [4] −2 (12) [4] −9 (6) [4]

Sylt-Rømø Bay
Arenicola flat 64 (14) [9] −32 (7) [9] −92 (51) [9] −132 (40) [8] −5 (5) [9] −1 (8) [8] −2 (48) [8] 55 (77) [7] 264 (379) [9] 74 (409) [8] n.m. n.m.
Nereis belt 79 (51) [8] −86 (23) [8] −35 (23) [8] −43 (26) [7] −4 (2) [7] −6 (2) [6] −41 (26) [6] −21 (29) [6] 402 (193) [7] 196 (109) [7] n.m. n.m.
seagrass bed 118 (56) [9] −98 (28) [9] −96 (37) [9] −65 (26) [7] −9 (2) [9] −7 (3) [7] −55 (46) [8] −85 (63) [5] −229 (107) [9] 5 (54) [6] n.m. n.m.
 223

Figure 4. Community oxygen gross (grey shading) and net production (+, white columns) and consumption (-) in light and dark bell jars with
and without macroalgae cover in sand (a) and mud (b) and in a seagrass bed of Zostera noltii (c) (August 1990 until December 1991). Biomass
of macroalgae and seagrass (above ground) is indicated by diamonds.

nitrate uptake was found in summer. This can be ex-
plained by distinctly higher nitrate concentrations in
tidal waters in winter and spring (up to 56 µmol l−1 ,
Asmus, 1986) (Table 2) in the Sylt-Rømø area, and the
correlation between nitrate concentrations and fluxes.
The higher the concentration, the more nitrate was
taken up in the Sylt-Rømø area. In summer, when ni-
trate concentrations were near zero, nitrate fluxes were
extremely low. In the Ria Formosa, the seasonal range
of nitrate concentration was smaller (up to 4.38 µmol
l−1 ) (Table 2) and there was no significant correlation
between concentrations and flux rates. However, ni-
trate fluxes increased with rising temperature in the
Ria Formosa (Spearman Rank correlation, all sites,
p<0.005, n=64). Nitrate uptake, as well as nitrate
release, were higher in summer than in winter.
Light intensity had a distinct impact on the nitrate
flux rates in the Ria Formosa. Nitrate uptake was sig-
224

Figure 5. Nitrate fluxes in light and dark bell jars (shaded). Positive rates show release, negative ones uptake by the enclosed community. (a)
Nitrate flux in sandy sediments with and without macroalgal cover. (b) Nitrate fluxes in seagrass beds of Zostera noltii in Ria Formosa with
bare sediments as control, and (c) in a Zostera noltii bed in the Sylt-Rømø Bay (redrawn after Asmus, 1986).

nificantly higher in light than in dark incubations, and Ammonium fluxes

on the other hand, release of nitrate was more intense
in dark than in light (Wilcoxon matched pairs test, all
sites, p=0.050, n=29). This was contrary to results
from the Sylt-Rømø Bay, where nitrate fluxes showed
no significant difference between light and dark bell
jars over the entire year. Nitrogen fluxes measured in
the 1990s were in the same order of magnitude than
those measured in the 1980s in the Sylt-Rømø area
(Kristensen et al., 1997; Asmus et al., 1998) so that
the differences are indeed site specific.
In the Ria Formosa, ammonium fluxes were mainly
measured at the muddy site. Release of ammonium
was more often registered than uptake (sign test Dixon
& Mood, n=17, p=0.02) (Figure 6b). Ammonium re-
lease rates was higher than uptake (U -test, p=0.050,
n=17). In light bell jars, ammonium uptake was signi-
ficantly higher than in dark bell jars where ammonium
release dominated (mud with macroalgae, mud, Wil-
coxon matched pairs test, p=0.05, n=8).

