Marine eutrophication review

[From the U.S. Government Printing Office, www.gpo.gov]

Science for Solutions

NOAA COASTAL OCEAN PROGRAM
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Decision Analysis Series No. 4
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MARINE EUTROPHICATION REVIEW

PART 1: QUANTIFYING THE EFFECTS OF NITROGEN ENRICHMENT
ON PHYTOPLANKTON IN COASTAL ECOSYSTEMS

PART 2: BIBLIOGRAPHY WITH ABSTRACTS

Kenneth R. Hinga
Heeseon Jeon
Noiffle F. Lewis

January 1995

U.S. DEPARTMENT OF COMMERCE
QH91.8 National Oceanic and Atmospheric Administration
.M34 Coastal Ocean Office
H45
1995

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Science for Solutions

9 0

NOAA COASTAL OCEAN PROGRAM
Decision Analysis Series No. 4

EMWT OF

MARINE EILITROPHICATION REVIEW

PART 1: QUANTIFYING THE EFFECTS OF NITROGEN ENRICHMENT
ON PHYTOPLANKTON IN COASTAL ECOSYSTEMS

PART 2: BIBLIOGRAPHY WITH ABSTRACTS

Kenneth R. Hinga
Heeseon Jeon
Nodile F. Lewis

Graduate School of Oceanography
University of Rhode Island
Narragansett, RI 02882

January 1995

U.S. DEPARTMENT OF COMMERCE
Ronald H. Brown, Secretary
National Oceanic and Atmospheric Administration
D. James Baker, Under Secretary property of CSC L:Lbrary
Coastal Ocean Office
M Q2@'
/4'4

Donald Scavia, Director
US Department of Commerce
NOAA Coastal Services Center Library
2234 South Hobson Avenue
Charleston, SC 29405-2413

This publicatinn qhntjlfi hp r*tpfi sq-

Hinga, Kenneth R., Jeeseon Heon and NodIle F. Lewis. 1995. Marine Eutrophication Review--Part 1:
Quantifying the Effects of Nitrogen Enrichment on Phytoplankton in Coastal Ecosystems; Part 2:
Bibliography with Abstracts. NOAA Coastal Ocean Program Decision Analysis Series No. 4. NOAA
Coastal Ocean Office, Silver Spring, MD. Part 1, 36 pp.; Part 2, 120 pp.

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Director
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MARINE EUTROPHICATION REVIEW

PART 1: QUANTIFYING THE EFFECTS OF NITROGEN ENRICHMENT
ON PHYTOPLANKTON IN COASTAL ECOSYSTEMS

TABLE OF CONTENTS

ABSTRACT 1

INTRODUCTION 1

DEFINING TERMS I

APPROACHES TO FINDING A RELATIONSHIP BETWEEN 2
EUTROPHICATION AND NUTRIENT INPUTS

NITROGEN AND PRIMARY PRODUCTION IN 4
MARINE ECOSYSTEMS

PHYTOPLANKTON ABUNDANCE AND OTHER MEASURES 5
THAT MAY BE RELATED TO EUTROPHICATION

OTHER COMPLICATING ISSUES 6

NITROGEN LOADINGS OR NITROGEN CONCENTRATIONS? 6

RELATIONSHIPS BETWEEN NITROGREN AND EUTROPHICATION 8

THE ROLE OF ANOTHER FACTOR 10

CURVE FITS TO CROSS-SYSTEM AND MERL DATA AND 11
COMPARISON TO THE EFFECT FACTORS FROM CHANGES
OVER TIME

CONCLUSIONS 11

TABLE 12

FIGURES 14

LITERATURE CITED 33

ABSTRACT

The results of three different approaches are used to investigate relationships between nitrogen
availability and phytoplankton primary production and abundance in coastal ecosystems. The
three approaches are controlled experiments in marine enclosures, history of changes in coastal
ecosystems, and cross-system comparisons. Most systems can be expected to have increased
primary production with increased nitrogen loadings or long-term average concentration.
Nitrogen availability alone is not a very precise predictor of phytoplankton production or
abundance. The magnitude of the change in phytoplankton production or abundance found for
most systems is typically in the range of 1.4 to 3 times for a doubling in nitrogen loading or
concentration. However, there are coastal ecosystems which have not followed the general
relationships found. Until the characteristics which sets those systems apart from other
systems can be identified, there is no guarantee that any individual system will have the typical
response.

INTRODUCTION

Perhaps the paramount question placed before the modem ecologist is "What will happen
if ... T' As this review considers anthropogenic nutrient inputs to coastal environments, the
question becomes: "What will happen if nutrient loads to a body of water increase?" And,
conversely: "What will happen if (usually at great expense) nutrient inputs to a coastal body of
water are reduced?" The answers to these questions are far from trivial to obtain. Ecology as a
predictive science is still in its infancy. There have been remarkably few planned and
controlled experiments at the ecosystem level to learn from, and to test the ecologist's ability to
accurately predict changes.

Nevertheless, marine ecologists can certainly observe present conditions in coastal ecosystems,
and they have observed what are presumed to be nutrient-related changes in estuaries.
Certainly some insights have been gained as to the behavior of coastal ecosystems under
increased nutrient loadings.

This review will consider relationships between nutrient loading and phytoplankton production
in coastal ecosystems. Specifically, the review will examine if there is a simple relationship
between nutrient loading and phytoplankton production, or surrogates for production, in
coastal ecosystems. In other words, how well can we predict how much change will occur in a
coastal ecosystem for any specific change in nutrient loading?

DEFINING TERMS

Before starting on any dialogue it is useful to make sure that participants are using the same
meanings for their vocabulary. The word "eutrophication" has had a variety of uses when
applied to coastal ecosystems, and probably many more uses when applied to other
ecosystems. For this review the definition of eutrophication as proposed by Nixon (1994) will
be used. His proposed definition is:

Eutrophication (noun) -- an increase in the rate of supply of organic matter to an ecosystem.

The rate of supply of organic carbon may be@ by either primary production by autotrophs within
the system (autochthonous carbon) or by an input of organic matter from outside the system
(allochthonous carbon). The central feature of the definition is the rate of supply of organic
carbon. At first consideration, the definition seems distant from the common informal use of
"eutrophication" to denote an increase in anthropogenic nutrient loading (i.e., cultural
eutrophication). However, use of Nixon's definition has considerable advantages.

First, the definition is simple. The definition is based upon a single parameter. There is a lot
to be said for simplicity.

Second, the definition makes it easy to separate eutrophication from its causes and its
consequences. The causes of eutrophication may include increased input of inorganic
nutrients, decrease in water turbidity, change in hydraulic residence time of the water, a decline
in grazing pressure or increase in direct inputs of organic matter. The consequences of an
increased rate of supply of organic carbon may include changes in ecosystem community
structure, increased rates of oxygen depletion, fish kills, etc.

Finally, focusing on the rate of organic carbon supply as the central issue makes it a
straightforward matter to classify the trophic status of coastal ocean ecosystems. The trophic
status of a coastal ecosystem is simply a measure of the rate of organic carbon supply.
Definitions of various trophic states were also proposed by Nixon (1994):

Organic Carbon Supply
g c M-2 year- 1,
oligotrophic <_100
mesotrophic 101-300
eutrophic 301-500
hypertrophic >500

The trophic classification states are value neutral. The definition does not imply that one
trophic state is better, more desirable, or more valuable than another.

APPROACHES TO FINDING A RELATIONSHIP BETWEEN EUTROPHICATION AND
NUTRIENT INPUTS.

In order to quantify "what will happen in a coastal ecosystem if nutrient inputs go up or
down," it is necessary to have or to establish a relationship between nutrient loading and rate of
carbon supply. Carbon supply can be from two sources, primary production within the
ecosystem and external loadings to the ecosystem. External loadings are likely to be the
dominant source of carbon in the vicinity of sewage outfalls with primary or no treatment. In
such areas one would not expect to find a cause-effect relationship between nutrient loadings
and carbon supply (unless one is considering the processes and procedures used in sewage
treatment plants themselves).

For most open-water areas of coastal ecosystems, the carbon supply is likely dominated by
autotrophic primary production within the system. A very useful relationship to identify would
then be a relationship between nutrient loadings and primary production. Some care must be
taken to make sure that coastal systems, or parts of coastal systems, with significant particulate
loadings are avoided in the development of a nutrient loading vs. production relationship from
empirical data.

A. number of approaches might be applied to the development of a relationship between nutrient
loading and primary production. These include:

1. Controlled experiments with marine ecosystems (mesocosm and enclosure experiments).

2. Historical records of the response of coastal ecosystems to changes in nutrient loads.

3. Cross-system comparisons of different coastal ecosystems to see if the primary production
in those systems is systematically related to nutrient inputs (comparative ecology).

4. Calculation (modeling) from first principles of physics, chemistry, and ecology.

Conducting controlled experiments with representative coastal ecosystems would be the most
direct way to determine the effects of nutrient loadings on coastal ecosystems. Such a direct
approach was instrumental in determining the response of lakes to nutrient loadings (i.e.,
Shindler et al., 197 1; Shindler 1973). Given that coastal ecosystems are considered public
domain and we are unlikely to want to deliberately damage such an ecosystem, it seems
unlikely that natural coastal ecosystems will be made available for direct experimentation.
Direct experiments are possible with coastal ecosystems in miniature. This is the realm of
enclosures and mesocosms. A variety of marine enclosures have been established to attempt to
create ecosystems in large containers that have chemical and biological properties of coastal
ecosystems. Descriptions of, and arguments for the use of, enclosures have been made by
Davies and Gamble (1979), Steele (1979), Parsons (1981), Lundgren (1985), and Santschi
(1988). It has been within the marine ecologist's ability for nearly two decades to establish
enclosures containing ecosystems that, over long periods of time (many months), have biota
and ecosystem function that are hard to tell apart from natural coastal ecosystems.

It seems remarkable that given this ability there has not been a widespread use of enclosures.
Direct experimentation is a powerful, if not the most powerful approach, in scientific
investigation. Although there have been a number of short term (i.e., up to about a month's
duration) nutrient experiments in marine enclosures, there has been only one long-term
eutrophication experiment in marine enclosures. There was a 28-month nutrient-addition
experiment at the Marine Ecosystems Research Laboratory of the University of Rhode Island
(Nixon et al., 1984; Kelly et al., 1985; Oviatt et al., 1986a, 1986b, 1989; Keller 1988, 1989;
Keller & Rice 1989; Hinga 1990). Results of that experiment will appear in the analyses later
in this paper.

One might expect it would be possible to learn a great deal from observations on the'changes
that have taken place in coastal ecosystems over time. However, the number of coastal
ecosystems where adequate measurements exist under two different loading conditions is
remarkably small. While current conditions and loadings are known in a good number of
coastal ecosystems, records of conditions under earlier loadings and records of earlier loadings
are relatively rare. Coastal ecosystems for which adequate data have appeared in the literature
will be considered below.

A comparative ecology approach was used by Volenweider (1976) to develop a quantitative
relationship between nutrient loadings in lakes and the abundance of lake phytoplankton. The
results of his comparative study demonstrated a clear relationship between phosphorus
loadings to lakes and the phytoplankton standing stock in lakes. Comparative approaches havc
been applied to marine ecosystems by Boynton et al. (1982), Nixon (1983), Nixon and Pilson
(1983) and Monbet (1992).

A modeling approach, while a very desirable goal, is likely out of reach for the near future.
Modeling the physics and chemistry of a coastal ecosystem on first principles may be currently
possible. Unfortunately, the first principles of ecology are not yet sufficiently understood to
permit useful predictioos. At the very least, an ecosystem model requires the ability to predict
when and why different species appear at various times. This ability is surely not yet achieved,
even in an unperturbed (by human activities) system. The ability to predict how communities
will change under novel conditions is even further off.

There are those who might argue that a functional model of the ecosystem is all that is needed
for prediction of at least some of the consequences of changes in nutrient loadings. In this
approach, specific species are not modeled. Related groups of species are assumed to have the
same function (nutrient dynamics, etc.). Functional modeling will certainly provide insights
into ecosystem functioning and perhaps eventually into effects of external changes on
ecosystems. Functional modeling is a valuable tool of the ecologist, but at present is more
likely to be useful as a diagnostic than a predictive tool. In order to use functional models to
provide predictions of response to change, it is necessary to know the coefficients, rates, and
equations that describe the functioning of the communities in the modeled system before and
after change in the parameters of interest. As noted above, it is not yet possible to confidently
predict which populations will comprise communities after a change in conditions. Unless one
can demonstrate that all coastal marine communities that could exist in a particular area are
functionally similar, or one can demonstrate how the functionality will change, it is
inappropriate to expect functional models to be reliable predictors of change.

Which of the possible approaches should be used to gain an understanding of the relationship
between nutrient loadings and the eutrophication of coastal ecosystems? The best answer is
probably "all of the above." When different independent approaches to the problem provide
the same results, then marine ecologists can start to have confidence that they have gained a
defensible ability to predict the response of coastal ecosystems to changes in nutrient loadings.

The history of lake eutrophication investigations provides a good example of the value of
having support from independent approaches. In the 1960's there was a great deal of
discussion and debate concerning the cause(s) of conspicuous thick algal blooms and algal-mat
growths appearing in many lakes. Were the mats and the blooms a result primarily of
phosphorus additions or of other agents? When the results of a direct experimental approach
(Shindler 1971, 1973) and a comparative approach (Vollenweider 1976) both clearly implicated
phosphorus as the primary causative agent, a firm basis was provided for the regulation of
phosphorus in detergents and other sources.

NITROGEN AND PRIMARY PRODUCTION IN MARINE ECOSYSTEMS

The nutrient central in discussions of coastal eutrophication is nitrogen. Early in the relatively
brief history of concern about marine eutrophication, Ryther and Dunstan (197 1) demonstrated
that additions of nitrogen stimulated growth of marine phytoplankton. Phosphorus additions
had little if any effect. Nutrient limitation studies have been conducted many times since (e.g.
Goldman, 1976; Laws and Redalje, 1979; Gran6li and SundbAck, 1985; D'Elia et al., 1986;
Gran6li et al., 1986; and Gran6li et al., 1988, Gran6li et al., 1990; Montgomery et al., 199 1;
Oviatt et al., 1994). In general, additions of nitrogen alone, as either nitrate or ammonium,
promote enhanced growth of phytoplankton (also see the review by Howarth, 1988).
However there are times when phosphorus limitation is found -in environments usually limited
by nitrogen (e.g. Gran6li etal., 1990) and marine environments where phosphorus limitation is
more common than nitrogen limitation (Myers and Iverson, 198 1). Silica limitation, resulting
from nitrogen and phosphorus enrichment, may also have an important influence on the
composition of phytoplankton communities.

With the general result that nitrogen is usually the limiting nutrient in coastal ecosystems, and
provided that long-term responses (i.e., yearly averages) are investigated, it seems reasonable
to attempt to find relationships between nitrogen and eutrophication, without consideration of
phosphorus. This assumption has been made in many previous studies, and will be adopted
here. Should it not be possible to find a satisfactory relationship(s) between nitrogen and
eutrophication, it may be necessary to retreat and consider phosphorus, and other factors.

PHYTOPLANKTON ABUNDANCE AND OTHER MEASURES WHICH MAY BE
RELATED To EUTROPHICATION

Primary production becomes the basis of the definition, as given above, of eutrophication in
coastal systems where external carbon loadings are not significant. Unfortunately, primary
production is not measured very regularly in many coastal ecosystems. It would be useful if
surrogate measures related to primary production based upon a more regularly measured
parameter could be found. This would expand the number of systems available for
investigation of nitrogen-eutrophication relationships.

The primary production of phytoplankton is dependent, in part, upon the abundance of the
phytoplankton. If phytoplankton is of low abundance, there are few "machines" to do the
work of production and the total production cannot be high. Hence phytoplankton abundance
and production may be expected to be somewhat related. Measurement of the standing stock of
phytoplankton is conducted much more often than measurement of production. The standing
stock of phytoplankton carbon is usually estimated by measuring chlorophyll-a and assuming a
fixed carbon to chlorophyll-a ratio.

The instantaneous (i.e., short-term) depth-integrated primary production (i.e., total production
under I m2 of surface) in marine systems can be estimated very well by knowing the
phytoplankton abundance, the depth of the photic zone (as calculated from the light extinction
coefficient), and the amount of incident light (Cole and Cloem, 1987; Keller, 1988, 1989).
Hence, if ambient light and turbidity were the same for all coastal ecosystems, chlorophyll-a
alone could be used to predict primary production.

Ambient light availability varies by latitude and local climate. Nixon (1983) assembled light
data for a variety of coastal embayments and found a modest range in annual average ambient
light ranging from 329 Ly day- I (Narragansett Bay, and New York Bay) to 461 Ly day- I (San
Francisco Bay). The effect of local climate was larger than the latitude effect (30 'N to 41 'N in
North America).

Light extinction also varies in coastal systems. Data also assembled by Nixon (1983) showed
attenuation coefficients ranging from about I m- I to 8 m- I with most of the high values at low
salinity. (This translates to I % light levels at about I to about 10 meters depth.) Of course, a
major determinant of light level in the water column is the amount of phytoplankton material
itself.

The abundance of phytoplankton is not always measured at multiple depths. It is more
common to find measurements of chlorophyll a, and of primary production made only on
near-surface samples. It is far less useful, from a whole ecosystem point of view, to consider
possible relationships between phytoplankton abundance and primary production for just near-
surface samples. It is the total carbon input to a coastal ecosystem that is relevant to
eutrophication as defined here. How well thetotal water column production can be estimated
from only near-surface measurements will not be considered in detail here.

If cross-system comparisons and consideration of changes in coastal ecosystems over time
were restricted to only those systems for which there is a good record of areal production, there
would not be much data to consider. In order to attempt to find relationships between nutrients
and eutrophication, measures of near-surface phytoplankton abundance, and near-surface
primary production will be considered. It will need to be kept in mind that mixing the various
types of measures of phytoplankton may obscure relationships for any single parameter.

OTHER COMPLICATING ISSUES

Phytoplankton abundance is often estimated from measurement of chlorophyll-a. It should be
noted that the carbon-chlorophyll ratio is not a constant (e.g., Falkowski, 1980). More
importantly, it appears that carbon to chlorophyll-a ratios vary systematically with
eutrophication (Nixon, 1992). The carbon to chlorophyll a ratio in the MERL eutrophication
experiment, referred to above, decreased by a factor of three from controls to the higher
enriched tanks. It seems probable that this is a result of shade adaptation and not a result of
changes in species composition (Oviatt et al., 1989). Such variability may interfere with
efforts to find a relationship between phytoplankton abundance and eutrophication.

Phytoplankton abundance may also be derived from direct counts of phytoplankton cells. A
change in cell count will not be directly proportional to change in biomass if there has been a
change in average cell size. In one of the ecosystems that will be considered later (Helgoland
Reede), cell counts were coupled with the volume per cell to provide a better estimate of
biomass change over time than raw cell counts.