Figure 6. Nitrate and ammonium fluxes in muddy sediments of Ria Formosa with and without macroalgal cover of (biomass is indicated by
diamonds).
High ammonium effluxes and the macroalgal bio-
mass correlated significantly with release of nitrate at
the mud site between June and December during algae
cover (multiple regression, R=0.999, no. of cases 4,
p=0.018, ammonium beta=0.91, biomass beta=0.44).
Highest ammonium release rates were monitored in
bell jars enclosing partially decaying macroalgae in
summer. In dark incubations, ammonium fluxes and
nitrate fluxes demonstrated a significantly positive cor-
relation both in bell jars with and without macroalgae
(r=0.758, n=9, Spearman R: p=0.01) indicating a pos-
225
sible nitrification of ammonium in the well oxygen-
ated enclosures. The seagrass bed took up ammonium
in the light bell jars and released small amounts of am-
monium in the dark as evidenced during a campaign in
September 1991 (Table 1).
Both in the Ria Formosa and Sylt-Rømø Bay, am-
monium fluxes were influenced by light emphasising
the importance of uptake by micro-and macrophytes.
In the Sylt-Rømø Bay, bioturbating macrofauna, es-
pecially the lugworm Arenicola marina increased am-
monium release from the sediment (Asmus, 1986).
226
Table 2. Annual range of nutrient concentrations (µmol l−1 ) in
incoming tidal waters in Ria Formosa and Sylt-Rømø Bay during
bell jar experiments.
Nitrate Nitrite Ammonium Silicate
Sand 0.3–1.7 0–0.12 3.0–13.4
Mud 1.1–3.5 0–0.33 2.4–11.1 6.5–29.2
Seagrass 0.2–4.4 0.30–0.58 5.8–11.1 3.6–12.7
Sylt-Rømø 0–56.6 0.19–2.31 0–10.2 5.5–87.2
Bay
n.m.=not measured, ∗ phosphate concentrations in 1984.
This bioturbation effect could not be documented
for Ria Formosa stations. Here, correlations between
ammonium, nitrate and phosphate fluxes imply that
ammonium effluxes were directly caused by reminer-
alisation.
Phosphate and silicate fluxes
Considering all sites, uptake of phosphate by the
benthic communities prevailed (sign test, Dixon &
Mood, p=0.010, n=61) (Figure 7). Phosphate fluxes
were dependent on concentrations in incoming wa-
ters (Table 2); phosphate was taken up at elevated
phosphate levels while efflux dominated at low con-
centrations (Spearman Rank, n=63, p=0.0005). In
general macroalgae and seagrass increased the tend-
ency for the uptake of phosphate by the benthic com-
munity. Low uptake rates of phosphate were meas-
ured at the sandy site with macroalgal cover, whereas
bare sand showed a tendency to release phosphate
(U -test, p=0.050, n=10). On mud flats with and
without macroalgal cover, phosphate was mainly taken
up except for some occasions during summer with
simultaneous high effluxes of ammonium (mud + al-
gae, June, August, dark bell jars). Here, phosphate
fluxes in light and in dark incubations were correlated
with the corresponding ammonium fluxes (PO4 3− flux
dark vs. NH4 + flux dark: r=0.879, n=9, Spearman
R: p=0.035; PO4 3− flux light vs. NH4 + flux light:
r=0.604, n=8, p=0.05). The seagrass bed took up
phosphate intensively, whereas the phosphate rates at
the control site with low shoot density varied between
release and uptake. Phosphate flux rates were signific-
antly positively correlated with nitrate fluxes in dark
incubations (all sites, r=0.50, n=30, p=0.01). Phos-
phate was taken up simultaneously with nitrate, nitrate
release was high when phosphate release increased.
Silicate concentrations in tidal waters were gen-
erally high (Table 2). Silicate was taken up by the
sand flat (Figure 8), whereas in most cases, silic-
Phosphate ate was released at the muddy site. In the muddy
0.35–0.86 seagrass bed, efflux of silicate dominated compared to
0.50–1.64 the control (U -test, p=0.050, n=24). In light bell jars
0.25–1.52 there was a significant positive correlation between
net oxygen production and silicate flux rates (n=32,
0∗ –4.91∗
p=0.003). Silicate uptake was elevated at high primary
production, showing the importance of diatoms. Tem-
perature had a promoting effect on silicate flux (n=63,
p=0.011).
In analogy to ammonium fluxes, silicate fluxes in
the Ria Formosa were mainly caused by the activity of
micro- and macrophytes with emphasis on benthic and
epiphytic diatoms. In the Sylt-Rømø Bay, bioturbating
macrofauna was important as well for the silicate flux
resulting in a silicate release from sandy sediments
(Asmus, 1986). The muddy seagrass bed in the Ria
showed silicate release, while a sandy seagrass bed in
the Sylt-Rømø Bay showed silicate uptake. This dif-
ference is probably due to a combination of different
sediment type and epiphytes.
Discussion
Ria Formosa versus Sylt-Rømø Bay
Functional differences between Ria Formosa and the
Sylt-Rømø Bay are mainly caused by varying climatic
conditions between southern and northern European
coastal areas. The effects of temperature, light, biotic
community composition and initial nutrient concentra-
tions on nutrient and oxygen flux rates in both study
sites are summarised in Table 3.
In the Sylt-Rømø Bay, the low winter temperatures
around freezing point are limiting factors for biolo-
gical and chemical processes. The promoting effect of
rising temperatures may reach optimal conditions in
summer, but high temperatures may only have harm-
ful effects in exceptionally hot summers during low
tide. At times, this may be a problem in the Ria For-
mosa, comparable to the nearby Spanish Palmones
River Estuary where green algae (Ulva species) suffer
from thermal stress during summer (Hernández et al.,
1997). In the muddy area of Ria Formosa, macroalgal
primary production was reduced at high summer tem-
peratures. Most marine algae die off above 35 ◦ C
(Lüning, 1985). Similar differences may be expected
for the light climate. In northern tidal flats, primary

Figure 7. Phosphate exchange rates between benthic communities and overlying water in Ria Formosa (light and dark bell jars, + release, -
uptake, August 1990 until December 1991).
production is light limited for a short period in winter
and light saturated in summer (Admiraal, 1977). In
Ria Formosa, even in winter, high photosynthesis
of seagrasses and micro- and macroalgae was found.
Damaging effects of a very high insolation in summer,
especially UV-radiation, are still to be investigated.
Both areas differ with respect to the level of eu-
trophication. Nixon (1995) defined eutrophication as
an increase in the supply rate of organic matter to
an ecosystem. We can only speculate about the sup-
ply of organic matter in the Ria Formosa. Probably
there is a large export of detritus from salt marshes to
227
tidal flats entering the food web and remineralisation
cycles. Hence, we can only judge indirectly on the
eutrophication level by inorganic nutrients. These con-
centrations, especially of nitrate, are relatively low in
the study sites of the Ria Formosa. This is at first sight
surprising, since sewage treatment is still in its initial
phase in the Algarve which is densely populated and
visited by numerous tourists. Additionally, the culture
of fruits and vegetables is intense. The negative ef-
fects of all these sources of pollution are reduced by
strong tidal exchange with Atlantic waters causing an
extremely short residence time.
228
Figure 8. Silicate uptake and release rates measured at 3 study sites (a) sand with and without macroalgae, (b) seagrass and control, (c) mud
with and without macroalgae in Ria Formosa.
In the Sylt-Rømø Bay, nutrient concentrations are
much higher, especially the winter values of nitrate.
The nutrient load of coastal waters of the North Sea
and the Wadden Sea is very high, and oxygen deple-
tion is mainly prevented by strong tidal hydrodynam-
ics (De Jonge et al., 1994). In the Sylt-Rømø Bay,
an elimination of nitrogen via denitrification is small
(annual range from 1 to 56 µmol m−2 h−1 ) depend-
ing largely on nitrate concentrations in the overlying
water (60–90% of the total denitrification) (Jensen et
al., 1996). Also, nitrification is low (annual means of
6.1–16 µmol m−2 h−1 ) with a low coupling to de-
nitrification (Jensen et al., 1996). Measurements of
denitrification are still missing in the Ria Formosa.
Although climatic conditions showed a strong im-
pact on oxygen and nutrient fluxes, the rates of ma-
terial exchange were not only simply higher in the
southern study site. The interplay of involved forces
determined the balance between uptake and release of
material by benthic communities.
North versus South
The promoting effect of rising temperatures on oxygen
and nutrient fluxes could be demonstrated in numerous
regional studies (Nixon et al., 1976, 1980; Zeitzschel,
1980, Table 4). Considering the complex interactions