Some comment must also be made concerning primary production measurements themselves.
There are a variety of basic techniques used to measure primary production and variability in
the practice of techniques based upon the same principle. Even the use of a single basic
technique, such as 14C-based production measurements, the depths of water samples, bottle
size and material, bottle agitation, and how bottles are incubated, are far from uniform in
practice. Such variability in technique undoubtedly affects results and may make comparisons
between different systems misleading.

NITROGEN LOADINGS OR NITROGEN CONCENTRATIONS?

Should a relationship be expected between eutrophication and nitrogen loading, or between
eutrophication and the achieved in-system nitrogen concentration (resulting from changes in
nitrogen loadings and the other processes affecting nitrogen concentrations)? A priori, there is
no way to determine which will be a more fruitful approach. Ideally, a description of the
availability of nitrogen is desired. It is not safe to assume that either inorganic nitrogen
concentrations or loadings of nitrogen adequately represent nitrogen availability.

Where measured nitrogen concentrations are used for a basis for a relationship, a long-term
average concentration of nitrogen must be used. In short time scales, the relationship between
nitrogen concentrations and phytoplankton abundance, may be inversely related, reflecting
short-term uptake by blooms. Further, consideration of just the dissolved species of nitrogen,
or dissolved inorganic nitrogen (DIN; nitrate plus nitrite, plus ammonia), may not represent the
real availability of nitrogen to the phytoplankton. Some portion of the particulate and dissolved
organic nitrogen may be subject to rapid remineralization and hence be biologically available.
If the fraction of the biologically available nitrogen in organic matter is variable in different
coastal ecosystems, use of DIN only may not provide clear relationships between nitrogen and
eutrophication.
If loadings are used as a basis for a relationship, do changes in loadings necessarily reflect a
corresponding change in availability of nitrogen to phytoplankton? Should loadings calculated
on a volume basis or loadings on an areal basis be used to represent availability? How does
flushing rate (or the inverse,,the residence time of water) affect the achieved availability of
nitrogen in the coastal ecosystem? The later question is addressed briefly as follows.

For a conservative (non-reactive) material introduced into an ideal well-mixed system,
loadings and achieved concentrations would be directly proportional. In a steady state

(constant loadings and concentration not increasing) a mass balance can be set up for a
conservative material where material entering the system must equal the material leaving the
system (or else the concentration of the material would either be rising or falling). Expressed
in mathematical shorthand:

Min = Mout

Min is the loading, at a constant rate, of the material M. (This assumes the offshore
concentration is negligible.) Mout depends upon how much water is exchanged per unit of
time between the system and more offshore waters.

Mout = volume of water lost per day * Concentration of M in water

[Following the units: g1day=1iters1day * g1liter]

As the flushing of a coastal system does not depend upon a material like nitrogen dissolved in
the water, the volume of water lost per day is constant (long-term average). Hence, in a single
ideal system, the achieved concentration of a conservative material is directly proportional to
the loadings. A doubling of a loading would lead to a doubling of achieved concentration,
once a new steady-state is achieved. (This requires a period of time equal to about three times
the residence time of the water in the system.) A similar exercise can be done for non-
negligible offshore concentrations, where the net result is that the net difference between the in-
system concentration and the offshore concentration is proportional to loading.

Nitrogen is not a conservative material. The forms of nitrogen undergo many reactions. As
nitrogen enters a coastal ecosystem from the landward side and passes through a coastal system
to the open ocean, it may be taken up from dissolved forms into organic matter and
remineralized (both in the water column and at the benthos) a number of times. In addition
there can be essentially permanent losses of nitrogen, through long-term burial in sediments
and denitrification (the conversion of fixed nitrogen to nitrogen gas). While there are very few
very well constrained nitrogen budgets for coastal ecosystems, it appears that burial is
generally a small term. Denitrification on the other hand may be quite significant and quite
variable. In a fast-flushing estuary, there may be little time for denitrification. In a system
where water has a long residence time, denitrification may account for a major loss. For
example, it is estimated than 90% of the nitrogen introduced into the Baltic Sea is denitrified
(Larsson et al., 1985).

The net result is that in coastal ecosystems with fairly short residence times, the achieved
concentrations of nitrogen in the system will be at least approximately proportional to changes
in nitrogen loadings. Even in systems where there is significant denitrification, if the fraction
of nitrogen lost to denitrification is approximately the same at different loadings, then the
proportionality between changes in concentration and changes in loading will hold.

For systems where there is a long residence time of water, it is probable that the relationship
between nitrogen loadings and concentrations will not be proportional. This is especially true
where the relative concentrations of the nitrogen species may have also changed over time. A
higher fraction of nitrate in the inputs (as may happen when sewage treatment. plants convert
more ammonia to nitrate to reduce the oxygen demand of their effluent) may be expected to
alter the fraction of nitrogen lost to denitrification. (Nitrate is the nitrogen species converted to
nitrogen gas during denitrificaion.)

RELATIONSHIPS BETWEEN NITROGEN AND EUTROPHICATION

There have been a few studies attempting to determine a general relationship between nitrogen
and eutrophication, or properties related to eutrophication. As might be expected from the
discussions above, a variety of different measures of nitrogen availability have been used.
Similarly, different measures of phytoplankton abundance and production have been
considered. The reader is advised to note the exact parameters plotted on each graph. Many of
the graphs appear similar but are of different parameters. In addition, some of the plots (those
generated new for this report) are provided in two versions showing the same data, a log-log-
axes version and a linear-axes version. The perspective obtained from inspection of plots may
be influenced by the log or linear construction of the plot. The dual versions are provided to
give multiple perspectives.

This section will look at some previous efforts to find relationships between nitrogen and
eutrophication from comparative studies and mesocosm experimentation. A new analysis of
the changes in coastal ecosystems over time will also be presented. In order to compare the
results obtained from the different measures of nitrogen availability and of phytoplankton
abundance and production, a factor-of-change of effect (abundance or production) per factor of
change in nitrogen (loading or DIN concentration) will be calculated. For example, it may be
found that in a system where the nitrogen loading doubled over time, the phytoplankton
abundance went up by a factor of 1.5. It may eventually be shown that it is inappropriate to
compare the results of abundance changes with production changes. For now, the approach
allows a quantitative look at eutrophication from a variety of types of information.

Nixon (1992) prepared plots of primary production and chlorophyll-a against DIN input (both
on an areal and volume basis) for a variety of marine systems ranging from the open ocean to
heavily nutrient-loaded estuaries (Figures I to 4). Both the production and abundance of
phytoplankton correlate with nitrogen loadings.

In viewing Figures I to 4 it should be kept in mind that the plots are log-log plots. Hence the
scatter of points is a significant magnitude. If a more limited range is examined, the
significance of the scatter becomes evident. Nixon and Pilson (1983) and Nixon (1983)
prepared cross-system plots for estuaries only. Their results are replotted in Figure 5. In these
graphs, the correlation between nitrogen and production is not so evident. Two estuaries with
the same DIN input rate may have up to a factor of five difference in production.

The results of two other cross-estuary approaches appear in Figures 6 and 7. Figure 6 shows
an effort similar to that of Nixon and Pilson (1983) except that the estuaries were broken into
segments (typically 4 segments), so as not to average high and low nutrient portions of
estuaries, and the segments were plotted individually. Nixon and Pilson (1983) used whole-
estuary averages. Figure 7 shows an annual average DIN vs. annual average chlorophyll plot
for individual stations in a variety of estuaries (from Monbet, 1992).

V;hile there may be a general trend of increasing phytoplankton abundance. with increasing
nitrogen availability, nitrogen alone is an imprecise predictor for a public official faced with a
decision as to whether the benefit will be worth the expense to reduce, for example, the
nitrogen inputs to an estuary by 30%. Estuaries, portions of estuaries, and individual stations
in estuaries with similar nitrogen loadings or DIN concentrations may have quite different (at
least from a practical management perspective) phytoplankton abundance and production.
From a historical perspective, examination of a portion, limited to high nutrient levels, of the
Vollenweider plot of corrected phosphate concentrations vs. chlorophyll (which was an

influential milestone in lake management), on a linear plot (Figure 8) shows scatter much like
that in Figures 5 and 6. While the general principle of phosphorus control on lake trophic
status was strongly supported by the original Vollenweider relationship, the relationship
established also did not have the precision to accurately (i.e., within a factor of 2) predict the
chlorophyll abundance in any individual lake.

The scatter in the cross estuary studies begs the question of how have individual coastal
ecosystems responded to changes in nitrogen. Will each individual coastal ecosystem respond
to nitrogen additions with an increase in phytoplankton abundance or production? If so, would
all ecosystems have the same magnitude of response for the same change in nitrogen loading or
concentration? Or, do coastal ecosystems of vary in sensitivity to changes in nitrogen?

What has been learned from controlled experiments? A 28-month eutrophication experiment
was conducted in the MERL marine enclosures using 9 enclosures. Three of the enclosures
were kept as controls and 6 enclosures received daily additions of nitrogen, phosphorus, and
silicate to create a gradient of loading. Figures 9, 10, and 11, show that measures of
phytoplankton abundance and production increase reasonably steadily with increasing nitrogen
loading or average concentration. The variability about the trend appears to result primarily
from influence of the benthic communities (which developed into quite different communities
in the different enclosures) on the phytoplankton in the water column. Nevertheless, at least
for this one environment, increases in nutrient loading resulted in eutrophication as defined
earlier.

What has been the record of changes in coastal ecosystems over time? Table I provides details
on a number of systems where a history of the conditions in the system is reasonably well
constrained. For most systems, the available parameters are phytoplanktori abundance (either
as directly-measured chlorophyll or as calculated chlorophyll from cell counts) and nitrogen
concentrations. The measured average abundance at two nitrogen concentration levels (usually
near the begining and end of the reported period of changes in nitrogen concentration) are
plotted in Figure 12. While many of the trends for individual systems combine to create an
overall trend consistant with those in Figures 1-4, is clear that the slopes of the trends are not
all parallel. For two of the systems, the change in abundance decreases with higher nitrogen
concentrations, running counter to the expected overall trend.

Additional systems and production data can be considered by calculating a normalized
parameter for each system. The parameter calculated here is the magnitude of change of
abundance or production that is calculated to result from a doubling in nitrogen concentration or
loading. A calculated "change factor" of 2, for example, indicates that there was a doubling in
abundance or production for a doubling in nitrogen loading or concentration. A change factor
of I would indicate that there was no change in abundance or production with a change in
nitrogen concentration or loading. A change factor of less than one indicates that there was a
decrease in abundance or production while there was an increase in nitrogen concentration or
loading. (See the legend for Figure 13 for details on the method of calculation.) The values
for the magnitude of the effect are plotted against the average DIN concentration for the system
(i.e., midpoint of the system's DIN concentrations).

Changes observed in different systems for a doubling in nitrogen ranged from a value of 0.60
to 6.99. The values 0.60 for a region of the Upper Chesapeake indicated that there was a
decrease in abundance of phytoplankton over a period in which there was increasing DIN
concentrations. Similarly, the value of 0.64 for the Upper Pamlico, indicated that there was an
increase in abundance of phytoplankton over a time period in which there was decreasing DIN
concentrations. Most of the calculated change factors fall between about 1. 1 and 3. It is
interesting that for the three stations in the system for which there are measures of both
abundance and production, Kanaehoe Bay, the increase in production was, at two of the three

stations, far greater than the increase in abundance. The Kanaeohe Bay results are from a
relatively short-duration study after a sewage diversion and may suffer from averaging a
relatively short period (Table 1). There was no clear trend in the magnitude of response in
relation to the overall DIN concentrations in the various systems. However, the data do not
clearly rule out a trend.

lt@ is not possible to determine how much of the range in factors found in Figure 13 and slopes
,in'Figure 12 may result from inadequacies in the data and the use of different measures of
phytoplankton abundance and production. It i's discouraging to note that two studies of a
similar portion of Chesapeake Bay (Upper Chesapeake Bay and Chesapeake Region VI in
Table 1) which encompass different but overlapping time periods, provide very pictures of the .
response. of p.hytoplankton abundance to increases in nitrogen concentrations. Yet, it seems
i 'bable that the variability can be accounted for simply by scatter. It is unlikely a single
mpro
tude of response to nitrogen increases applies to all the Systems. The two systems with
running counter to the general trend are particularly telling. There are also systems
where, without any change in DIN, phytoplankton concentrations have increased. (Such
examples cannot be plotted on Figure 113 as the effect value would be infinity.) For example,
the. Bokniseck station in the Keil Bight had increases in phytoplankton abundance and
production over a period when there is no apparent increase in DIN (see Gerlach, 19.90).

THE ROLE OF ANOTHER FACTOR

Perhaps it is simply unrealistic to expect that nitrogen availability alone will be a precise
Predictor of (long@term) phytoplankton abundance or production in coastal ecosystems. At the
.yery least, phytoplankton abundance is also controlled by grazing. Phytoplankton production
Is also controlled by light availability that may in turn depend upon inorganic particles,
unrelated to nitrogen, suspended in the water. Coastal ecosystems are also subject to
anthropogenic inputs of other chemicals that may alter phytoplankton response to nitrogen.

Revisiting the data assembled by Monbet (1992), shown earlier in Figure 7, demonstrates that
anadditional, somewhat subtle, factor helps control the abundance of phytoplankton in coastal
ecosystems. Monbet (1992) divided the estuaries in his data set into macrotidal (tidal range >2
m) and rnicrotidal (tidal range < 2 in). The macrotidal and microtidal points fall into 2 nearly
distiinct'groups as shown in Figure 14. For the same concentration of nitrogen, macrotidal
estuaries generally support lower phytoplankton abundances than microtidal estuaries.

Presumably, the higher tidal range estuaries have greater mixing energies which in turn exhibits
some type of control on phytoplankton abundance. Two mechanisms can be readily
hypothesized. First, more rapid vertical mixing may keep individual phytoplankton cells out of
the photic zone a sufficient fraction of the time to slow overall growth. Second, more vertical
mixing may bring phytoplankton near to the benthos where they are subject to more rapid direct
grazing by benthic organisms. Cloem (1991 and personal communication) had previously
noted that phytoplankton abundance in Northern San Francisco Bay was correlated to the neap-
spring cycle of the tides.

Only one of the estuary segments shown in Figure 6 would be classified macrotidal (with a
tidal range of only 2.1 m) by Monbet's definition. Nevertheless, if the data in Figure 6 are
sorted into those segments with tidal ranges @!1.2 m and <1.2 m, the set with the higher tidal
ranges falls low relative to the set with the lower tidal range (Figure 15). Although these data
do not provide a strong test of the eff 'ect of tidal height (lacking data from systems with high
nitrogen concentrations and low tidal range), they are consistent with the trend found by
Monbet (1992).

CURVE FITS TO CROSS-SYSTEM AND MERL DATA AND COMPARISON TO THE
EFFECT FACTORS FROM CHANGES OVER TIME

Once the cross-estuary data is sorted into groups by tidal height, it is possible to calculate. the
magnitude of the change in abundance for a change in nitrogen for each group. This allows a
comparison of the cross-estuary data to changes over time data. Figure 14 shows the Monbet
(1992) data with power functions fit to the two groups of data. Of the types of equations for
which curve fits were tried (power, linear, log, and exponential) the power curve provided the
best combination of correlation coefficients and normal distribution of residuals for both the
Monbet (1992) data and the NIERL data. Figure 15 shows the MERL abundance data
(previously shown in Figure 9) fit with a power function. In Figure 16, a power function is fit
to the data from segments of estuaries with tidal ranges less than 1.2 in. Changes in abundance
expected from doublings in nitrogen concentration may be calculated directly from the fitted
curves. Both groups of Monbet (1992) data and the MERL experiment indicate that a doubling
of nitrogen concentrations will lead to approximately a 1.4 times increase in phytoplankton
abundance. The data from the estuary segments with tidal heights <1.2 in indicates an increase
of only 1. 1 times for a doubling of nitrogen concentrations (although the correlation coefficient
is rather low). Doubling values from the cross-estuary studies and the MERL experiment are
within the range of those found in changes over time in individual estuaries (Figure 13).

CONCLUSIONS

Cross-system comparisons, history of changes in coastal ecosystems, and experimentation all
support the general idea that if an individual coastal ecosystem is enriched with nitrogen it will,
probably have higher average abundances of phytoplankton and higher average phytoplankton
primary production.

Unfortunately, cross-system comparisons, based only upon nitrogen loading or
concentrations, do not provide a very precise predictor of the phytoplankton abundance or
production that will occur in individual systems. Other factors in addition to nitrogen need to
be taken into account. Tidal height in coastal systems has been shown to have a large influence
on the abundance of phytoplankton that will be supported by a given average level of DIN in
the system. It appears that factors that may influence phytoplankton in addition to nitrogen and
tidal height will need to be considered in order to be able to provide a more precise predictive
relationship for phytoplankton abundance and production.

The magnitude of the response for a doubling in nitrogen loading or average concentration is
most likely to result in an increase in abundance or production by a factor between 1.4 and 3.
The responses of most coastal ecosystems will probably fall within this range.

However, there are systems that have not responded in the "normal" manner to nitrogen
enrichment. Some coastal ecosystems have had changes in phytoplankton abundance without,
or in the opposite direction of, changes in nitrogen concentration, Some coastal ecosystems do
not follow along the trend of nitrogen vs. abundance or production curves, even when the
systems are sorted by tidal height.

There is, at present, no guarantee that in any individual system, reductions in nitrogen loadings
will result in proportionate reductions (i.e., between 1.4 and 3 for a doubling in nitrogen) in
phytoplankton abundance or production. There is no characteristic of estuaries identified here
which will identify coastal ecosystems that will have unusual responses to nitrogen loadings.

Table 1. Sources of information for changes in coastal ecosystems over time (for Figure 12). The factor change in [DIN] or loading is tthc
measured high nitrogen concentration to measured low concentration or loading for the system over the period of time considered.