Table 3. Summary of effects of different ecosystem variables on nutrient flux rates in the Ria Formosa and in the Sylt-Rømø Bay.
Statistically significant correlations are indicated by ∗
Temperature Light
Ria Formosa
Community Oxygen Production
Microbial Oxygen Production ∗ ∗
Community Oxygen Consumption
Nitrate Flux ∗ ∗
Ammonium Flux ∗ ∗
Phosphate Flux
Silicate Flux ∗ ∗
Sylt-Rømø Bay
Community Oxygen Production ∗ ∗
Community Oxygen Consumption ∗ ∗
Nitrate Flux
Ammonium Flux ∗ ∗
Silicate Flux
regulating the material exchange between sediment
and water, the question arises whether this relationship
is still valid in a biogeographical comparison between
northern and southern areas. Only a small selection
of the meanwhile huge number of studies in temper-
ate regions can be presented (Table 4). For reasons
of comparison, consideration is given mainly to stud-
ies, in which light and dark benthic chambers have
been used in shallow waters, since nutrient fluxes vary
with water depth (Banta et al., 1995). Core incuba-
tions are in many respects comparable to bell jars as
far as the size of the enclosed sediment area is con-
cerned (Asmus et al., 1998), and in this context they
are mainly considered to fill in geographical gaps. Ex-
change rates of bare sandy to muddy sediments with a
natural macrofauna population, except dense aggrega-
tions of bivalves, are compared from northern regions,
with ice cover in winter, to southern sediments of coral
reefs, river estuaries and a mangrove system (Table 4).
Oxygen, nitrate and silicate fluxes showed no clear
North South trend. In shallow tidal and subtidal flats,
oxygen production by microphytobenthos is high (e.g.
Narragansett Bay, Chesapeake Bay, Ria Formosa,
Sylt-Rømø Bay, Table 4). The oxygen consumption
displays a broad range between −4 (Ems Dollard es-
tuary, Van Es, 1982) and −314 mgO2 m−2 h−1 (Bay
of Cadiz, Forja et al., 1994) without relation to an-
nual medians of temperature. The adaptation of micro-
and macrobenthic populations to their environmental
229
Concentrations Presence of Macrophytes
in the water column or Macrofauna
∗
∗
∗
∗ ∗
∗
∗
∗ ∗
∗
temperature ranges seems to prevent a biogeograph-
ical increase in community oxygen flow from North to
South. Nitrate has been released in several studies, but
uptake of nitrate by sediments is frequently recorded
in relation to ambient nutrient concentrations (Boyn-
ton et al., 1980). Silicate is most frequently released
from sediments caused by temperature dependent dis-
solution processes (Lewin, 1961) and is enhanced by
bioturbation (Helder & Andersen, 1987; Gehlen et al.,
1995). Silicate uptake is caused by benthic diatoms in
the studies cited here. In coral reefs, net silicate fluxes
between sediment and water are small or negligible
(D’Elia & Wiebe, 1990). Reef sediments consist to
a large part of eroded deposits from corals and other
carbonate bearing organisms as molluscs (for review
see Alongi, 1989).
In most study areas, the release of ammonium
dominates over ammonium uptake. However, in areas
of high microphytobenthic activity, ammonium is
taken up intensively (Rizzo, 1990; Sundbäck et al.,
1991). Nixon et al. (1976) measured a strong increase
in ammonium efflux between temperatures just above
0 and up to 24 ◦ C. Additionally, Fisher et al. (1982)
found high values of ammonium release between 15
and 25 ◦ C. A similar trend is evident in a worldwide
comparison. In areas of low annual mean temperatures
(9 – 15 ◦ C), small ammonium release rates are found,
while in warmer regions (15 – 20 ◦ C), also higher
ammonium effluxes are measured (Table 4, Figure 9).
230
Table 4. Oxygen flow (mg O2 m−2 h−1 ) and nutrient flux rates (in µmol m−2 h−1 ) measured in northern and southern areas. Oxygen units
were converted from ml O2 to mg O2 by the factor 1.43, and mmol O2 to mg O2 by the factor 32. o.m.=organic matter=Loss on ignition., l. and
d.=light and dark bell jars