Phytoplankton Annual
System Years Salinity Average [DIN Parameters Averages Sources
% timoles/liter and Measured JNJ or
factor change in Loading
WIN] or loading

upper Chesapeake Bay 1965-1979 5-10 79 Chlorophyll JNO) Price et al., 1985
2.Ox

Upper Chesapeake
Bay (2)
Region IV (3) 1960-1969 8-16 14 Chlorophyll tDINJ Harding, 1994
1980-1990 1.2x
Region V (4) 1960-1969 4-13 22 Chlorophyll [DIN] Harding, 1994
1980-1990 1.8x
Region VI (5) 1960-1969 0-10 44 Chlorophyll IDIN) Harding, 1994
1980-1990 2.0

Kanaoe Bay
Station OF 1976-1979 2.2 Chlorophyll and [DIN] Smith ct al., 1981 a, b
2.9x Primary
Production per
volume.
Station SE 1.976-1979 0.97 [DIN] Smith et al., 1981a, b
1.5x
Station CE 1976-1979 34-35 0.79 [DIN] Smith et al., 198 1 a, b
lAx

Tampa Bay 1972-1978 Johansson and Lewis, 1992, and
1985-1989 22-25 7.15 Chlorophyll Loading Y-N Johansson, Personal Communication
3.2x

Table 1, continued. Phytoplankton Annual
System Years Salinity % Average [DIN] Parameters Averages Sources
gmoles/liter and Measured [NJ or
factor change in Loading
[DIN] or loading

Eastern Oosterschelde 1980-1989 27-30 52 h1orophyll and [DIN] Wetsteyn and Kromkamp, 1994
(71re Netherlands) 2.lx Primary Nienhuis and Small, 1994
Production per
area

Helgoland Reed (North 196 2-1984 30-32 22.6 Cell volumes [DIN] Gerlach, 1990
Sea) 1.6x

Halsskov Rev (Belt 1975-1982 -20 7.2 Priamary [Y-N] (6) Nielsen and Ertebjerg, 1984
Sea, Baltic) 3.7x Production per
area

Gulf of Finland (Baltic) 1972-1985 6 22.6 Chlorophyll [Y-N] (6) Kononen and Niemi, 1984
1.3x Konenen,1988

Tolo Harbor 1978-1985 25 4.4 Cell Counts [DIN] Hodgkiss and Chan, 1983, 1987
5.Ox

Tolo Channel 32 1.9 Ce 11 Counts tDINI Hodgkiss and Chan, 1983, 1987
3.2x

Upper Pamlico 1972-1992 0.5-5 13.6 Chlorophyll [DIN] Stanley, Personal Communication
1.9x
I .Not defined, from discussion appers; to be total DIN.
2. Change was calculated as the average of 1960-1969 compared to the average of 1980-1989
3. Patuxent River to South River/Annapolis.
4. South River/Annapolis to Bay Bridge/Magothy River.
5. Bay Bridge/Magothy River to susquehanna Flats.
6. Not defined. W

Figure 1. From Nixon (1992). Labeled systems are: (1) Sargas@o Sea; (2) North Pacific
gyre; (3) Choptank River estuary; (4) the Patuxent estuary; (5) Pamlico River estuary; (6)
Potomac River estuary; (7) main stem of the Chesapeake Bay; (8) Patapsco River estuary;
(9) Continental Shelf off New York; (10) Georges Bank; (11) Peru Upwelling; (13)
Delaware Bay; (14) Kaneohe Bay; (15) Baltic Sea. Open triangles are from the MERL
eutrophication experiment. For data sources and further information see Nixon (1992).

1000
0 141011r-@7
13 16

0 CIy 015b '14a

100
2b 014b
0
- ---------------
2a

cc L --------
a.
10 - I I I fill2 a I I till I i I I
0.1 10 10 101 101
DIN INPUT, mmol m-3 y I

Figure 2. Primary Production by phytoplankton (14C uptake) as a function of the annual
input of dissolved inorganic nitrogen per unit area of a wide range of marine ecosystems.
Systems are identified in Figure 1. The line represents a Redfield carbon to nitrogen
molar ratio of 6.625 and gives a rough indication of how much carbon would be fixed at a
given rate of nitrogen input without any recycling of nitrogen.

1000

Q -
7
10,P
0 13@6 0
A
Cr 5r A 9
E 14a- A4&
0 '@' 0.- 15 b
0 100 2b
14b
0
EC
CL 2a

10 -j
0.01 0.1 1 10 100
DIN INPUT, moles m- 2 Y- 1
5
/A

L4 64,N
115

2 b

Figure 3. Mean annual concentrations of phytoplankton chlorophyll as a function of the
annual input of dissolved inorganic nitrogen per unit volume of a wide range of marine
ecosystems. Systems are identified in Figure 1.

100

Cl)

tM
E
10 3@
Q

0 0

20 To
z 14b
------------

0.1
0.1 10 10, 10 3 ioA
DIN INPUT. mmol m-3 Y- I

Figure 4. Mean annual concentrations of phytoplankton chlorophyll as a function of the
annual input of dissolved inorganic nitrogen per unit area of a wide range of marine
ecosystems. Systems are identified in Figure 1.

100 =-
z

'E
07 5
E A6 A
- 10 7
4

2a 2
-j '0011
Cc

10
z
< 1 14 b
z

0.1 1 1 1 1 tillII II III
0.01 0.1 1 10 100
DIN INPUT, moles M-2 y-1

Figure S. Redrawn from Nixon (1983) and Nixon and Pilson (1983).
Coastal ecosystems represented are: Long Island Sound, Kaneohe Bay,
Chesapeake Bay, Narragansett Bay, Patuxent River, Apalachicola Bay,
South San Francisco Bay, North San Francisco Bay, and
Lower New York Bay.

500-

e4 400-

N 300-0

200

100

DIN input rate, ffirnol rn-3 yr-I

1000-

100

DIN input rate, mmol rn-3 yr"I

Figure 6. Unpublished data (compiled by K. Hinga, V. Berounsky, B. Kopp, A. Keller,
A. Desbonnet, S. Pavignano, D. Stanley, M. Pilson, S. Nixon, and V. Lee). Each point
represents a segment of an estuary. Estuaries were divided into typically four segments
and data averaged for the segment. Averages may include up to 10 years of data.
Estuaries represented are: Narragansett Bay, Delaware Inland Bays, Galveston Bay,
Tampa Bay, San Francisco Bay, Buzzards Bay, Long Island Sound, Delaware Bay,
Albemarle/Pamlico Bays, Barataria Bay, and Hudson River Estuary.

100

0 0
Z 10 - 00
49
MV

1 10 100 1000
Average DIN, "Aiter

25 -

41 0

00
5

0
0 40 80 120 160 200
Average DIN, gg/liter

Figure 7. Data from Monbet (1992).

100-
0
0
=L 0 0
0
0
0 OCPO 0 0
0 @s 6)0 0 0
0 (g (1 0 0 0 (9
08 0
0 00m(s 0 0
d 10- 00 0 0 0 0
0 0
>% 00 0 88 0 0 0
00 0
Cp C@90 000 0 0 opo- 0
(9 000 6p 00 0
0 0
8qgcg 0
0 0000V 00
0 00 0 9)

Annual Average DIN Concentration, gmoles I- I
100-

0
75-

0
50- 0
0 0 0
CPO 0 0
0
25- 0
0 0 0 0
0
0
�bp 0 0. 0
0

Annual Average DIN Concentration, gmoles I- I
10 EO
Cb

I Cl

Figure 8. Vollenweider (1976) data (estimated) for 22 lakes with high phosphorus
concentrations.

20
41

0 40 80 120 160 200
Average Phosphorus Concentration, g&4iter

Figure 9. Data from MERL eutrophication Experiment (Frithsen et al., 1985).

40
0

20
10 0

0 50 100 150 200 250 300 350 400
MERL Average DIN, pmoles/liter

100

1 10 100 1000
MERL Average DIN, pmoles/liter

Figure 10. Data From MERL Eutrophication Experiment (Frithsen et al., 1985).

100-

0 10-
E

CL
0
0

E
cc

100 1000 10000

Nitrogen addition rate
mmol/m3/yr

Figure 11. Data From M[ERL Eutrophication Experiment (Oviatt et al., 1986 and Keller,
1088). 14C production is from 14C uptake of water samples from the MEERL enclosures.
Net production is the result of whole-enclosure oxygen changes over a 24-hour period.

1000-

14C PRODUCTION

0 NET PRODUCTIVITY

E
co Net production, 28 month average
1 4C production, I year of data
100 - . . . . I . . . . .
100 1000 10000

Nitrogen addition rate
mmol/m3/yr

Figure 12a. Chlorophyll concentrations at two different nitrogen
concentrations for each system or station. See Table I for details
on. each system.

100,

to-

0.1
8

[DIN] or Total [N]

U. Chesapeake q Gulf of Finland

Chesapeake IV Helgoland

Chesapeake V Oosterschelde

Chesapeake VI Kaneohe, OF

U. Pamlico Kaneohe, SE

Tolo Harbor 10 Kaneohe, CE

Tolo channel

Figure 12b. Legend same as Figure 12a.

30-

25-

20-

15-

10-

[DIN] or Total [N]

U. Chesapeake Gulf of Finland

Chesapeake IV Helgoland

Chesapeake V Oosterschelde

Chesapeake VI Kaneohe, OF

U. Pamlico Kaneohe, SE

Tolo Harbor Kaneohe, CE

Tolo channel

27
Figure 13. For each system (See Table 1), the data for the "effect" (chlorophyll,
production, cell counts, or cell volumej was regressed against the nitrogen concentration
over the period of time when nitrogen and effects were changing. From the resultant
linear relationship between effect and nitrogen, the predicted effect level was calculated
for nitrogen values of 2/3 and 4/3 of the mean concentration for the system while the
change was taking place. Thefactor for change in effect is the ratio of the calculated
effects at 4/3 a d 2/3 of the mean nitrogen concentration. For all but four of the systems
there was an increase in nitrogen over time. Two of the systems, Tampa Bay and
Kanaehoe Bay, the change in nitrogen concentrations (and effect) were decreases
resulting from sewage diversion. In the Oosterschlede nitrogen concentrations decreased
as a result of a building of a storm-surge barrier. In the Pan-dico river, the nitrogen
concentrations were also going down over time.

4-
K

3- V
x
2-01
A
+

Average [DDfl

Upper Chesapeake Kanaeohe Bay. Chl

A Tampa Bay, Loadings Gulf of Finland

13 Helogland Reede Tolo Harbor

V Tolo Channel x Kanacoe Bay, Production

+ Halskov Rev Upper Pamlico

Ches IV 0 Ches V

Ches VI Oosterschelde. Chl

Oosterschelde, Prod

Figure 14. Data from Monbet (1992) fitted with a power function. Data is divided into
stations with tidal ranges >2 m (crosses) and <2 rn (open circles) and each set fit
separately.

100-

+ - >2m Chlorophyll Y = 0.626xo.121 r2 = 0.501
0
0 - <2m Chlorophyll y = 3.785xo- 491 r2 = 0.421
p. 75-

0
50- 0

0 0 0
+
C9 0 +
0 +
@U 25
0 +
+ +
+

Annual Average DIN Concentration, gmoles 1-1

100-

0
0
0
0 000 0 +
0 +
0 0 +
0 0
0 + ++
0 008 +
0 0 +
10- - 0
80 8 + + +
+
CP +++ + +++
&0 + + + +
0 + + ++ + +
+
+ 0 +
+ +
> ++ +
+ + +
0 ++ +
0 + +
++ @++ + +

++ +

Annual Average DIN Concentration, gmoles 1-1

Figure 15. Data shown in Figure 6 divided into two groups by tidal height in the
segment.

0 CHL Ave, jig/liter, Tide Range <1.2 m

0 CHL Ave, gg/liter, Tide Range 2:1.2 rn

10
0 0

0 0

0 40 80 120 160 200
AVG DIN (uM)

100

0
0
& 0
10 0 0
0 0

10 100 1000
AVG DIN WM)

Figure 16. Data from MERL eutrophication experiment fitted with a power function.

y 1.8683 xA(0.64976) R= 0.96035
100

10, 100 1000
MERL Average DIN, prnolestliter

Figure 17. Data from Figures 6 and 14 with a power function fit to the data with tidal
height <1.2 m.

-0- OHL Ave, gg/liter, Tide Range <1.2 m

0 OHL Ave, gg/liter, Tide Range @1.2 m

100
y 6.124 XA (0. 19155) R- 0.32712

0
0
IF 0
10 0 0
0 0

0
C)

10 100 1000
AVG DIN (uM)

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Kononen, K. (1988) Phytoplankton summer assemblages in relation to environmental factors
at the entrance to. the Gulf of Finland during 1972-1985. Kieler Meeresforschungen,
Sonderheft 7:281-294.
Kononen, K. and A. Niemi (1984) Long-term variation of the phytoplankton composition at
the entrance to the Gulf of Finland. Ophelia, supplementurn 3: 101-110.

Larsson, U., R. Elmgren, and F. Wulff (1985) Eutrophication and the Baltic Sea: Causes and
consequences. Ambio 14:9-14.

Laws, E. A. and D. G. Redaije (1979) Effect of sewage enrichment on the phytoplankton
population of a subtropical estuary. Pacific Science 33:129-148.

Lundgren, A. (1985) Model ecosystems as a tool in freshwater and marine research. Archiv
F. Hydrobiologie, Suppl.Bd. 70 2:157-196.

Monbet, Y. (1992) Control of phytoplankton biomass in estuaries: A comparative analysis of
microtidal and macrotidal estuaries. Estuaries 15:563-57 1.

Montgomery, R. T., B. F. McPherson, and E. E. Emmons (199 1) Effects of nitrogen and
phosphorus additions on phytoplankton productivity and chlorophyll a in a subtropical
estuary, Charlotte. Harbor, Florida. Report No. Water-Resources Investigations Report
91-4077.

Myers, V. B. and R. 1. Iverson (198 1) Phosphorus and nitrogen limited phytoplankton
productivity in northeastern Gulf of Mexico coastal estuaries. In: B. J. Neilson and L. E.
Cronin (ed.) Estuaries and Nutrients. Clifton, New Jersey, Humana Press.

Nielsen, A. and G. )Ertebjerg (1984) Plankton blooms in Danish waters. Ophelia,
Supplementurn 3:181-188.

Nienhuis, P. H. and A. C. Smaal (1994) The Oosterschelde estuary, a case-study of a
changing ecosystem; an introduction. Hydrobiologia 282/283:1-14.

Nixon, S. w. (1983) Estuarine Ecology- A Comparative and Experimantal Analysis Using 14
Estuaries and the MERL Microcosms. Graduate School of Oceanography, University of
Rhode Island, Final Report to EPA for Grant No. X-003259-01.

Nixon, S. W. (1992) Quantifying the relationship between nitrogen input and the productivity
of marine ecosystems. Advanced marine Technology Conference, No.5 Japan, pages 57-
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Nixon, S. W. (1994) Coastal marine eutrophication: some social causes and future concerns.
Ophelia , In Press.

Nixon, S. W. and M. E. Q. Pilson (1983) Nitrogen in estuarine and coastal marine
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Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P. Donaghay, B. Sullivan, S. Seitzinger, D.
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Oviatt, C. A., A. A. Keller, P. A. Sampou, and L. L. Beatty (1986a) Patterns of productivity
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Oviatt, C. A., D. T. Rudnick, A. A. Keller, P. A. Sampou, and G. T. Almquist (1986b) A
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Oviatt, C. A., P. Lane, 1. F. French, and P. Donaghay (1989) Phytoplankton species and
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Santschi, P. H. (1988) A need to study interactive effects. Applied Geochemistry 3:8 1.

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Stanley, D. W. (1994) Personal Communication.

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Vollenweider, R. A. (1976) Advances in defining critical loading levels for phosphorus in
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Wetsteyn, L. P. M. J. and J. C. Kromkamp (1994) Turbidity, nutrients, and phytoplankton
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MARINE EUTROPHICATION REVIEW

PART 2: BIBLIOGRAPHY WITH ABSTRACTS

TABLE OF CONTENTS

INTRODUCTION 1

ASSEMBLY OF THE BIBLIOGRAPHY 1

BIBLIOGRAPHY FORMAT
Subject Lists 2
Main Reference List 2

ELECTRONIC VERSIONS OF THE BIBLIOGRAPHY 2

RESOURCES USED IN THE BIBLIOGRAPHY (TABLES 1-4) 3

SUBJECT LISTS
Phytoplankton Abundance 6
Phytoplankton Composition 8
Red Tides And Nuisance Blooms 9
Macrophytes 9
Oxygen Concentration 10

MAIN REFERENCE LIST 11

APPENDIX: Keyword List 117

INTRODUCTION

Modem societies have increased the rate of supply of nutrients to estuaries and coastal marine environments
from many sources over the rates in the pre-industrial era and times of lower population density. The addition
of nutrients, or eutrophication, is considered a major problem in coastal areas. Typical symptoms attributed
to eutrophication are: increases in algal biomass and productivity, nuisance algal blooms, algal mats and
scums, increased bacterial activity, odor problems, turbidity, and depletion of oxygen in bottom waters. Long
term and secondary effects may include changes in the composition of pelagic and benthic communities,
reduction of species diversity, decreased abundance of submerged aquatic plants due to epiphetic or
planktonic shading, disruption of food chains, and the possible occurrence of toxic phytoplankton blooms
(such as "red tides"). On a positve note, a reasonable level of nutrient enrichment may increase primary
productivity without much negative effect and may be converted into increased abundance and yield of
desireable and harvestable species.

It is likely that all the consequences of estuarine eutrophication are not known. Even for those known effects,
a clear understanding of the magnitude of many of the effects resulting from increases in nutrient loadings is
lacking. Equally important, there are likely some effects attributed to nutrient additions to coastal
environments which are not actually caused by the increases in nutrients. This may occur because other
materials may be added to the marine environment, or there are other alterations of the environment which are
occurring simultaneously with nutrient additions.

In order to make a step toward a better understanding of the effects of eutrophication, the NOAA Coastal
Ocean Program commissioned the assembly of this bibliography on marine cutrophication. Literature which
may be useful in understanding marine eutrophication is in diverse sources. This bibliography is intended to
facilitate the finding of literature either for the scientist or manager who may want to develop a clearer picture
of marine eutrophication. The bibliography is not a compilation of data sources which might be used for new
analysis of eutrophication.

ASSEMBLY OF THE BIBLIOGRAPHY

The 438 references in this bibliography were assembled by searching a number of data bases, utilization of
symposia volumes concerning eutrophication, special journal volumes on eutrophication, and books on
nutrients in the marine environment. The data searched, and the years covered by the sources are listed in
Table 1. The keywords used for those searches are listed in Table 2.

The symposia volumes and books which were used for sources are listed in Table 3. Special journal volumes
are listed in Table 4. The judgement for including papers from these sources was based upon the content of
the individual papers. Our intent was to include papers which had research results or other insights
concerning marine eutrophication and not to include papers which simply mentioned the topic. Nor did we
attempt to compile a listing of data sources. There is not always a clear distinction between these two
categories and other preparers might have sorted papers somewhat differently.

Finally, papers which were listed in the references of the papers located by the above two techniques were
included where they met the criteria for inclusion.

BIBLIOGRAPHY FORMAT

Subject Indexes

There are five lists identifying papers contaning information in five major subject divisions. These are:

phytoplankton abundance
macrophytes
red tides and other nuisance blooms
phytoplankton community assemblage
oxygen concentrations

Each reference in these lists gives author and year. The complete reference is found in the main reference list
described below.