Study area Sediments Water Bell jars Volume; area Temperature◦ C

Depth (m) Min. Max. Median

USA, Atlantic Coast,

North to South
Narragansett Bay sand to silt-clay, o. m. 1.5 – 4.5% subtidal 5.8 – 7.3 d. 27 l; 0.24 m2 0 24 12
Potter Pond, Rhode Island mud, o.m. 15% <1 l.+d. 3l; 0.02 m2 −1.5 28 13.3
Potter Pond, Rhode Island sand, o.m. 2% <1 l.+d. 3 l; 0.02 m2 −1.5 28 13.3
Narragansett Bay different sediments d. 27 l; 0.25 m2 5 20 12.5
Chesapeake Bay, Patuxent estuary silts to clays, o.m. 1.5 – 2.6% 3 d. 38 l; 0.3 m2 3 29 16
Chesapeake Bay, Potomac River sandy mud to fluid mud subtidal d. 32 – 38 l; 0.16 – 0.3 m2 16
Chesapeake Bay, Cherrystone Inlet sand, silt-clay, o.m. 0.7 – 5.2% subtidal 1 l.+d. 7.2 l; 0.07 m2 6.5 32.4 19.5
Pamlico Sound, North Carolina organic rich <5 l. 62 l; 0.31 m2 1 24 12.5
Pamlico Sound, Cape Lookout Bight organic rich, org. carbon 2–3% >5 l. 4 l; 400 cm2 5 27.5 16.3
25 – 35 l; 0.2 – 0.4 m2
Georgia Bight fine sand, o.m. 0.7 – 2.5% 5, tides 2 d. 47.5 l; 0.25 m2 10 28 19

Siberia, Pacific Coast sand to mud, org. carbon 0.2% – 1.6% 3–7 l.+d. 2 l; 15 l; 0.05 m2 −1.6 20.5 9.5

Australia, Pacific Coast

Port Phillip Bay sand, sandy mud, mud, clay 7 – 24 d. 7 l; 0.073 m2 15

USA, Pacific Coast

San Francisco Bay 1.50 12 17 14.5
Tomales Bay sand to fine mud 4–6 l. 33 l; 0.2 m2 9 19 14

Europe, North to South

Baltic Sea, Gulf of Bothnia soft silt subtidal cores, lab. 400 ml; 50 cm2 4 15 9.5
Baltic Sea muddy sand, org. carbon 0.49% 20 l. 2094 l; 3.14 m2 8 12 10
Königshafen sand, o.m. 0.5% intertidal <1 l.+d. 4 – 18 l; 600 – 700 cm2 0 19.6 9.8
Königshafen sand – mud, o.m. 0.5 – 6% intertidal < 1 l.+d. cores 0.5 l; 50 cm2 3.6 22.2 12.9
Ems-Dollard estuary mud, silt, sand, org. carbon 0.5 –14% intertidal l.+d. 707 cm2 3.8 19.1 11.5
Ems-Dollard estuary mud, silt, sand, org. carbon 0.5 –14% intertidal l.+d. 3–6l 3.8 19.1 11.5
Shannon estuary mud, o.m. 1.4 – 8% intertidal d. cores 50 cm2 10 20 15
Tamar Estuary, SW England mud, org. carbon 1.0 – 2.57% intertidal lab. 314 cm2
Bay of Morlaix, France fine sand intertidal, 1 – 4 d. 40 – 50 l; 0.2 m2 6 18 12
Marennes-Oléron, France mud shallow intertidal l.+d. 0.3 l; 40 cm2 10 28 19
Atlantic, Bay of Cadiz clay, org. carbon 0.78 – 2.16% 0.5 – 8.5 d. 59.1 – 89.8 l; 0.385 m2 18.4
Atlantic, Ria de Formosa sand to mud intertidal aquaria 17 20 18.5
Atlantic, Ria de Formosa sand to mud intertidal <1 l.+d. 17.7 l; 650cm2 12 23.3 17.7

Mediterranean Sea
Thau lagoon &Carteau bay under + outside mussel culture, mud subtidal, 3.5 – 4 l.+d. 13.8 l; 0.11 m2 5.3 22.8 14.1
Thau lagoon &Carteau bay org. carb. 2.9 – 8.2%, o. m. 10.6 – 29.4%
Thau lagoon, France under + outside oyster culture, mud 5 l. 15 l; 0.114 m2 4 26 15
Golfe de Fos, France u.+o. mus. cult., muddy sand, sandy mud ca. 5 l. 17 l; 0.13 m2 7.6 20 13.8
Alfacs Bay, Ebro Delta sand to mud subtidal <1.5 – 5 d. 7 l; 700 cm2 19.5
Alfacs Bay, Ebro Delta sand to mud, o.m. 1 – 10.4% subtidal <1.5 – 5 d. 7 l; 700 cm2 19.5

Tropics
Great Barrier Reef, Australia carbonate sands, o. m. 0.3 – 2.8% 2 – 16m light 315 l; 0.9 m2 25
Heron Island, Australia coral sands 0.5 – 8 l.+d. 900 cm2 25
Kaneohe Bay, Hawaii, eutrophied sand to mud, org. carbon 2.9% 1 – 15−17 l.+d. 262 l; 7854 cm2 25
Kaneohe Bay, after sewage diversion l.+d. 262 l; 7854 cm2 25
Coconut Island, Kanehoe Bay sand + coral rubble, sandy silt subtidal, 1 – 1.5 l.+d. 262 l; 7854 cm2 25
St. Croix, US Virgin islands sand, org. carbon 0.19 – 0.59% subtidal d. 8 – 11 l; 0.049 m2 25
Papua New Guinea, Fly Delta mud intertidal l.+d. 1 l; 0.007 m2 31 32 31.5
New Caledonia sands + mud subtidal, 6.7 – 17 l.+d. 51.6 – 72.5 l; 0.2 m2 23.8 27.8 25.8

Joyuda Lagoon, Puerto Rico mangrove-fringed lagoon cores 35.26 cm2 25

Continued on p. 231
 231

Nitrate Ammonium Phosphate Silicate Oxygen flux Authors

Range Mean Range Mean Range Mean Range Mean Range Mean
from to from to from to from to from to

< −40 120 0 400 0 60 −150 −10 Nixon et al. (1976, 1980)
−80 80 20 440 −10 50 −190 133 Nowicki & Nixon (1985a, b)
−100 20 0 60 −10 10 −75 ca. 100 Nowicki & Nixon (1985a, b)
12 500 4.2 233 41.7 392 Elderfield et al. (1981)
−674 100 −105 1584 1 295 −171 −21 Boynton et al. (1980)
−300 250 −129 1083 −25 167 41.7 1042 −112 −5.2 Callender & Hammond (1982)
−71 28 −44 615 5.3 42 −219 212 Reay et al. (1995)
0 6.4 0 454 −8.3 46 −100 −26 Fisher et al. (1982)
0 0 0 28 1023 525.5 −4 97 46.5 Klump & Martens (1981)

ca. 2 ca. 35 10.9 ca. 25 ca. 104 57.7 ca. 8 ca. 50 22.3 −51 −142 Hopkinson (1985, 1987)

13 30 0.7 129 0.6 38 18 492 −39.3 54.8 Propp (1977) Propp et al. (1980)

−1.7 37.9 4.2 500 1.7 83.3 66.7 770.8 −136 −25 Berelson et al. (1998)