Main Reference List

The references are listed alphabetically by author. The year of publication appears next, followed by the title
of the work in bold text. The complete citation to the work appears next. Keywords highlighting the content
of the referenced work appear in brackets after the citation. Keywords supplied with the original work were
used when available. Keywords were provided for other references. Appendix A is a compiled list of the
keywords found in the bibliography. The complete abstract of the paper follows the citation unless there was
no abstract.

ELECTRONIC VERSIONS OF THE BIBLIOGRAPHY.

The primary version of the bibliography was assembled in Endnote PIUSTM software for the MacintoshTM.
The file is about 985 kilobites and is supplied upon request with the hard copy. ASCII versions of the main
reference list for both MacintoshTm and PCTM Compatible computers can be supplied upon request (about 500
kilobites).

RESOURCES USED IN THE BIBLIOGRAPHY

Table 1. Data Bases Utilized For Preparation Of Bibliography

Electronic Data Bases Years Covered
Aquatic Sciences and Fisheries Abstracts 1978-1992
Science Citation Index 1989-1991
NOAA Libraries Catalog
Government Documents Catalog Service 1976-1991
Paper Copy Abstracts
Aquatic Sciences and Fisheries Abstracts 1970-1977
Deep Sea Research 1970-1991
Dissertation Abstracts 1970-1990
Pollution Abstracts 1970-1991

Table 2. Words Used In Searches Of Electronic Databases

Abnormality Enrichment Organic Matter
Abundance Epifauna Pathology
Algae Epiphytes pH
Anoxia Estuaries Phosphate
Assemblage Eutrophication Phosphorus
Biomass Eutrophied Photosynthesis
Bivalves Fauna Pollution
Bloom Fish Population
Brown Tides Function Primary Production
Chlorophyll Growth Primary Productivity
Clams Hypoxia Red Tides
Coast Infauna Redox
Community Light Response
Coral Macroalgae Seagrass
Cycles Macrofauna Sewage
Deficiency Macrophyte Species Composition
Denitrification Meiofauna Standing Crop
Depletion Metabolism Structure
Diatom Microalgae Sublethal
Dinoflagellates Nitrate Submeraed Vascular Plants
Discharge Nitrification Sulfate
Diversity Nitrite Sulfur
Dynamics Nitrogen Turbidity
Eelgrass Number Waste
Effluents Nutrients Water Quality
Energy Organic Zooplankton
Enriched

Table 3. Symposia Volumes And Edited Books With Papers On Eutrophication
Which Were Used In The Assembly Of The Bibliography

on Nutrients and Eutrophication: The
Anderson, D. M., A. W. White, and D. G. Baden Limiting Nutrient Controversy, W.K. Kellog
(1985) Toxic Dinoflagellates. (Proceedings Biological Station, Michigan State
of The Third International Conference on University, I I - 12 February, 197 1).
Toxic Dinoflagellates, St. Andrews, New Lawrence, Kansas, American Society of
Brunswick, Canada 8-12 June, 1985). New Limnology and Oceanography and the Allen
York, Elsevier. Press.

Carpenter, E. J. and D. G. Capone (1983) LoCicero, V. R. (1975) Toxic Dinoflagellate
Nitrogen in the Marine Environment. New Blooms. (Proceedings of the First
York, Academic Press. International Conference on Toxic
Dinoflagellate Blooms, Boston, 4-6
Cosper, E. M., V. M. Brice1j, and E. J. Carpenter November, 1974) (MIT Sea Grant Report
(19 89) Novel Phytoplankton Blooms: MITSG 75-8). Wakefield, Massachusetts,
Causes and Impacts of Recurrent Brown Massachusetts Science & Technology
77des and other Unusual Blooms. (Papers Foundation.
from a conference at Stony Brook, NY, 27-28
October, 1988). Berlin, Springer-Verlag. Neilson, B. J. and L. E. Cronin (1981) Estuaries
and Nutrients. Clifton, New Jersey, Humana
Dennis, L. and H. H. Seliger (1979) Toxic Press.
Dinoflagellate Blooms. (Proceedings of the
Second International Conference on Toxic Okaichi, T., D. M. Anderson, and T. Nemoto
Dinoflagellate Blooms, Key Biscayne, (1988) Red Tides: Biology, Environmental
Florida, 31 October - 5 November, 1978). Science, and Toxicology. (Proceedings of the
North Holland, Elsevier. International Symposium on Red Tides,
Takamastu Perfecture, Japan, 10-14
Gran6li, E., B. Sundstr6m, L. Edler, and D. M. November, 1987). New York, Elsevier.
Anderson (1990) Toxic Marine
Phytoplankton. (Proceedings of the Fourth Smayda, T. J. and Y. Shimizu (1993) Toxic
International Conference on Toxic Marine Phytoplankton Blooms in the Sea.
Phytoplankton, Lund, Sweden, 26-30 June, (Proceedings of the Fifth International
1989). Now York, Elsevier. Conference on Toxic Marine Phytoplankton,
Newport, Rhode Island, U.S.A., 28 October
Hallegraeff, M. and J. L. Maclean (1989) Biology -I November 199 1). Amsterdam, Elsevier.
Epidemology and Management of
Pyrodinium Red Tides. (Proceedings of the Tyson, R. V. and T. H. Pearson (199 1) Modern
management and training workshop, Bandar and Ancient Continental SheyAnoxia.
Seri Begawan, Brunei Darussalam, Special Publication No. 58 of The Geological
Philippines, 23-30 May, 1989). Manila, Society. London, The Geolocical Society.
Fisheries Department, Ministry of
Development. Vollenweider, R.A., R. Marchetti, and R. Viviani
(1992) Marine Coastal Eutrophication.
Kennedy, V. S. (1984) The Estuary as a Filter. (Proceedings of an International conference,
Orlando, Academic Press. bologna, Italy, 21-24 march 1990. Reprinted
from the journal Science of the Total
Likens, G. E. (1972) Nutrients and Environment, Supplement, 1992) Amsterdam,
Eutrophication: The Limiting Nutrient Elsevier.
Controversy. (Proceedings of the Symposium

Table 4. Special journal volumes concerning eutrophication used in the assembly
of the bibliography

MarinePollution Bulletin Volume 23 (1991)
Ambio Volume 19, Issue 3 (1990) Special Issue: Environmental Management and Appropriate
Marine Eutrophication. Use of Enclosed Coastal Seas. (Proceedings of

the International Conference on the

Kieler Meeresforschungen, Sonderheft 6 (1988) Environmental Management of Enclosed Coastal
The Baltic Sea Environment, History- Seas'90, held in Kobe, Hyogo Prefecture, Japan,
Eutrophication-Recruitment-Ecotoxicology. 3-6 August, 1990).
(Proceedings of the 10th Symposium of the Baltic
Marine Biologists, 29 September - 3 October Netherlands Journal of Sea Research Volume 19,
1987). Issue 3&4 (1984). Nutrients in Estuaries:

Eutrophication and Other Aspects.
Marine Pollution Bulletin Volume 20, Issue 7 (Proceedings of the ES13AJHydrobiological
(1989) Pollution in the Far East. Society Symposium, Texel, The Netherlands, I I-
13 September, 1984).

6
SUBJECT INDEXES

SUBJECT INDEX: Phytoplankton Abundance

Words Searched for: abundance; bloom; chlorophyll; chl a; algal growth
(142 references)

Andersin, A.-B., J. Lassig, L. Parkkonen, and H. Gran6li, E., S. Schulz, U. Schiewer, D.
Sandler (1978) Gedziorwska, W. Kaiser, and M. Plinski
Anderson, D. M. (1987) (1988)
Anderson, D. M. (1992) Grassle, J. P. and J. F. Grassle (1984)
Arzul, G. and P. Gentien (1990) Gray, J. S. (1979)
Austen, M. C., R. M. Warwick, and M. C. Rosado Haas, L. W., S. J. Hastings, and K. L. Webb
(1989) (1981)
Beatty, L. L. (1991) Hagmeier, E. (1978)
Beukema, J. J. (1990) Hardy, J. T. and Z. Zubayli (1976)
Beukema, J. J. (1991) Heindnen, A. P. (199 1)
Beukema, J. J. and G. C. Cadde (1986) Hinga, K. R. (1992)
Beukema, J. J. and G. C. Cad6e (1991) Hobbie, J. E. and J. J. Cole (1984)
Birch, P. B. and J. 0. Gabrielson (1984) Hodgiciss, I. J. and B. S. S. Chan (1987)
Bodeanu, N. (1992) Howells, G. P. (1972)
Boni, L., E. Carpen6, D. Wynne, and M. Reti' Hull, S. C. (1987)
(1989) Hungspreugs, M., W. Utoomprkpom, S.
Brand, L. E., M. D. Gottfried, C. C. Baylon, and Dharmvanij, and P. Sompongchaiyakul
N. S. Romer (199 1) (1989)
Breuer, G. and W. Schramm (1988) Imai, I., S. Itakura, and K. Itoh (1991)
Brockmann, U., G. Billen, and W. W. C. Gieskes Jochem, F. and B. Babenerd (1989)
(1988) Johansson, J.O.R. and R.R. Lewis, 111 (1992)
Brockmann, U. and E. Dahl (1990) Josefson, A. B. (1987)
Brockmann, U. H., E. Dahl, and K. Eberlein Josefson, A. B. and R. Rosenberg (1988)
(1985) Kaiser, W., A. Irmisch, D. Nehring, F. Georgi,
Brown, J. R., R. J. Gowen, and D. S. McLusky and G. Bruel (1990)
(1987) Kat, M. (1987)
Cad6e, G. C. (1986) Kautsky, N., H. Kautsky, U. Kautsky, and M.
Cad6e, G. C. (1990) Waern (1986)
Cadde, G. C. (1992) Keller, A. A. (1989)
Cad6e, G. C. and J. Hegeman (199 1) Keller, A. A. and R. L. Rice (1989)
Cederwall, H. and R. Elmgren (1980) Keller, A. A. and R. L. Rice (1990)
Cosper, E. M., W. Dennison, A. Milligan, E. J. Kononen, K. and A. Niemi (1984)
Carpenter, C. Lee, J. Holzapfel, and L. Krbncke, 1. (1990)
Milanese (1989) Laws, E. A. and D. G. Reda1je (1979)
Cosper, E. M., C. Lee, and E. J. Carpenter (1990) Leppdkoski, E. (1980)
Costa, J. E. (1988) Maclean, J. L. (1984)
Dauer, D. M. and W. G. Conner (1980) Maclean, J. L. (1989)
Dauvin, J.-C. and F. Gentil (1989) Maestrini, S. Y. and E. Gran6li (1991)
Dzurica, S., C. Lee, E. M. Cosper, and E. J. Mattson, J. and 0. Lind6n (1983)
Carpenter (1989) McLusky, D. S., M. Teare, and P. Phizacklea
Emara, H.I., M.A. Shriadah, Th.H. Moustafa and (1980)
M.S. El-Deek (1992) Millner, R. S. (1980)
Fanuko, N. (1984) Mingazzini, M., A. Rinaldi, and g. Montanari
Friligos, N. (1985) (1992)
Gerlach, S. A. (1990) Morton, B. (1989)
Giovanaridi, F. and E. Tromellini (1992) Moshiri, G. A., N. G. Aumen, and W. G.
Gowen, R. J., P. Tett, and K. J. Jones (1992) Crumpton (1981)
Gran6h, E., P. Carlsson, P. Olsson, B. Sundstr6m, Murakami, A. (1973)
W. Gran6li, and 0. Lindahl (1989) Nielsen, A. and G. A@rtebjerg (1984)
Grandli, E., W. Graneli, and L. Rydberg (1986) Nishijima, T. and Y. Hata (1991)

Nixon, S. W., A E. Q. Pilson, C. A. Oviatt, P. Silva, E. S. (1985)
Donaghay, B. Sullivan, S. Seitzinger, D. Skjoldal, H. R. and 1. Dundas (1991)
Rudnick, and J. Frithsen (1984) Smayda, T. J. (1989)
Okaichi, T. (1987) Smayda, T. J. and A. W. White (1990)
Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst Soulsby, P. G., D. Lowthion, and M. Houston
(1984) (1982)
Olsson, G. E., B. Sundstr6m, and L. Edler (1987) Spencer, C. P. (1985)
Oviatt, C. A., P. Lane, F. F. 111, and P. Donaghay Stockner, J. G., D. D. Cliff, and K. R. S.
(1988) Shortreed (1979)
Paerl, H. W. (1988) Stockner, J. G. and K. S. Shortreed (1988)
Pastorok, R. A. and G. R. Bilyard (1985) Sukhanova, 1. N., M. V. Flint, G. Hilbaurn, V.
Pearson, T. H. (1980) Karamfilov, A. I. Kopylov, E. Matveea,
Pearson, T. H. (1987) T. N. Rat'kova, and A. F. Sazhin (1988)
Pearson, T. H., A. B. Josefson, and R. Rosenberg Takahashi, M. and N. Fukazawa (1982)
(1985) Takano,H.(1987)
Pearson, T. H. and R. Rosenberg (1976) Tamminen, T. (1990)
Pearson, T. H. and R. Rosenberg (1978) Taslakian, M. J. and J. T. Hardy (1976)
Pearson, T. H. and S. 0. Stanley (1979) Thompson, G. B. and J. Ho (198 1)
Pekkari, S. (1973) Tsijtsumi, H., T. Kikuchi, M. Tanaka, K. Imasaka,
Perciasepe, R. (1992) and M. Miyazaki (1991)
Persson, L.-E. (1987) Vidakovi'c, J. (1983)
Phillips, D. J. H. and S. Tanabe (1989) Vukadin, H. (1991)
Porumb, F. (1992) Walker, D. 1. and R. F. G. Ormond (1982)
Price, K. S., D. A. Flerner, J. L. Taft, G. B. Wallstr6m, K. (1988)
Mackieman, W. Nehlsen, R. B. Biggs, N. Wang, Z. (1987)
H. Burger, and D. A. Blaylock (1985) Weigelt, M. and H. Rumohr (1986)
Raman, A. V. and K. P. Prakash (1989) Weston, D. P. (1990)
Renk, H., J. Nakonieczny, and S. Ochocki (1988) Widbom, B. and R. Elmgren (1988)
Revelante, N. and A Gilmartin (1976) Wittberg, M. and W. Hunte (1992)
Ritz, D. A., M. E. Lewis, and M. Shen (1989) Wong, P. S. (1987)
Rosenberg, R. (1985) Wu, R. S. S. (1982)
Rosenberg, R., J. S. Gray, A. B. Josefson, and T. Wyatt, T. and I Horwood (1973)
H. Pearson (1987) Yang, D. B. (1987)
Schiewer, U., R. B6rner, and N. Wasmund (1988) Yi, S. K., J.-S. Hong, and J. H. Lee (1982)
Seliger, H. H. (1989) Zingone, A., M. Monitresor, and D. Marino
Sellner, K. G. and M. M. Olson (1985) (1990)

SUBJECT INDEX: Phytoplankton Composition

Words searched for: assemblage; community structure; species composition; species richness
(25 references)

Abdul-Hussein, M. M. and C. F. Mason (1988) Oviatt, C. A., P. Lane, F. F. 111, and P. Donaghay
Anderson, N. J., B. Rippey, and A. C. Stevenson (1988)
(1990) Pastorok, R. A. and G. R. Bilyard (1985)
Beauchamp, S. T. and J. Kerekes (1989) Schultz, S., G. Frtebjerg, G. Behrends, G. Breuel,
Bodeanu, N. (1992) P. Ciszewski, U. Horstmann, K.
Crema, R., A. Castelli, and D. Prevedelli (1991) Kononen, E. Kostrichkina, J. Leppdnen,
DeGroodt, E. G., H. F. J. Los, T. A. Nauta, A. A. F. Mohlenberg, 0. Sandstr6m, M.
Markus, and I. deVries (1992) Viitasalo, and T. Will6n (1992)
Dortch, Q., D. Milstead, N.N. Rabalais, S.e. Skjoldal, H. (1992)
Lorenz, D.G. Redaije, M.J. Dagg, M.E. Sukhanova, I. N., M. V. Flint, G. Hilbaum, V.
Turner, and T.E. Whitledge (1992) Karamfilov, A. 1. Kopylov, E. Matveea,
Gerlach, S. A. (1990) T. N. Rat'kova, and A. F. Sazhin (1988)
Grandli, E., 14. Persson, and L. Edler (1986) Swartz, R. C., F. A. Cole, D. W. Schults, and W.
Josefson, A. B. (1987) A. DeBen (1986)
Kononen, K. and A. Niemi (1984) Thompson, G. B. and J. Ho (198 1)
Kr6ncke, 1. (1990) Tomascik, T. and F. Sander (1987)
Mdkinen, A., 1. Haahtela, H. Ilvessalo, and J. Valente, R. M., D. C. Rhoads, J. D. Germano, and
Lehto (1984) V. J. Cabelli (1992)
Moore, C. G. and T. H. Pearson (1984) Wittberg, M. and W. Hunte (1992)
Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P.
Donaghay, B. Sullivan, S. Seitzinger, D.
Rudnick, and J. Frithsen (1984)

SUBJECT INDEX: Red Tides and Nuisance Blooms

Words searched for: red tide; brown tide; nuisance blooms
(38 references)

Maclean, J. L. (1989)
Anderson, D. M. (1992) Morton, B. (1989)
Arzul, G. and P. Gentien (1990) Murakami, A. (1973)
Aubert, M. (1992) Nishijima, T. and Y. Hata (1991)
Cho, C.-H. (1991) Nixon, S. W. (1989)
Cosper, E. M., W. Dennison, A. Milligan, E. J. Ochi, T. (1987)
Carpenter, C. Lee, J. Holzapfel, and L. Okaichi, T. (1987)
Milanese (1989) Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst
Cosper, E. M., C. Lee, and E. J. Carpenter (1990) (1984)
Doucette, G. J. (1989) Phillips, D. J. H. and S. Tanabe (1989)
Dzurica, S., C. Lee, E. M. Cosper, and E. J. Reguera, B. and Y. Oshima (1990)
Carpenter (1989) Seliger, H. H. (19.89)
Fiedler, P. C. (1982) Sellner, K. G. and M. M. Olson (1985)
Hodgkiss, 1. J. and B. S. S. Chan (1987) Silva, E. S. (1985)
Hungspreugs, M., W. Utoomprkporn, S. Smayda, T. J. (1992)
Dharmvanij, and P. Sompongchaiyakul Sukhanova, I. N., M. V. Flint, G. Hilbaurn, V.
(1989) Karamfilov, A. 1. Kopylov, E. Matveea,
Imai, I., S. Itakura, and K. ltoh (199 1) T. N. Raftova, and A. F. Sazhin (1988)
Jingzhong, Z., D. Liping, and Q. Baoping (1985) Takahashi, M. and N. Fukazawa (1982)
Keller, A. A. and R. L. Rice (1989) Wang, Z. (1987)
Keller, A. A. and R. L. Rice (1990) Wong, P. S. (1987)
Lam, C. W. Y. and K. C. Ho (1987) Wyatt, T. and J. Horwood (1973)
Liaci, L. S. (1980) Yang, D. B. (1987)
Maclean, J. L. (1984)