−45.8 −4.2 −21 4.2 66.7 46 −4.2 4.2 4.2 −83.3 458.3 246 −61 −17 −40 Hammond et al. (1985)
−4.2 2.5 −0.42 51 117 82 5.4 17.1 10 58 196 151 −20 −4.9 −12.5 Dollar et al. (1991)

2.8 11.7 0.2 30.9 −77.4 −15.3 Nedwell et al. (1983)

N: 18.3 2.3 66.7 −10.7 Balzer (1984)
−354 173 −290 371 −2638 1493 −195 416.5 Asmus (1986) Asmus & Asmus (1985)
−492 26 −96 150 −115.2 352 Kristensen et al. (1997)
−196 −4 Van Es (1982)
−208 0 −250 83 −8 42 −146 −21 −88 −16 Rutgers Van der Loeff et al. (1981)
−62 0 −31 40 300 170 0 80 40 −24.6 −7.2 Brennan & Wilson (1993)
−250 375 62.5 −24 170 −7.4 23 −210 131 Watson et al. (1993)
−120 60 51 369 210 −163 −22 Boucher & Boucher-Rodoni (1988)
−159.8 −34.3 11.7 209.4 −5.3 27 49.1 529.8 Feuillet-Girard et al. (1997)
0 0 0 258.3 1525 737.5 20.8 379.2 183.3 258.3 1479 616.7 −314.4 −92.1 −183.3 Forja et al. (1994)
−7.3 −1.0 −4.2 0 38.0 19.0 −0.3 4 1.9 −2.4 31.7 14.6 Falcao & Vale (1990)
−156 108 −24 −71 292 110.5 −88 63 −12.5 −314 440 63 −90.5 191 this study

15 382 −69 21 Barranguet et al. (1994)

−158 23.8 −30 600 −14.9 53.2 −160.9 84 Mazouni et al. (1996)
−59 5 3.5 366 −3.2 116.2 15.5 1206 Baudinet et al. (1990)
−22 8 44 100 70 −86.7 −38.3 −54.4 Vidal et al. (1989)
0 160 −2 −76.2 Vidal et al. (1992)

5.96 22.67 0 1.17 2.38 33.58 Hansen et al. (1987)

−4.4 4.1 Sorokin (1992)
4.2 54.9 3.0 128.8 −18.8 Smith et al. (1981)
5 37.1 1.9 127.5 −11.3 Smith et al. (1981)
−38 354 62.5 80 1146 45.8 −4.2 71 20.8 0 0 −170.8 Henderson (1981)
−131 141 −5 −90 372 71 Williams et al. (1985)
−20 −16.2 13.3 −16.6 3 Alongi (1991)
−3.2 0.43 6.8 30.3 −140.2 −27.8 −95 Boucher et al. (1994)
Boucher & Clavier (1990)
−6.4 0 −2.7 −7.4 250 50 Correodor & Morell (1989)
Morell & Corredor (1993)
232
Figure 9 Ammonium flux rates (median (dark diamond), minimum
(light triangle) and maximum (dark triangle)) from geographical
regions with different annual median water temperatures (see also
Table 4). The positions of the Sylt-Rømø area and the Ria Formosa
are indicated.
Figure 10 Phosphate flux rates (median (above), minimum (light
diamond) and maximum (dark diamond)) from regions with differ-
ent median annual water temperatures (see also Table 4).
Simultaneously, in all these areas, there are studies
presenting only small ammonium release rates as well.
Going further to tropical regions, we mostly find lower
values. Phosphate flux rates show a very similar pat-
tern as ammonium fluxes. Release is generally higher
Figure 11 Cumulative curves of means of maximal fluxes of
ammonium (N) and phosphate (•) in relation to annual median
values of temperature in intervals of 2 ◦ C (ammonium vs tem-
perature: y=−4304.6+515.344∗ x−8.91xˆ2+0.0842857, r=0.9778;
phosphate vs temperature: y=−761.706+98.324∗ x−1.993∗ xˆ2-
0.1 304285, r=0.9829). These curves are based on data from a
literature survey (Table 4).
and more frequent than uptake, and higher phosphate
effluxes are found at elevated temperatures in temper-
ate regions. In subtropical and tropical studies, lower
ranges and means of phosphate fluxes are recorded
(Table 4, Figure 10).
A plot of cumulative means of maximum am-
monium and phosphate release rates versus annual
median temperatures (Figure 11) display a steep, al-
most linear increase in temperate areas. However,
the incline is small at higher temperatures of sub-
tropical and tropical areas (Polynomial function fit-
ting: second order polynom, ammonium vs temper-
ature: y=−4304.6+515.344∗x−8.91xˆ2+0.0842857,
r=0.9778; phosphate vs temperature: y=−761.706+
98.324∗x−1.993∗xˆ2−0.1 304285, r=0.9829; STAT-
ISTICA, 1994). In subtropical and tropical areas,
lower flux rates are found in organically poor sed-
iments. But also high rates are recorded, as high
ammonification in a Brazilian tropical lagoon rich in
labile organic material (Fernex et al., 1992). Elev-
ated nutrient fluxes are also found in tropical studies
in eutrophic areas, as in the coastal ecosystem of the
Kaneohe Bay (Henderson, 1981; Smith et al., 1981).
In the study by Smith et al. (1981), definitely lower
ammonium and phosphate fluxes were measured after
the reduction of the eutrophication by sewage input
into the bay. The entire level of nutrient flux was lower
than in studies of temperate areas. In comparison to the
hypoxic, organic rich sediments of temperate areas,
sediments of coral reefs are more often poor in organic
matter and well oxygenated. Nutrient concentrations
in porewater and in the water column are lower than

in temperate areas (Williams et al., 1985; Capone et
al., 1992; Charpy-Roubaud et al., 1996). The res-
ulting flux rates between sediment and water were
often too low to be measured in bell jars. Thus, other
methods have been applied, as calculation of diffusion
from porewater profils (Capone et al., 1992; Charpy-
Roubaud et al., 1996), tracer studies (Sorokin, 1992)
or flow respirometric input-output balances (Charpy-
Roubaud et al., 1990; for review see D’Elia & Wiebe,
1990). The flux rates from these studies cannot be
compared directly to those of benthic chambers due
to methodological differences. Hence, they are not
included in Table 4. However, their results give fur-
ther evidence of generally low nutrient fluxes in the
tropics. In these areas, dissolved nutrients are taken
up rapidly by organisms and also remineralisation is
very intense. Thus, the nutrient cycles in coral reefs
seem to be quite closed. While in northern areas, the
pool of organic labile substrate of detritus is large
and forms an omnipresent basis for nutrient remin-
eralisation, in coral reefs most nutrients seem to be
incorporated in living organisms and to a lesser extent
in detritus (Hatcher, 1985; Sorokin, 1992). N-fixation,
nitrification and denitrification are very important in
tropical shallow waters. Furthermore, bacteria can be a
dominating nutrient sink in tropical sediments. Alongi
(1991) revealed in an extensive benthic chamber study
that the release of nutrients in the Fly delta in Papua
New Guinea was extremely low presumably due to
uptake of nutrients by benthic bacteria.
In a geographical view, temperature dependence
of material fluxes is a phenomenon of the temperate
zone. Flux rates in subtropical and tropical regions are
limited by other factors and interactions of environ-
mental conditions and processes. In each study area,
either in cold or warm regions, certain ecosystem fea-
tures may dominate the material flux, overriding the
ecologically basic temperature impact. The regionally
different interactions of chemical, physical and biolo-
gical processes determine to which degree oxygen and
nutrients are either taken up or are released by benthic
communities.
Acknowledgements
We wish to thank Asmus Petersen and Birgit Lembke
for asssistance during the field measurements in the
Ria Formosa. This study is based in part on results
from the German-Portuguese research project “Die
233
Biologie der Ria Formosa” sponsored by the Ministry
for Research and Technology (grant no. 03FO562A).
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