SUBJECT INDEX: Macrophytes

Words searched for: seaweeds; macroalgae.
(20 references)

Baden, S. P., L.-O. Loo, L. Pihl, and R. Mdkinen, A., I. Haahtela, H. Ilvessalo, and J.
Rosenberg (1990) Lehto (1984)
Birch, P. B. and J. 0. Gabrielson (1984) Menesguen, A. (1992)
Cheney, D. P. (1992) Pekkari, S. (1973)
Conolly, N. J. and E. A. Drew (1985) Quinlan, A. V. (1982)
Fletcher, R. L., V. Cuomo, and 1. Palomba (1990) Raffaelli, D., J. Limia, S. Hull, and S. Pont (1991)
Harlin, M. M. and B. Throne-Miller (198 1) Schramm, W., D. Abele, and G. Breuer (1988)
Hull, S. C. (1987) Smith, D. W. (1984)
Kautsky, N., H. Kautsky, U. Kautsky, and M. Soulsby, P. G., D. Lowthion, and H. A. C.
Waem (1986) Montgomery (1985)
Lee, V. and S. Olsen (1985) Tubbs, C. R. and J. M. Tubbs (1983)
Lenzi, M. (1992) Vogt, H. and W. Schramm (199 1)

SUBJECT INDEX: Oxygen Concentration

Words searched for: anoxia; hypoxia
(65 references)

Andersin, A.-B., J. Lassig, L. Parkkonen, and H. Oviatt, C. A., A. A. Keller, P. A. Sampou, and L.
Sandler (1978) L. Beatty (1986)
Andersson, L. and L. Rydberg (1988) Oviatt, C. A., J. G. Quinn, J. T. Maughan, J. T.
Aneer, G. (1987) Ellis, B. K. Sullivan, J. N. Gearing, P. J.
Baden, S. P., L.-O. Loo, L. Pihl, and R. Gearing, C. D. Hunt, P. A. Sampou, and
Rosenberg (1990) J. S. Latimer (1987)
Boesch, D. F. (1983) Parker, C. A. (1983)
Brockmann, U., G. Billen, and W. W. C. Gieskes Parker, C. A. and J. E. O'Reilly (199 1)
(1988) Pastorok, R. A. and G. R. Bilyard (1985)
Degobbis, D. (1989) Pearson, T. H. (1980)
Dethlefsen, V. and H. v. Westernhagen (1983) Perciasepe, R. (1992)
Dortch, Q., D. Milstead, N.N. Rabalais, S.e. Phillips, D. J. H. and S. Tanabe (1989)
Lorenz, D,G. Reda1je, M.J. Dagg, M.E. Phoel, W. C., K. L. Webb, and C. F. D'Elia (1981)
Turner, and T.E. Whitledge (1992) Polak, J. and G. D. Haffner (1978)
Emara, H.I., M.A. Shridah, Th.H. Moustafa, and Portnoy, J. W. (199 1)
M.S. El-Deek (1992) Price, K. S., D. A. Flemer, J. L. Taft, G. B.
Friligos, N. (1981) Mackieman, W. Nehlsen, R. B. Biggs, N.
Gerlach, S. A. (1990) H. Burger, and D. A. Blaylock (1985)
Griggs, G. B. and T. S. Hopkins (1976) Rabalais, N. N. and D. F. Boesch (1986-1987)
Herdendorf, C. E. (1986) Rabelais, N. N., R. E. Turner, J. W.J. Wiseman,
Hognestad, P. T. (1987) and D. F. Boesch (1991)
Hosie, D. A., R. F. Uglow, L. Hagerman, T. Reys, E. and M. Merino (199 1)
Sondergaard, and W. Weile (1991) Riisgkd, H. U. and E. Poulsen (198 1)
Justi'c, D. (1987) Rosenberg, R. (1985)
Justi'c, D. (1991) Rosenberg, R., B. Hellman, and B. Johansson
K6hler, A. and F. H61zel (1980) (1991)
Lam, C. W. Y. and K. C. Ho (1987) Rosenberg, R. and L.-O. Loo (1988)
LeppAkoski, E. (1980) Rydberg, L., L. Edler, S. Floderus, and W.
LISS, L. 1. S. S. (1990) Gran6li (1990)
Malone, T. C. (1991) Sales, D., A. G6mez, and D. Cantero (1983)
Millner, R. S. (1980) Sampou, P. and C. A. Oviatt (1991)
Mills, C. A., S. I. Heaney, C. Butterwick, J. E. Sampou, P. A. (1989)
Corry, and J. M. Elliott (1990) Stachowitsch, M. (1991)
Morton, B. (1989) Tolmazin, D. (1985)
Nehring, D., S. Schulz, and W. Kaiser (1984) Topping, G. (1976)
Nehring, D. (1992) Van Es, F. B., V. A. M.A., L. A. Bouwman, and
Nilsson, P., B. J6nsson, I. L. Swanberg, and K. H. G. J. Schr6der (1980)
Sundbdck (199 1) Vollenweider, R.A., A. Rinaldi, and G. Montanari
Nixon, S. W. and V. M. Berounsky (1984) (1992)
Nixon, S. W., C. A. Oviatt, J. Garber, and V. Lee Weigelt, M. and H. Rumohr (1986)
(1976) Welsh, B. L. and F. C. Eller (1991)
Ochi, T. (1987) Whiteledge, T. E. (1985)
Okaichi, T. (1987) Wu, R. S. S. (1982)

MAIN REFERENCE LIST

Abdul-Hussein, M. M. and C. F. Mason (1988) The first records of "dead" bottoms in the
The phytoplankton community of a Bornholm basin are from 1948, when no
eutrophic reservoir. Hydrobiologia 169:265- macrofauna was recorded below 80 m. Records from
277. 1954 show that the deepest parts of the Eastern
lphytoplankton; reservior; community; pH; Gotland Basin and the deep area between Oland and
composition; assemblage) Gotland were devoid of macrofauna at that time, but
The dynamics of the phytoplankton community that the deep areas of the northernmost Baltic
of a eutrophic reservoir are described for a two year proper and the Gulf of Finland were still populated.
period. Fifty-eight species were recorded, 25 of The change continued and during the 1960s the
them common. Bacillariophyta dominated during communities dominated by lamellibrachs in the
the winter and early spring and Chlorophyta during Bornholm and Gda'nsk Deeps disappeared, and were
late spring, to be replaced by a bloom of subsequently replaced by polychaete communities.
Cyanophyta. The mean and peak biomass of These have been wiped out during periods of bad
phytoplankton was 8.6 mg 1-1 and 40.8 mg 1-1 in oxygen conditions, but quickly re-established when
1981, and 8.3 mg 1-1 and 37.6 mg 1-1 in 1982. conditions improved. The larnmelibrach
Temperature accounted for 67.3% and pH for 8% of community has not been restored. In the Northern
the variation in total phytoplankton biomass over Central Basin and the Gulf of Finland the
the two year period, using regression technique. depopulation of the deep bottoms probably began
Both horizontal and vertical patchiness, measured later, in the late 50's. In the 70s practically no
as an index of mean crowding, were recorded in the macrofauna has been recorded below the permanent
reservoir. Horizontal aggregations were associated halocline in the Central Basin (except the
with spring blooms of Chlorophyta and summer southernmost parts of it) and the Gulf of Finland.
blooms of Cyanophyta, while vertical aggregations During the 60s and 70s the area with periodically
were most marked during the summer bloom of unfavorable oxygen conditions has covered about
Cyanophyta. Concentrations of phytoplankton were 100,000 km2, which is 25% of the total area of the
influenced by wind, the prevailing southwesterly Baltic Sea.
wind accumulating algae in the northeasterly arm of
the reservoir during much of the year. Anderson, D. M. (1987) Toxic algal blooms
and red tides: A global perspective. In:
Andersin, A.-B., J. Lassig, L. Parkkonen, and H. T. Okaichi, D. M. Anderson and T. Nemoto (ed.)
Sandler (1978) The decline of macrofauna in Red Tides: Biology, Environmental Science, and
the deeper parts of Baltic proper and the Toxicology. Elsevier, New York.
Gulf of Finland. Kieler Meeresforschungen Itoxic; bloorn}
Sonderheft 4:23-30. The literature on toxic algal blooms and red tides
(abundance; benthic; macrofauna; oxygen; hypoxia; documents a global increase in the frequency,
Baltic; hydrogen sulfide) magnitude, and geographic extent of these events
An attempt is made to describe the large-scale over the last two decades. Some of this increase is
changes in the benthic soft bottom macrofauna in undoubtedly a result of the increased awareness and
the deep parts of the Bornholm Basin, the Gulf of analytical capabilities of the scientific community,
Gda'nsk, the Central Basin and the Gulf of Finland, but a strong correlation between the number of red
from the beginning of Baltic zoobenthos research to tides and the degree of coastal pollution or
the present day. The authors also try to correlate utilization of coastal waters for aquaculture argue
these changes with fluctuations in the oxygen that there are other contributing factors. It also
content and salinity in near-bottom water layer. The appears likely that toxic algal species have spread
paper surveys the literature and presents recent and within regions over spatial scales of hundreds of
earlier unpublished results. kilometers, moving with major water currents and
During the later part of last century and the first storms. Long distance transport of species across
decades of the twentieth century no area of the oceans may have occurred as well, but the evidence
Baltic Sea seemed to have been totally devoid of is not conclusive and the hypothesis controversial.
macrofauna. Unfortunately there are considerable
gaps in our knowledge of the time before the Anderson, D. M. (1992) Toxic dinoflagellate
middle of this century. The most striking decline blooms and red tides in New England and
has taken place, generally speaking, after the abroad: A biogeographic and physiologic
exceptionally great inflow in 1951-1952, and the perspective. 31st Annual Symposium of the
subsequent prolonged stagnation. Northeast Algal Society, Woods Hole.
Idinoflagellate; red tide; bloom; Alexandrium; Gulf
of Maine}

12
Toxic red tides of the dinoflagellate Alexandrium been reported from different parts of the estuary. In
fundyense have become persistent phenomena over this paper, we have calculated trends in nutrient and
the last two decades in the southern Gulf of Maine, oxygen concentrations within the surface and deep
having been common occurrences for hundreds of waters of the Kattegat and adjacent waters. This has
years further to the north. This paper will explore been done with available data for the past decades,
the bloom dynamics of this organism in the Gulf, with reference to nutrient supply and phytoplankton
as well as the physiological characteristics of the production.
cells that influence their toxin production. Oxygen concentrations within the deep water
Additional studies will be presented that relate to decreased from 4.58 to 4.08 ml 1-1 between 1971
the population biology of Alexandrium, both and 1982, indicating a 50% increase in oxygen
within the Gulf of Maine, and throughout the consumption. Concentrations of Tot-N, Tot-P and
world. Through a combination of traditional field inorganic nitrogen increased simultaneously, both
phytoplankton ecology, physical oceanographic in the surface water during the winter and in the
measurements, remote sensing, toxin analysis, deep-water during the summer. Changes in Tot-N
morphological analysis, and molecular studies, the and Tot-P were dominated by the Baltic water,
complex and fascinating characteristics of this while local supply to the Kattegat dominated the
important organism are gradually becoming known. changes in inorganic nitrogen. Increases in Tot-N
Topics to be emphasized include the long-distance and Tot-P suggest a successively increasing
transport and confinement of Alexandrium biomass.
populations within a coastal current that originates The importance of local nutrient supply to the
in the rivers of southern Maine; the potential Kattegat was studied comparing expected nutrient
impact of environmental conditions on the concentration within the surface water (due to
production of toxins in Alexandrium; the role of exchange with adjacent water) with actually
dormant cysts in the bloom dynamics and observed concentrations.
population distribution of Alexandrium; the
possible global dispersal of Alexandrium Aneer, G. (1985) Some speculations about
populations via ship traffic; and forensic RFLP the Baltic herring (Clupea harengus
analysis of globally distributed Alexandrium membras) in connection with the
populations. eutrophication of the Baltic Sea. Canadian
Journal of Fisheries and Aquatic Science 4 2
Anderson, D. M., A. W. White, and D. G. Baden (suppl.1):83-90.
(ed.) (1985) Toxic Dinoflagellates JBaltic; herring; filamentous; algae; egg;
(Proceedings of The Third International mortalityl
Conference on Toxic Dinoflagellates, St. In this paper the hypothesis is put forward that
Andrews, New Brunswick, Canada 8-12 Baltic herring's (Clupea harengus membras)
June, 1985). Elsevier, New York. spawning time, spring or autumn, is determined by
No abstract feeding conditions during the adult phase and thus
not generally fixed. The present "absence" or
Anderson, N. J., B. Rippey, and A. C. Stevenson autumn spawners is thought to be the result of
(1990) Change to a diatom assemblage in improved feeding condition during the latest decades
* eutrophic lake following point source as a result of the eutrophication of the Baltic Sea.
* u t r i e n t re-direction: a During two spawning ground studies carried out in
palaeolimmnological approach. Freshwater 1978 and 1982 unusually high mortality rates were
Biology 23:205-217. noted for eggs in situ. In 1982, during 4 week close
{diatom; assemblage; lake; nutrient} to peak of spawning, the mortlaity increased
No abstract. substantially, especially for eggs among
filamentous algae. A significant difference was
Andersson, L. and L. Rydberg (1988) Trends in noted between eggs on coarser algae and those
nutrient and oxygen conditions within among filamentous algae (p < 0.001). During this
Kattegat: Effects of local nutrient period the average mortalities were 33 and 75%,
supply. Estuarine, Coastal and Shelf Science respectively. Very low levels of oxygen were
26:559-579. measured at night among the filamentous algae. An
Initrogen; oxygen; phosphorus; nutrient; Baltic} increase in the amounts of this type of algae as a
The Kattegat forms the outer part of the Baltic response to eutrophication might constitute a new
estuary. It is characterized by a stable two-layer hazard to the reproductive success of the Baltic
stratification maintained by approximately equal herring.
supplies of low saline water from the Baltic and
high saline oceanic water from the Skagerrak. The Aneer, G. (1987) High natural mortality of
nutrient supply to these waters increased rapidly Baltic herring (Clupea harengus) eggs
during the past decades and oxygen deficits have

caused by algal exydates? Marine Biology
94:163-169. Anttila, R. (1973) Effect of sewage on the
JBaltic; herring; egg; toxic algal exudates; oxygen) fish fauna in the Helsinki area. OIKOS
Results from field studies in 1978 and 1982 Supplementum 15:226-229.
together with results from a laboratory study carried (sewage; fish; species)
out in 1984 indicate that the unusually high natural The results given are based on fishery studies
mortality of Baltic herring (Clupea harengus L.) begun by the City of Helsinki in 1969 on the
eggs observed in situ is presumably caused by toxic effects of sewage on the fish fauna and fishing in
exudates released by filamentous brown algae the Helsinki sea area. The main research methods
during a limited period which happens to coincide are test fishing with a series of nets, and fishery
with the peak of spawning. Oxygen deficiency is inquiries.
no longer believed to be the major cause of the Sewage has brought about considerable changes in
observed unnaturally high mortality. It cannot, the quality and quantity of fish fauna in the
however, be entirely ruled out. Helsinki sea area. The changes in the fish fauna are
similar to those generally recognized as deriving
Ansari, Z. A., B. S. Ingole, and A. H. Parulekar from sewage. The fish species susceptible to
(1986) Effect of high organic enrichment pollution have suffered a decline and/or moved to
of benthic polychaete population in an purer areas. Such species are brown trout Salmo
estuary. Marine Pollution Bulletin 17:361-365. trutta(L)., whitefish Coregonus lavaretus (L.),
(Benthic; polychaete; productivity; diversity; burbot Lota lota (L.) and in Leusiscus idus (L.). Of
biomass; density; sewagel the major edible fish, pikeperch Lucioperca
The benthic polychaete fauna of an estuarine lucioperca L. has withstood pollution best.
region receiving domestic sewage and wastes from a The fish biomass in the more eutrophic bays is
nearby fish landing jetty was compared to that of a considerable in size compared with pure areas. In
site having normal organic enrichment. The addition to the deterioration in the fish species
population density, biomass and species diversity structure there are also flavour defects in fish over a
were greater at the sewage receiving site. Some sea area of 3,000-4,000 hectares. Flavour defects in
species showed preference for sewage site and their fish are caused by the secondary phenomena of
numbers were significantly greater at the sewage sewage and oil that have entered the water. The
site while other species showed no differences. mercury and pesticides contents observed in fish
Total counts and the biomass were significantly have been relatively low.
higher at the site of organic enrichment. A
quantitatively enhanced polychaete population Arndt, H. (1988) Dynamics and production
indicates that input of high organic matter of a natural population of Brachionus
stimulate benthic productivity. plicatilis (Rotaria, Monogonota) in a
eutrophicated inner coastal water of the
A *ntonius, A. (1985) Coral diseases in the Baltic. Kieler Meeresforschungen, Sonderheft
Indo-Pacific: A first record. P.S.Z.N.I: 6:147-153.
Marine Ecology 6:197-218. frotifer; Brachionus; growth; zooplankton; Balticl
(coral; disease; Indo-Pacific; Red Sea; Philippines) The dynamic and production of a natural
The so-called "band" diseases of reef corals, the population of the rotifer Brachionus Plicatilis
White Band Disease (WBD) and the Black Band (O.F.MlOLLER) were studied by regular field
Disease (BBD), were observed in the Red Sea and in sampling at routine stations in the shallow Darss
the Philippines. Since they were previously known Zingst estuary, southern Baltic (3-7 %- S).
only from the western Atlantic, this is the first Investigations of the horizontal distribution
record for the Indo-Pacific region. WBD is neither revealed significant population growth during the
infectious nor contagious and the pathogens is summer months at salinities above 3 %o S and at
unknown. BBD is highly infectious and contagious those stations characterized by high eutrophication.
and is caused by the cyanophyte Phormidium The first individuals hatch from resting eggs in
corallyticum. Susceptibility to WBD occurs May. Significant reproduction occurs from June to
throughout the order Scleractinia, but the effects of September, when temperatures are above 15 'C.
the disease are most severe among Acroporidae. Mixing rates were highest during the exponential
Susceptibility to BBD seems to be restricted to growth phase. Instantaneous rates of growth, birth,
Faviidae, with Platygyra and Goniastrea species the and mortality were estimated. Mean P/B ratios for
most heavily afflicted. WBD cannot be influenced the growing season were high (around 0.7 d-I) and
by any parameter tested so far. BBD can be were in the range of values obtained from mass
enhanced by light and by water eutrophication, and cultures in the field under subtropical conditions. In
its advance can be stopped by antibiotics. An active Barther Bodden annual biomass production for 1982
WBD seems to be a precondition for the origin of
BBD. was 1.1 g fw m-3. In its natural habitat, B.

14
plicatilis serves as a food source for fish juveniles promoting or inhibitory telemediators (exocrines)
and the mysid Neomysis integer, The seasonal present a very important control mechanism, in
dynamics of the rotifer population were not particular as far as the species compositions and
significantly affected by predators. periodic successions of phytoplankton are
concerned. However, this mechanism also
Arts, G. H. P., G. V. d. Velide, J. G. M. Roelofs, ultimately influences the overall equilibria. We
and C. A. M. V. Swaay (1990) Successional must take into account many facts which show the
changes in the soft-water macrophyte importance of this last factor.
vegetation of (sub)Atlantic, sandy, Such massive, semi-monospecies dinolflagellate
lowland regions during this century. blooms are, almost without exception,
Freshwater Research 24-.287-294. environmentally harmful, many of them also
Ivegetation; macrophytes) leading to the toxicity of sea food and relevant
SUMMARY human pathology. However, in specific cases,
1. Considerable changes in macrophyte vegetation such as the DSP intoxications induced by a number
can be noticed in 146 originally soft waters, when of species of Dinophysis, even the slight increase
data on the recent aquatic vegetation are compared in population densities, e.g. >200 cells dm-3, may
with historical information from the period 1900- cause quite dramatic pathology among shellfish
60. Changes in nutrient status (N, P, and C) and consumers and, of course, also important economic
accumulation of organic material can be regarded as losses for mariculture activities.
the operative factors.
2. The processes observed in soft waters are Austen, M. C., R. M. Warwick, and M. C. Rosado
acidification, eutrophication and water hardening. (1989) Meiobenthic and macrobenthic
Which process dominates depends on the type of community structure along a putative
soft water. pollution gradient in Southern Portugal.
3. Acidification as well as eutrophication of water Marine Pollution Bulletin 20:398-405.
bodies may ultimately result in the total (benthos; Portugal; community; diversity;
disappearance of all aquatic macrophytes,, with the abundance; sewage)
exception of the floating-leafed nymphaeids Macro- and meiobenthos were sampled from Ria
Nymphaea alba (L.) and Nuphar lutea (L.) Sm. Formosa, Portugal along putative sewage pollution
Observed successional stages are described and gradients in summer 1987 and winter 1988. The
summarized. status of the benthos was assessed using
multidimensional scaling ordination, ABC plots,
,Arzul, G. and P. Gentien (1990) Chemical and a variety of univariate indices of community
typology of coastal sediments in relation structure. Meiobenthos in both seasons and
to red tides. In: E. Grandli, B. Sundstr6m, L. macrobenthos in winter appeared to respond to
Edler and D. M. Anderson (ed.) Toxic Marine sewage enrichment only in the immediate vicinity
Phytoplankton. Elsevier, New York. of the sewage outfalls and channels but ABC plots
[red tide; phytoplankton; bloom; organic matter; indicated that macrobenthic communities were
sediments; copper; nutrients; nitrogen; phosphorus; moderately stressed throughout the area in summer.
Alexandrium) In the summer, human digging for shellfish results
The study of different chemical parameters in considerable sediment turnover. Since meiofauna
characterizing the coastal sediments showed may be less affected by physical sediment
possible relationships between high levels of disturbance than macrofauna this may explain why
organic matter and total copper concentrations, and the macrofauna communities were apparently
"red tide" occurrences. Sediment extracts had toxic disturbed relative to the meiofauna even beyond the
effects on the growth of Alexandrium tarnarensis in influence of the sewage. Sampling meiofauna with
April, however, it was not observed in sediment the macrofauna significantly increased our
elutriated in July. understanding of mans impact in this area. There
was little correlation between macro- and
Aubert, M. (1992) Sanitary consequences of meiobenthos in diversity, abundances, number of
eutrophication and related ecological taxa and biomass and these idices gave no
disequilibria in the marine environment. indication of a organic enrichment gradient.
In: R. A. Vollenweider, Marchetti and R. Viviani
(ed.) Marine Coastal Eutrophication. Elsevier, Axelard, D. M., G. C. B. Poore, G. H. Arnott, J.
Amsterdam. Bauld, V. Brown, R. R. C. Edwards, and N. J.
Inuisance bloom, dinoflagelate, Dinophysis) Hickman (1981) The effects of treated
Considering the rates and dynamics of marine sewage discharge on the biota of Port
primary productivity it appears quite clear that Philip Bay, Vitoria, Australia. In: B. J.
besides the essential factors such as nutrient pools Neilson and L. E. Cronin (ed.) Estuaries and
and inputs of energy, the function of growth- Nutrients. Humana Press, Clifton, New Jersey.

15
fbacteria; nitrification; denitrification; sewage; (oxygen; hypoxia; seaweeds; macroalgae; increase;
microalgae; macrophytes; macrobenthos; decrease; macrofauna; lobster; fish; floating;
zooplankton; phytoplankton; Australia; diversity; mortality)
fish} The southern Kattegat is susceptible to
The Werribee sewage-treatment farm contributes eutrophication due to shallow mean depth (23m)
more than half of the total nitrogen and phosphorus and a strong halocline which reduces intrusion of
input to Port Philip Bay. This study attempted to oxygen to bottom waters. The effects of
determine the fate of these nutrients and their effect eutrophication were first observed in the area in the
on the biota of the Bay. This was addressed by autumn of 1980. Since then investigations on
comparing community composition, biomass, primary producers, fish, lobsters, and benthic
productivity, or process rates in the Werribee area infauna have been conducted to document the effects
of the Bay with that in Bay areas more remote from of eutrophication. Above the halocline in the
nutrient discharge. Laholm Bay a change in the macrophyte species
Rates of bacterial nitrification and denitrification from Fucus spp. to filamentous green algae has
were greatest in sediments closest to the sewage been observed. Mortality of benthic macrofauna,
discharge point. Up to 15 percent of the inorganic mainly bivalves, has been observed in most years
nitrogen discharge may be lost via sequential and seasons. The recruitment of flatfish has not
nitrification-denitrification in a 4 km2 area. been negatively affected. Around the halocline the
Epibenthic microalgal biomass and productivity benthic infauna were seriously affected by oxygen
were found to be five times greater at Werribee than deficiency. Below the halocline, fish disappeared
at a control station. The Werribee macrophyte and lobsters emerged from burrows when oxygen
community showed reduced species diversity, saturation declined below 40%. When oxygen
dominance of fast growing opportunistic species, saturation decreased below 15% Norway lobsters
loss of large brown algal species, and occasional were immobilized and their blood-pigment
algal blooms, as compared to communities in concentration decreased. Benthic infaunal species
nutrient-poor areas of the Bay. Classification emerged from the sediment. Lobsters died when
analysis revealed offshore and nearshore groups of saturation dropped to 10%, while many infaunal
macrobenthos in a 2 km2 area near Werribee; species tolerated levels of 7-5% for some weeks.
species diversity was greatest offshore. The Analysis of stomach content indicated that neither
Werribee offshore macrofauna was typical of that fish nor lobsters died from lack of food, but from
along the whole northwestern coast of the Bay. In hypoxia. Subsequent to the reoxygenation of the
summer, fish biomass at Werribee was equal to that bottom water during winter, flatfish and benthic
found at stations remote from sewage discharge, infauna recovered whereas cod and lobster
however, community composition differed. At population did not.
Werribee, the nearshore fish community was Beatty, L. L. (1991) The response of benthic
dominated by juveniles and small species, while suspension feeders and their grazing
offshore older and larger fish were more abundant. impact on phytoplankton in eutrophied
Phytoplankton productivity decreased with coastal ecosystems. Dissertation for Ph.D.,
increasing distance from sewage outfalls only The University of Rhode Island.
during summer. Nitrogen is probably the nutrient (benthos; MERL; nutrient; composition;
critical to phytoplankton biomass production in the
Bay but light and/or temperature may limit abundance; biomass)
productivity over much of the non-summer period. The eutrophication of coastal waters from land-
Baywide phytoplankton productivity was similar to derived nutrients affects all biological aspects of the
that of non-eutrophic coastal marine waters. estuarine ecosytem. Changes to the benthic
Zooplankton standing crop was low compared to community can be pronounced. Not only do
that for many estuaries and marine embayments. benthic species respond to nutrient-induced
Densities of zooplankton in the Werribee area were phytoplankton blooms, but these animals may
highly variable, and crustacean species found at have a subsequent impact on the system.
Werribee were also distributed throughout the Bay. A sixteen-month study was conducted with daily
These findings suggest that sewage discharge had nutrient loading. One of the nutrient levels included
affected benthic more than planktonic communities, enhanced silicon to promote diatom growth and
but that the measurable impact of the discharge is potentially lead to a healthier food chain. Cores
limited to a few hundred meters around the outfalls. were taken monthly to follow changes in benthic
species composition. Laboratory experiments were
Baden, S. P., L. 0. Loo, L. Pihl, and R. performed on five occasions from March to October
Rosenberg (1990) Effects of eutrophication to determined the role of benthic grazing in control
on benthic communities including fish: versus nutrient-enriched systems. Cores were taken
Swedish west coast. Ambio 19:113-122. from mesocosms and removal rates of natural
assemblages of phytoplankton were determined.

16
A two to five-fold increase in abundance and higher in the second year. The observed increases in
biomass was observed in enriched over control annual production between years in the colored
mesocosms. A shift in species composition lakes were largely due to changes in euphotic depth
included the rapid response of facultative resulting from variations in hydrology and DOC
suspension-feeding spinoid polychaetes, Polydora export from the lake catchments. Lower discharges
ligni and Streblospio benedicti, and amphipod, in the colored lakes in 1980 were accompanied by
Ampleisca abdita and Corophium sp., to lower DOC concentration in the clear water lake did
phytoplankton compared to the other nutrient not produce significant changes in water color,
systems, although differences were not statistically light extinction coefficient nor annual production
significant by the second summer. Zooplankton between year. Rates of primary production at light
initially responded to phytoplankton blooms but optimum (P-max) were consistently higher in the
adult benthic suspension feeders dominated the most colored, acidic lake indicating that relatively
grazer biomass within three months of nutrient high rates of autotrophic production will occur
additions. under acidic conditions if nutrient supply is
Benthic grazers could remove up to 75% of water maintained.
column phytoplankton per day. Rates were higher
in nutrient-enriched systems where animal biomass Becker, D. S., N. L. Gerrish, and K. K. Chew
was greater than in control closures. Phytoplankton (1981-1982) Influence of organic
>10 um and water temperatures over 15'C were enrichment on dernersal fishes. Coastal
also significant factors in observed feeding rates. Ocean Pollution Assessment News 1:50-52.
Benthic species successfully competed with (fish; organic enrichment; demersal; Puget Sound)
zooplankton for food resources; alttough their No abstract
weight-specific feeding rates are lower than those
for copepods, opportunistic characteristics of Bell, P. F. (1991) Status of eutrophication
spinoid polychaetes and amphipods and their ability in the Great Barrier Reef lagoon. Marine
to deposit feed when water column phytoplankton Pollution Bulletin 23:89-93.
is depleted contribute to their success. (lagoon; phytoplankton; nutrients)
The results of this study are most applicable to Historical data on the levels of nutrients and
shallow, well-mixed coastal areas. As water column phytoplankton in the GBR lagoon are reviewed.
depths decrease and/or food is more accessible to the The results indicate that background levels of P-
benthos, benthic-pelagic coupling increases in P04 and phytoplankton have increased significantly
importance. over the past 50-60 years and that the levels appear
be at or above the eutrophication threshold level for
Beauchamp, S. T. and J. Kerekes (1989) Effects coral reef waters. Other data indicate that river
of acidity and DOC on phytoplankton discharge probability has a major impact on the
community structure and production in nutrient status of the GBR lagoon, but other factors
three acid lakes (Nova Scotia). Water, Air, such the nitrogen-fixing blue-green alga,
and Soil Pollution 46:323-333. Trichodesmium, could also be important.
acidity; lakes; phytoplankton; assemblage) Trichodesmium has the ability to introduce large
Phytoplankton community structure varied amounts of new nitrogen and it appears that the
between the three lakes and between years within increased phosphorus levels could be driving its
lakes. The Beaverskin Lake phytoplankton growth. To-date little effort has been made to assess
community was dominated by cyanophytes and the impact of eutrophication on the coral reef
chlorophytes in the summer and chrysophytes in communities. Because the background nutrient
the winter. Kejimkujik Lake was dominated by levels are relatively high both run-off and sewage
bacillariophytes in the summer of 1979 but no discharges could have serious impacts on nearby
single group dominant in 1980 or 1981. coral reef communities. Tertiary treatment (i.e.
Pebbleoggitch Lake phytoplankton consisted nutrient removal) of sewage should be required for
mainly of chlorophytes in 1979 but low biomass all discharges in the vicinity of coral reefs and
and no dominant groups characterized this lake special precautions need to be exercised when
during the growth season of 1980. Daily integral designing run-off drainage systems.
planktonic primary production measured
simultaneously in the three lakes showed that in Berman, T. and Z. Dubinsky (1985) The
both years annual planktonic primary production autoecology of Peridinium cinctum fa.
was highest in the clear water lake, Beaverskin westii from Lake Kinneret. Verhandiunsen
Lake, which also had lower dissolved organic der Internationalen Verlaasbuchhandlune 22:2850-
carbon (DOC) concentration compared to the two 2854.
dystrophic lakes. In the clear water lake annual lautoecology; Peridinium; pH; temperature;
production was similar between years but in the dinoflagellate; Lake Kinneret; freshwaterl
two colored lakes annual production was 40% No abstract

character of the last three winter. The stock of
Berounsky, V. M. and S. W. Nixon (1985) mussels declined during 1990 to unprecedently low
Eutrophication and the rate of net levels. All intertidal mussel beds were completely
nitrification in a coastal marine destroyed by commercial fishermen to stock their
ecosystem. Estuarine, Coastal and Shelf Science subtidal plots. It is recommended that extensive
20:773-781. tidal-flat areas should be closed for such fishery
(nitrification; estuaries; nitrogen; nutrients) activities.
Rates of nitrification were calculated for four large
(13 m3) estuarine-based microcosms that had been Beukema, J. J. (1991) Changes in
subjected to inorganic nutrient enrichment. composition of bottom fauna of a tidal-
Calculated rates were based on two years of weekly flat area during a period of
nitrate and nitrite measurements and ranged from a eutrophication. Marine Biology 111:293-301.
maximum of 0-55 umol NO-2+3 produced I- I day- I lbenthos; faunal composition; tidal-flat; Dutch
in the control tank (no enrichment) to over 13 Wadden Sea; production; abundance; biomass)
During the last 20 yr the western half of the
umol NO-2+3 produced I-1day-l in the most Dutch Wadden Sea has undergone significant
enriched tank (receiving 18.6 umol NH4 I- I day- 1). eutrophication: concentrations of P and N
Almost all NO-2+3 production was pelagic, little compounds and planktonic algae have roughly
doubled, as has primary production. Though
was benthic. Net NO-3 production or net NO-2 oxygen levels are often low in summer, anoxic
production dominated the net nitrification rates areas are small and rare due to strong tidal mixing.
during different seasons. Good correlations were During the 1970 and 1990 period, macrozoobenthos
found between various oxidation rates and substrate was sampled annually at 15 stations at Balgzand, a
concentrations. The calculated net nitrite production 50-km2 tidal-flat area in the westernmost part of
rates were 10 to 1000 times higher than previously the Wadden Sea. Not only did the estimates of total
reported rates for open ocean systems, numbers, biomass, and production double during
demonstrating the potential importance of these two decades, but significant changes in the
nitrification to estuarine systems. composition of the benthic community were
observed, too: (1) the numerical proportion of
Beukema, J. J. (1990) Long-term and recent polychaetes increased at the expense of molluscs
changes in the benthic macrofauna living and crustaceans, (2) the overall mean weight per
on tidal flat in the western part of the individual of the macrozoobenthos decreased
Wadden Sea. Proceedings of the 7th (numbers of individuals of small-size species
International Wadden Sea Symposium, Ameland, increased more rapidly than those of large-sized
The Netherlands. species), and (3) though absolute numbers and
lbentbic; macrofauna; tidal flat; Wadden Sea; biomass of all feeding types increased, the share of
species abundance; biomass) carnivores declined and that of deposit feeders
The bottom fauna living on tidal flats in the increased; the proportion of suspension feeders
westernmost part of the Wadden Sea were followed showed little change. This study refers to true
quantitatively by biannual sampling at 15 fixed macrobenthos only (1-min sieve) and further
stations throughout the 1970-1990 period. During excludes two taxa (Corophium spp. and Hydrobia
this period, the following long-term changes were ulvae) which occasionaly exercised an undue
observed; influence on numbers. Mass mortalites caused by
1. biomass doubled and numerical abundance more low oxygen concentrations were of a small-scale
than doubled; as a consequence the size of a'mean' nature only. Total number of species fluctuated
macrobenthic animal declined; without a clear trend. As a consequence of the
2. the abundance of more than half of the species increasing numerical densities, trends in species
studied increased significantly; numbers were slightly increasing when expressed
3. among these species, small deposit-feeding per unit area and slightly decreasing when estimated
worms were over-represented and some species per 100 individuals (by rerefaction).
increased more than other species.
The above changes are attributed to the recent Beukema, J. J. and G. C. Cadde (1986)
eutrophication of the area (CADtE 1992). A Zoobenthos responses to eutrophication
favorable effect is the enhanced stock and of the Dutch Wadden Sea. Ophelia26:55-
productivity of the benthos. Detrimental effects 64.
(caused by prolonged periods of oxygen depletion) lbenthos; Dutch Wadden Sea; biomass; secondary
occurred incidentally and locally. recent (1988- production; abundancel
1990) changes include a marked increase in the During the last few decades nutrient
species which are known to be sensitive to low concentrations in the Dutch coastal water have
winter temperatures and are ascribed to the mild increased significantly. Values observed for rates of

18
primary production as well as for concentrations of (estuary; classification; susceptibilityl
chlo rophyll and for particulate organic matter No abstract
have been higher during recent years than estimates
from earlier periods. Thus more food will have been Birch, P. B. and J. 0. Gabrielson (1984)
available for herbivorous benthos during recent Cladophora growth in the Peel-Harvey
years. estuarine system following blooms of the
Both biomass and annual production of the cyanobacterium Nodularia spumigena.
macrozoobenthos living on the tidal flats in the Botanica Marina 27:17-2 1.
western part of the Wadden Sea doubled during the Ibloom; cyanobacteria; estuary; macroalgae;
1970 to 1984 period. More than half of the species Cladophora; Nodularia; seaweeds}
contributed to this increase. In the species studied In recent years severe blooms of the nitrogen-
in most detail, Macoma balthica, both reproductive fixing cyanobacterium Nodularia spumigena have
success and growth rate increased significantly occurred during spring and early summer in the
during the 15-year period. eutrophic Peel-Harvey Estuary. Following
Though a causal relationship between decomposition of these blooms there are blooms of
simultaneous increases of nutrient levels, primary Cladophora and other macroalgae, and more recently
production and secondary production cannot be of a benthic species of Oscillatoria. Data on
proved, such a relationship seems to be natural. So nitrogen and phosphorus in estuary water and in
far, obvious detrimental effects such as mass Cladophora tissue, combined with a decomposition
mortalities from lack of oxygen appear to be rare in study of Nodularia, indicate that large amounts of
the well-mixed Wadden Sea. nutrients are recycled from Nodularia to other
algae. Control of Nodularia blooms is now of
Beukema, J. J. and G. C. Cadde (1991) Growth primary importance not only because of serious
rates of the bivalve Macoma balthica in problems caused directly by it, but also because of
the Wadden Sea during a period of the nutrients made available upon its decay for
eutrophication: relationships with other nuisance algae and bacteria.
concentrations of pelagic diatoms and
flagellates. Marine Ecology Progress Series Birch, P. B., J. 0. Gabrielson, and K. S. Hamel
68:249-256. (1983) Decomposition of Cladophora.
lphytoplankton; bloom; flagellates; Wadden Sea; Botanica Marina 26:165-17 1.
diatom; bivalve; growth; Macoma; secondary jalgae; Cladophora; estuary; Australia;
productionj decomposition; nitrogen; phosphorus}
Probably as a consequence of eutrophication, both Cladophora aff. albida is a benthic algA which
the length of the annual phytoplankton blooms and grows in nuisance proportions in the shallow (2m)
the mean concentrations of flagellates (dominated Peel-Harvey estuarine system in Western Australia.
by Phaeocystis pouchetii) have increased in the It grows as small (1-3 cm) ball-like clumps in beds
western part of the Wadden Sea during the last I - 10 cm deep, the lower self-shaded sections of
decades. Planktonic diatoms, on the other hand, which are in various stages of decomposition.
have hardly increased, but fluctuate heavily from Samples of Cladophora (half of which were pre-
year to year. Among the benthic tidal-flat fauna, killed by freezing) were buried in a bed in nylon
which has increased in total numbers and biomass mesh for up to one year. Initially live Cladophora
during the last decades, the bivalve Macoma was very resistant to decay; only after two months
balthica was studies in detail. The 15 year (1974 to did significant decomposition begin, resulting in
1989) patterns of its annual growth and condition loss of 50-60% of dry weight, carbon and nitrogen
were compared with abundance patterns in losses were slower (20-30% in two weeks, 50%
phytoplankton components (during the growing after three months).
seasons of M. balthica). Fluctuations in M. It was estimated that up to 60% of the
balthica were found to parallel fluctuations of Cladophora pool of nitrogen and phosphorus could
planktonic diatoms but not those of P. pouchetii. It be available for recycling in one year. This reduces
is concluded that M. balthica responded by faster the dependence of Cladophora on external inputs to
growth and higher condition to better feeding maintain production.
conditions in years of high diatom abundance.
Diatoms rather than P. pouchetii or other Blackburn, S. I. and Y. Oshima (1989) Review
flagellates will be the most important food source of culture methods for Pyrodinium
for M. balthica. bahamense. In: G. M. Hallegraeff and J. L.
Maclean (ed.) Biology, Epidermiology and
Biggs, R. B., T. B. DeMoss, M. M. Carter, and E. Management of Pyrodinium Red Tides. Fisheries
L. Beasley (1989) Susceptibility of US Department, Ministry of Development, Manila.
estuaries to pollution. Review in Aquatic The dinoflagellate Pyrodinium bahamese, both
Sciences 1:189-207. var. bahamense and var. compressum, has been

19
successfully cultured in seawater-based media by developed between 1960-1988, almost 60% reached
several workers. These cultures were sometimes the greatest development during the years 1983-
kept for many years but all of them were eventually 1988. The intensity of the blooms, the growth of
lost. This tropical dinoflagellate requires the mass species number, and the increase in
temperatures of 24-300C, dilute nutrient media and density induced high levels of global photplankton.
soil extract, while a pH optimum of 8 has also Continually rising, the mean biomass for the
been observed. Recommendations for renewed Romanian Black Sea areas was ten ti;mes greated in
culture attempts are provided. 1983-1988 than in 1959-1963.
Boddeke, R. and P. Hagel (1991) Boesch, D. F. (1983) Implications of oxygen
Eutrophication of the North Sea depletion on the continental shelf of the
continental zone. Netherlands Institute for Northern Gulf of Mexico. Coastal Ocean
Fisheries Research, E:7. Pollution Assessment News 2:26-28.
[North Sea; shellfish; fishl foxygen; hypoxia; Gulf of Mexico; nutrientl
North Sea landings of demersal fish rose after No abstract
1963 to more than I millions tons from a rather
constant level of about 400,000 tons since 1909. Boni, L., E. Carpend, D. Wynne, and M. Reti
Landings of bivalves from the Dutch coastal zone (1989) Alkaline phosphatase activity in
increased greatly after 1950. From the data Protogonyaulax tamarensis. Journal of
presented, a structural change in productivity of the Plankton Research 11:879-885.
Southern and Central North Sea in the period 1950- (bloom; pH; alkaline phosphatase; indicator;
1985 as the result of eutrophication of the dinoflagellates)
continental coastal waters emerges as the most A non-toxic strain of the marine dinoflagellate
likely cause of the enhanced commercial Protogonyaulax tamarensis (= Gonyaulaz
production. Therefore, the continuing decrease of tamarensis) has been isolated from a bloom in the
the discharge of phosphate to the southern North Adriatic Sea, off the Emilia-Romagna coast.
Sea since the beginning of the eighties, is likely to Cultures of the cells were grown in the laboratory
have negative effects on the production of fish and in enriched seawater at various initial ambient
shellfish in the S-E North Sea. orthophosphate (Pi) concentrations, ranging from
0.3 to 40 uM. The growth rate varied from 0.3 to
Bodeanu, N. (1992) Algal blooms and 0.8 divisions day-1, depending on the Pi levels.
development of the main phytoplanktonic From measurements of kinetic parameters, the
species at the Romanian Black Sea binding of the artificial substrate p-
littoral in conditions of intensification nitrophenylphosphate to the P. tamarensis alkaline
of the eutrophication process. In: R. A. phosphatase was quite strong (KM = 50 uM).
Vollenweider, R. Marchetti and R. Viviani (ed.) Maximal activity was observed at pH 8.4, although
Marine Coastal Eutrophication. Elsevier, the pH-activity curve was broad, in contrast to that
Amsterdam. of other alkaline phosphatases. Protogonyaulax
Ispecies abundance, species composition, Black tamarensis alkaline phosphatase, measured over a
Sea) 24 h period, exhibited an apparent diurnal
The growth of mineral salts and organic matter in fluctuation in activity, in common with the
the western Black Sea, caused by intensification in enzyme from other dinoflagellates.
the 1980s and 1990s of anthropic activity in the
pontic basin, the Danube and other tributary rivers, Boni, L., L. Mancini, A. Milandri, R. Poletti, M.
determined the increase of the frequency and Pompei, and R. Viviani (1992) First cases of
magnitude of algal blooms with their detrimental diarrhoetic shellfish poisoning in the
consequences for ecosystems. More than 20 Northern Adriatic Sea. In: R. A.
monospecific algal blooms have occured in Vollenweider, R. Marchetti and R. Viviani (ed.)
Romanian coastal waters between 1983 and 1988. Marine Coastal Eutrophication. Elsevier,
Of the eight species responsible for the Amsterdam.
phenomenon, five achieved the greatest blooms (Adriatic Sea, toxicity)
ever known at the Romanian littoral. Shellfish poisoning occurs when toxic
With the exception of algae which produce photoplankton species are present in their
monospecific blooms, remarkable development of environment. Because shellfish such as mussels
numerous other species has been recorded. The are filter feeders they concentrate toxins. It is also
number of species with high numerical densities well known that sometimes also small amounts of
(over 100,000 cells 1-1), increasing from one toxic algae can produce serious shellfish poisoning.
period to the next, reached 72 in 1983-1988, In the Adriatic Sea, dinoflagellate red tides are a
compared with 61 in 1971-1982 and only 38 in recurring phenomenon, but no shellfish poisoning
1960-1970. Of the total mass species which

has been described despite the presence of and chemical variables from 63 different estuarine
potentially toxic species such as Alexandrium spp. systems. Data were analyzed statistically to test
(Protogonyaulax) and Dinophysis spp. In June hypotheses regarding algal productivity and factors
1989 several cases of food poisoning characterized regulating temporal patterns. Prior to statistical
by vomiting, abdominal pain and diarrhoea were analysis, estuarine systems were classified into four
recorded along the coast of Emilia-Romangna and groups based on criteria of physical circulation and
Marche, facing the north-westem Adriatic Sea. The geomorphology. Analysis of grouped data indicated
poisoning was attributed to shellfish ingestion, that algal production and biomass were consistently
mostly mussels. Microscopic investigations high in warm periods of the year in a broad
revealed the presence of Dinophysis spp. in spectrum of estuaries and that ratios of available
seawater and mussel guts. Mytilus nitrogen to phosphorus were low during periods of
galloprovincialis, Venus gallina, Tapes high production, except in highly eutrophic
semidecussatus, and Venus verrucosa have been systems. In general, phytoplankton production and
monitored for toxin by mouse bioassay and biomass exhibited weak qprrelations with a variety
diarrhoetic shellfish poisoning (DSP) was detected of physical and chemical state variables, perhaps
only in mussels. Shellfish samples from Friuli indicating the significance of rate processes as
Venezia Giulia, Veneto, Emilia-Romagna and opposed to standing stocks in regulating these
Marche were monitored for toxin and from June 22 important features of estuarine systems. A six-year
a temporary prohibition on mussel fishing for the time series of measurements of algal production and
period of DSP was imposed. chlorophyll a at stations in middle Chesapeake Bay
exhibited considerable year-to-year variability, with
Borum, J. (1985) Development of epiphytic a three-fold range in peak values. Summertime
communities on eelgrass (Zostera marina) maxima were strongly related to annual loadings of
along a nutrient gradient in a Danish both nitrogen (N) and phosphorus (P) but annual
estuary. Marine Biology 87:211-218. production appeared to be sustained primarily on
feelgrass; epiphytes; community; estuary; recycled nitrogen and phosphorus. To generalize
biomass; phytoplankton; nutrients) from these findings, N and P loading rates were
The effect of nutrient enrichment on epiphyte estimated for 14 different estuarine systems, and a
development was examined by following the significant positive relationship was obtained
seasonal development of epiphyte biomass on between phytoplankton production and nitrogen
eelgrass (Zostera marina L.) at four localities along (but not phosphorus) inputs.
a nutrient gradient in Roskilde Fjord, Denmark
between March and December 1982. In the most Brand, L. E., M. D. Gottfried, C. C. Baylon, and
nutrient-poor area, epiphyte biomass followed a N. S. Romer (1991) Spatial and temporal
distinct bimodal seasonal pattern with maxima in distribution of phytoplankton in
spring and early fall. Low nutrient availability and Biscayne Bay, Florida. Bulletin of Marine
a high rate of eelgrass leaf renewal kept epiphyte Science 49:599-613.
biomass at a low level throughout the summer [phytoplankton; abundance; distribution; Florida;
period. Unlike phytoplankton, the epiphytic pigments)
community was not stimulated by nutrient Biscayne Bay provides a study in contrasts. North
enrichment during spring, however from May Biscayne Bay has undergone substantial changes
though August, the biomass of both components during the twenties century resulting from the
increased exponentially with increasing development of the metropolis of Miami around it
concentrations of total N in the water. Along the while south Biscayne Bay has been relatively little
nutrient gradient, phytoplankton biomass increased affected by the dramatic human population increase
5- to 10-fold, while epiphyte biomass increased SO- in South Florida.
to 100-fold. Thus differences in nutrient conditions The phytoplankton at 24 stations throughout
among study sites were more clearly reflected by Biscayne Bay were examined monthly for I year.
epiphytes than phytoplankton. Phytoplankton were found to be 5 to 10 times
higher in abundance in the north bay compared to
Boynton, W. R., W. M. Kemp, and C. W. Keefe the south and even more abundant near canal
(1982) A comparative analysis of mouth. Eutrophication from freshwater runoff is
nutrients and other factors influencing the likely cause of the higher abundance of
estuarine phytoplankton production. In: phytoplankton in the north bay as well as the
V. S. Kennedy (ed.) Estuarine Comparisons. higher fluorescence response indices and
Academic Press, New York. chlorophyll to phaeopigment ratios in the
(phytoplankton; productivity; estuary; biomass; phytoplankton in the north. Phytoplankton are
production; NIP ratio; nitrogen; phosphorus} most abundant in the north bay right after the first
We reviewed data concerning phytoplankton major rains of the rainy season, thought to be the
production, chlorophyll a, and associated physical result of the large amounts of nutrients flushed into

the bay at that time. Phytoplankton are least Pollution of the North Sea. An Assesssment.
abundant at the end of the dry season after a long Springer-Verlag, Berlin.
period of little runoff. Phytoplankton abundance JNorth Sea; nutrients; oxygen; effects; bloom;
shows virtually no seasonality in the south bay review)
where land runoff is relatively insignificant. The No abstract
south Biscayne Bay ecosystem remains dominated
by benthic macrophytes while the north bay shifted Brockmann, U. and E. Dahl (1990)
to an ecosystem dominated by a relatively dense Distribution of organic compounds
plankton community with a few macrophytes during a bloom of Chrysochromulina
remaining. polylepis in the Skagerrak. In: E. Gran6li,
B. Sundtr6m, L. Edler and D. M. Anderson (ed.)
Breuer, G. and W. Schramm (1988) Changes in Toxic Marine Phytoplankton. Elsevier, New York.
macroalgal vegetation of Kiel Bight [bloom; Chrysochromulina; phytoplankton;
(Western Baltic Sea) during the past 20 inorganic nutrients; organic; Skagerrak; Kattegatl
years. Kieler Meeresforschungen, Sonderheft Nutrient elements, bound in dissolved organic
6:241-255. substances, can become a major controlling factor
(Baltic; Kiel Bight; Furcellaria; Phyllophora; for algal growth when inorganic nutrients are
Phycodrys; abundance; biomass; coveragel depleted. During the late phase of the extended
A large scale quantitative survey of the benthic bloom of Chrysochromulina polylepis in the
vegetation of the Kiel Bight (Western Baltic Sea) Skagerrak/Kattegat during spring 1988,
has been performed by analysis of underwater concentrations of particulate and dissolved organic
television observations and samples obtained by substances (C, N P) were estimated together with
SCUBA diving during 1985-86. This investigation hydrographic measurements, and nutrient and
was compared to a serniquantitative survey carried phytoplankton analyses. Along a profile through
out in 1962-64 by Schwenke (1964, 1964). For the the Skagerrak between Skagen and HA116 a stable
total study (2571 km2), distinct changes in and most biologically rich pycnocline at 7-25 in
biomass and species composition have been depth separated two distinct chemical regimes: (i) a
observed. There is an increase of biomass above the surface layer where nitrate (<0.2 ug at N dm-3) and
12 in level (probably with exception of the 6 in phosphate (<0.02 ug at P dm-3) were nearly
level) and a decrease below 12 m. Extensive depleted and dissolved organic compounds (DOM)
Furcellaria lumbricalis populations have reached maximum concentrations (>10 ug at N dm-
disappeared. Furcellaria lumbricalis has been 3 and >0.05 ug at P dm-3), (ii) lower layers with
replaced by Phyllophora truncata and phycodrys higher inorganic nutrients and lower DOM. As a
rubens which are the predominant species at result of this, the pycnocline exhibited a minimum
present. Among other possible causes for these concentration of total dissolved nitrogen (<10 ug at
changes, the role of commercial stone fishing and
effects of increasing eutrophication in the Kiel N dm-3) together with maximum cell densities
Bight are discussed. (>16XI06 cells dm-3) of C. polylepis. Because of
these vertical nutrient profiles, the cells could
Brey, T. (1986) Increase in utilize both inorganic and organic bound nutrient
macrozoobenthos above the halocline in elements from the adjacent layers. The main N-
Kiel Bay comparing the 1960s with the source was however probably nitrate as seen from
1980s. Marine Ecology Progress Series 28:299- mixing curves. High concentrations of dissolved
302. carbohydrates (>3 umol Glc-eq dm-3) in the mixed
Imacrozoobenthos; Kiel Bay; biomass; mollusc) layer, significantly negative correlated with
The macrobenthos of sandy sediments in the 9 to salinity, indicate that the dominating source for
13 m water depth area of Kiel Bay was investigated. DOM was the Baltic outflow.
Data from 1961 to 1965 were compared with data
from 1982/83. Biomass of total macrobenthos was Brockmann, U. H., E. Dahl, and K. Eberlein
significantly higher in 1982/83 (14.6 g ash-free dry (1985) Nutrient dynamics during a
weight m-2) than in 1961 to 1965 (4.7 g AFDW Gyrodinium aureolum bloom. In: D. M.
M-2). This increase is mainly caused by mollusc Anderson, A. W. White and D. G. Baden (ed.)
species. Eutrophication, changes in predation Toxic Dinoflagellates. Elsevier, New York.
pressure and changes in physical disturbance are fnutrient; dinoflagellate; phytoplankton; blooml
discussed as possible reasons. Monospecies cultures of Gyrodinium aureolum
were investigated in two 16 and 20 in deep
Brockmann, U., G. Billen, and W. W. C. Gieskes enclosures for 20 and 22 days respectively.
(1988) North Sea nutrients and Nutrients were added at the beginning and after 10
eutrophication. In: W. Salomons, B. L. or 12 days, simulating coastal front mixing. In
Bayne, E. K. Duursma and U. F6rstner (ed.) spite of nutrient exhaustion in the upper layer

22
(nitrate below 0. 1 ug at N dm-3) the cells did not deep-basin samples; (111) sill samples; (IV)
migrate to the nutrient-sufficient lower layer. After Skagerrak samples. Characteristic species of the
two days of nutrient depletion, G. aureolum groups were identified with a pseudo F-test.
maintained its high cell number. However, the cells Detrended Correspondence Analysis indicated
decreased in the second enclosure during 5 days of seasonal faunal changes at 42 and 72 in depths. At
nutrient deficiency and did not increase following both depths a shift in numerical dominance from
the addition of nutrients, so that other species amphipods to mysids was found in winter, The
become dominant, including diatoms. most pronounced changes seemed to occur at 42 m.
Changes in composition and abundance of species
Brown, J. R., R. J. Gowen, and D. S. McLusky during a renewal of the bottom water indicated that
(1987) The effect of salmon farming on mysids were influenced by presumed near-bottom
the benthos of a Scottish sea loch. currents, while amphipods and decapods were not.
Journal of Experimental Marine Biology and Comparisons of the present amphipod fauna with
Ecology 109:39-5 1. the fauna in 1933-37 revealed significant
[benthos; abundance; biomass; fish farming; differences. The fauna in the deep basin was
diversity) impoverished and the Lilljeborgidae, earlier
. The effects of waste from a salmon farm on the represented by three species are most likely caused
benthos of a ljordic sea loch on the western coast of by eutrophication of the fjord.
Scotland have been studied. Within 3 in of the
floating cages the sediments was highly reducing, Cad6e, G. C. (1986) Increased phytoplankton
and dissolved oxygen content of the water overlying primary production in the Marsdiep area
the sediment ranged from 35 to 75% saturation. At (Western Dutch Wadden Sea). Netherlands
@!15 in from the cages the sediment was Journal of Sea Research 20:285-290.
oxygenated, and the dissolved oxygen content of the (phytoplankton; primary production; Dutch
water overlying the sediment was 50-80% Wadden Sea; phosphate; eutrophication}
saturation. Sedimentary redox potential and Annual phytoplankton primary production in the
dissolved oxygen content of bottom water showed a Marsdiep tidal inlet increased from ca 150 g C m-2
seasonal variation. The benthic fauna showed in the period 1964 to 1976, to ca 300 g C m-2 in
marked changes in species number, species 1981/1982 and 1985. This increase is considerable,
diversity, faunal abundance, and biomass in the but comparable to that observed in the outer Ems
region of the fish farm, with four zones of effect estuary, eastern Wadden Sea, from 240 g C m-2 a- I
identified. Directly beneath, and up to the edge of in 1972/1973 to 400-500 g C m-2 a-1 in
the cages, there was an azoic zone. A highly 1976/1980. Although the increase is most probably
enriched zone, dominated by Capitella capitata due to eutrophication, as illustrated by the regular
(Fabricius) and Scolelepis fuliginosa (Clapar@de), increase in phosphate in the Marsdiep area since
occurred from the edge of the cages out to =8 m. A 1950, it is difficult to explain why this affected
slightly enriched "transitional" zone occurred at !@25 primary production no earlier than the late
m, and a "clean" zone at the distances >25 m. This seventies. Primary production has probably not
study showed that salmon farming had similar increased in the more turbid inner parts of the
effects on the benthos as other forms of organic Wadden Sea, where light is the limiting factor, and
enrichment, but the effects were limited to a small P values were already higher than in the inlet areas.
area in the immediate vicinity of the cages.
Buhl-Jensen, L. and J. H. Fossa (1991) Cad6e, G. C. (1986) Recurrent and changing
Hyperbenthic crustacean fauna of the seasonal patterns in phytoplankton of the
Gullmarflord area (western Sweden): westernmost inlet of the Dutch Wadden
species richness, seasonal variation and Sea from 1969 to 1985. Marine Biology
long-term changes. Marine Biology109:245- 93:291-289.
258. (phytoplankton; abundance; Dutch Wadden Sea;
(benthic; fauna; change; amphipods; disappearancel distoms; flagellates; turbidityl
Distribution of 118 species of Amphipoda, Data for phytoplankton composition and
Mysidacea and Decapoda, sampled in 1984 and abundance in the Marsdiep are presented for the
1985 with an epibenthic sledge along a transect period from 1969 to 1985 inclusive. Only a few
from the Skagerrak to the inner Gullmarfjord species dominated the phytoplankton. A recurrent
(Sweden), is described. Amphipods were richest in pattern was observed in the seasonal succession: in
species in the Skagerrak, while most mysid species winter, total cell numbers were invariably low, but
were taken at the sill. Deca-impoverished and the freshwater algae, sluiced into the Wadden Sea from
Lilljeborgidae, earlier represented by three species, Ussel Lake, showed highest densities in winter. A
had disappeared since 1930s. The changes are most diatom spring peak was observed around mid-April,
likely caused by eutrophication of the fjord and followed by a Phaeocystis pouchetii peak about

23
three weeks later. Later in summer usually two lphytoplankton; trend; Marsdiep; Wadden Sea;
more diatom peaks followed by non-diatom peaks Phaeocystis; blooml
were present. The exact timing of the spring peak Published and unpublished data on phytoplankton
varied from year to year, with the extremes being of the Marsdiep tidal inlet were studied. Most older
late March and early May. A relatively late spring data going back to 1897, are based in net-
peak usually coincided with a relatively high phytoplankton only, the earliest quantitative
turbidity in the preceding winter. An increase in (Uterm6hl) data being from 1965. Phaeocystis sp.
total cell numbers was found over the 17-year bloomed in the Marsdiep after a spring diatom
observation period. Diatoms decreased from 1969 to peak, at least as long ago as 1897. Summer and
1974 but have increased since then, reaching values autumn peaks of Phaeocystis sp., frequent now,
above those of 1969 during recent years. Flagellates were also observed in 1898 and 1899. The duration
showed a consistent increase over the entire of the Phaeocystis blooms in 1897 to 1899 was
observation period. shorter than observed after 1978, but longer than
in the early 1970s. The recent (1987 to 1989)
Cad6e, G. C. (1990) Increased bloom. Nature duration of Phaeocystis blooms is 2 to 3 times that
346:418. of 1897-1899. The increase surpasses normal yearly
(phytoplankton; bloom; North Sea; Phaeocystis; variation and can be related to anthropogenically
nutrient) caused increase in nutrient concentrations. A
No abstract number of diatom species, at present numerically
dominant in the spring peak, are not mentioned as
Cad6e, G. C. (1992) Trends in Marsdiep dominant in the earlier periods of observation. They
phytoplankton. Proceedings of the 7th are small and passed through the nets used.
International Wadden Sea Symposium, Biddulphia sinensis, at present often abundant, is an
lphytoplankton; Marsdiep; diatom; bloom; immigrant in the North Sea since 1903, and for
phosphorus; nitrogen; North Sea; primary that reason absent from the earliest Marsdiep
production; composition; biomass) observations. No clear trend in duration of diatom
Phytoplankton cell-counts indicate an almost blooms is apparent during 1965 to 1989.
continuous increase in non-diatom plankton since Anthropogenic eutrophication did not affect diatom
the early 1970s. Particularly Phaeocystis blooms blooms. Marsdiep records in the literature of
increased in duration. Diatoms fluctuated more Phaeocystis globosa, P. pouchetii and P. sp. all
irregularly. Man-induced eutrophication - increased refer to the same species.
in phosphate and nitrogen, but not in silicate - can
explain such trends. They are comparable to those Caldwell, J. W. (1985) Effects of elevated
found in Helgoland, but different from (climate- turbidity and nutrients on the net
induced) trends observed in the open North Sea production of a tropical seagrass
during the Continuous Plankton Recorder survey. community. Dissertation for Ph.D., The
Such climatic effects are apparently overruled by University of Florida.
eutrophication effects in coastal waters. Historical [turbidity; nutrients; seagrass; dredging;
data on Phaeocystis for the Marsdiep area confirm production; shading)
the changes observed; they indicate blooming Dredging effects on seagrass communities in the
periods lasting some 50 days in 1897 and 1899, Florida Keys were examined by (1) comparing
before eutrophication started. This blooming period impacts on net production resulting from dredging
has tripled in the late 1980s. and natural weather events, (2) determining changes
Annual primary production was measured less in community photosynthetic efficiency, (3)
regularly, but the first 14C production data were evaluating shading and nutrient effects on net
collected in 1963-1965. Primary production has production, and (4) developing a systems dynamics
remained c. 150 g carbon (C) m-2 a- I up to the model.
early 1970s, but doubled in the early 1980s. Net community production was estimated during
Production in 1990 might be relatively low due to numerous meteorological and dredging events using
freshwater (including nutrient) discharge from the the Odum-Hoskins oxygen technique in flow-
Usselmeer. though field microcosms. In other experiments,
Chlorophyll-a showed an increase parallel with shading and nutrients (phosphorus, nitrate, and
the primary production increase in the late 1970s, ammonia) were manipulated to simulate dredge
but the annual variation is large. Comparison with plume conditions. The model examined the
the 1951-1953 data is hampered because of relationships between seagrass biomass, water
differences in the methods used. column and sediment nutrients, detritus, and
consumers.
Cadde, G. C. and J. Hegeman (1991) Historical The greatest depression in net community
phytoplankton data of the Marsdiep. production resulted from severe thunderstorms and
Hydrobiological Bulletin 24:111-118. dredging events, respectively. Net community

24
production measured two years after dredging
showed an approximate four-fold decreased. Carpenter, E. J. (1970) Effects of phosphorus
In field microcosm experiments, significant mining wastes on microorganisms of the
interaction occurred between shading and nutrient Palmico River estuary, North Carolina.
concentration. Significant metabolic reduction Dissertation for Ph.D., North Carolina State
occurred due to shading , even with higher nutrient University at Raleigh.
conditions. All but the lowest concentration (Palmico River estuary; blue-green algae; sewage;
resulted in significantly increased production in the phosphorus; mining; photosynthesis)
light. Qualitative comparison with a control To test the effects of phosphorus mining wastes
showed an enhancement of production only in the (SPW) and domestic sewage on microorganisms of
light. the Palmico River estuary, six 1400 m3 artificial
The model of seagrass production was most estuaries and eight 15.1 m3 static pools were
sensitive to changes in nutrient-seagrass constructed. Concentration, excluding controls, of
relationships, seagrass production estimates, and SPW in the pools was 1 percent in estuaries ranged
seagrass-light interactions. Recovery of seagrass from I percent to 51 percent of the total water
biomass following numerous dredging events (3.5 volume. The present day concentration of SPW in
years) was longer than that from estimated total the lower Palmico River is 0.7 percent (v/v) and is
annual thunderstorms encountered (I year) but expected to rise as other companies begin mining
shorter than recovery from hurricane events (4.1 operations. Three experiments were carried out, two
years). (54 and 44 days long) in the estuaries and one (15
The effects of short-term dredging on net days long) in the plastic pools.
community production as shown by field The experiments showed that the addition of SPW
experimentation and the model were less severe to estuarine water does not immediately increase the
than some weather events because of differences in photosynthesis or biomass of phytoplankton. This
duration and intensity. Lowered post-dredging is most likely because phosphorus does not limit
photosynthetic efficiency, as defined by unit the growth of phytoplankton in the river. As
biomass production per unit light, was a result of shown in a nutrient limitation study, nitrogen
degraded system function possibly from sediment limits algal photosynthesis in the Palmico estuary.
scour from the dredging cut. The major deleterious However, blue-green algae cell numbers (Anabaena
effect of dredging resulted from shading, although sp. and Spirulina sp.) were 77.2 percent higher
some production enhancement could occur from (.010<p<.025) than controls in estuaries containing
nutrient release. The model suggested that dredging approximately 13 percent SPW. Although cell
effects were prolonged because of current scour numbers were 33.2 percent above controls in
from the dredging cut, that recovery time was estuaries with 3 percent SPW, the difference was
comparable to hurricane events, and that more not significant at the 5 percent level. Also, blue-
investigation is needed in seagrass-nutrient and green algae (Anabaena torulosa) numbers were 404
light-production relationships. percent higher (.Ol<p<.025) than controls in
plastic pools with 1 percent SPW (v/v) and 295
Cambridge, M. L. and A. J. McComb (1984) percent above (.010<p<.025) controls with 1
The loss of seagrass in Cockburn Sound, percent SPW and 0. 1 percent (v/v) artificial sewage.
Western Australia. 1. The time, course The growth of blue-green algae was probably
and magnitude of seagrass decline in stimulated by the addition of phosphorus in the
relation to industrial development. SPW and their nitrogen fixing ability allowed then
Aquatic Botany 20:229-243. to obtain sufficient nitrogen in waters where this
Iseagrass; effluents; Australia) nutrient is limiting. This increase in blue-green
The areas of seagrass meadows in Cockburn algae may be potentially harmful to the estuary by
Sound, a marine embayment in Western Australia, changing the algal species composition and by
were estimated from historical aerial photographs adding fixed nitrogen to an estuary where
supplemented by ground surveys, studies on photosynthesis is nitrogen limited:
meadows in adjoining areas, and coring for rhizome An interaction occurred in the pools between
remains. Ten species of seagrass with different SPW and the artificial sewage mix which resulted
habitat tolerances are recorded for the area, with in photosynthesis 28.8 percent above the controls.
Posidonia sinuos Cambridge et Kuo forming the The existence of this interaction shows the
most extensive meadows. It is estimated that from importance of sewage treatment facilities for towns
1954 to 1978 the meadow area was reduced from in the Palmico River basin. Rate of phosphorus
some 4200 to 900 ha., based on leaf detritus uptake by microorganisms was 23.1 percent higher
production from 23000 to 4000 t (dry wt.) y- 1. The (p<0.005) than controls in plastic pools with I
major loss of seagrass occurred during a period of percent SPW and 74.9 percent higher (p<005) than
industrial development on the shore, and the controls in pools with both SPW and artificial
discharge of effluents rich in plant nutrients. sewage. This indicated that microorganisms are

taking up the added phosphorus in SPW. The constitute the first statistically confirmed biological
uptake in pools with both SPW and sewage was evidence of this eutrophication, and demonstrate
3.2 x that in the pools with SPW added alone. This that eutrophication is a major process affecting the
uptake may reflect the primary productivity entire Baltic Sea, and not just restricted coastal
interaction which occurred in the pools. No changes areas.
in the Vmax of glucose by bacteria was seen in the
studies. This was probably because the rate of Cederwall, H. and R. Elmgren (1990)
formation of glucose did not increase in an Biological effects of eutrophication in
experiment of such short duration. the Baltic Sea, particularly the coastal
zone. Ambio 19:109-112.
Carpenter, E. J. and D. G. Capone (ed.) (1983) [Baltic; review)
Nitrogen in the Marine Environment. The reported biological effects of the increased
Academic Press, New York. nutrient load on the Baltic Sea are summarized,
No abstract with some comparisons with the Kattegat and
Skagerrak. Interest is focused on the coastal zone,
Caspers, H. (1987) Changes in the benthos where effects are more obvious than in offshore
at a sewage-sludge dumpsite in the Elbe areas, but from which results have not often been
Estuary. In: J. M. Capuzzo and D. R. Kester published internationally. Reports demonstrate
(ed.) Oceanic Processes in Marine Pollution. Vol. 1. environmental degradation over extensive coastal
Biological Processes and Wastes in the Ocean. ar'eas of the Baltic Sea. Recorded effects include
Robert E. Krieger Publishing Company, Malabar, increased nutrient levels; increased algal blooms,
Florida. chlorophyll a concentrations, and primary
(sewage; fauna; settlement; stabilityl productivity; decreased water transparency and
Twenty years of investigations on the benthos in decreased depth penetration of Fucus vesculosus;
the outer part of the Elbe Estuary, which includes a increased deposition of organic matter on the
sewage-sludge dumpsite, show that the community bottom and increased frequency and severity of
structure was highly unstable. Some changes were oxygen deficency in bottom waters; and reduction
brought about by hydrological factors and the of bottom fauna. It is concluded that for most
variable pattern of sedimentation. Most of the Baltic countries efforts to reduce local
changes, however, resulted primarily from eutrophication are likely to have important positive
biological factors, such as the predominance of one effects, even when reductions in discharges of
or two species, the availability of physical space, nutrients are relatively insignificant in comparison
predation, and the life span of the species. The to the total nutrient load on the Baltic Sea.
continual arrival of larvae, belonging to various
taxa, caused a more rapid fluctuation in the benthic Cerco, C. F. (1981) Nitrification in the
community structure than is usually encountered in upper tidal James River. In: B. J. Neilson
freshwater habitats. Communities in the coastal and C. E. Cronin (ed.) Estuaries and Nutrients.
marine area of the Elbe Estuary might best be Humana Press, Clifton, New Jersey.
designated as astatic and characterized as unstable lbacteria; nitrification; ammonia; model}
but highly resilient. A field and model study of the nitrification
process in the upper tidal portion of the James
Cederwall, H. and R. Elmgren (1980) Biomass River, Virginia, has been completed. Attention was
increase of benthic macrofauna devoted to the enumeration of nitrifying bacteria
demonstrates eutrophication of the Baltic and to the determination of the fate of constituents
Sea. Ophelia, Supplementum 1:287-304. involved in the nitrification process. Ammonia and
labundance; macrofauna; benthic; l3alticl nitrite oxidizers are present in the James River
Twenty-eight stations around the Baltic islands of water column in concentrations of 10-1 - 10
bland and Gotland, sampled by Hessle in 1920 and mpm/ml and in the bottom sediments in
1923 were revisited in 1976-1977. Using methods concentrations of 102 - 105 mpm/ml. Elevated
similar to those of Hessle, we demonstrate a populations are observed in the vicinity of waste
statistically highly significant increase in discharges suggesting that nitrification of these
macrofauna biomass above the halocline (median wastes begins immediately upon discharge.
over 4 times higher, based on 23 stations) over the Examination of three sets of field data supports the
intervening 55 years. Neither differences in hypotheses that nitrification occurs. Of particular
methods, nor interannual fluctuations can explain interest is the observation, during two instances, of
the large increase, which is considered to represent a nitrate removal concurrent with nitrification.
real long-term change, apparent over the entire
Baltic Sea. The only plausible cause is Cheney, D. P. (1992) Project development to
eutrophication of the Baltic, leading to higher food harvest the beach-fouling alga Pilayella
supplies for the benthic fauna. These results littoralis in Nahant Bay, MA. 31st Annual

26
Symposium of the Northeast Algal Society, Woods seasonal patterns of growth and tissue
Hole. composition in L. digitata Lamour and L.
(seaweeds; beach; fouling; Pilayella; Nahant Bay) saccharina (L.) Lamour. Marine Ecology
ne beaches of Nahant Bay, located on the north 6:181-195.
shore of Massachusetts, have been perturbated for Iseaweeds; growth; nutrient; Laminarial
many years by an unusual, presumably mutant, Dissolved - nitrate, nitrite, ammonium and
free-living form of the brown alga Pilayella phosphate were monitored for 2.5 years along a
littoralis. Normally, P. littoralis grows as a small coastal eutrophication gradient originating at the
filamentous plant attached to larger algae in the St. Andrews sewer outfall. Linear growth rates and
littoral zone during late winter and spring months. tissue composition (phosphorus, various nitrogen
However, ever since at least 1903, large masses of fractions and storage carbohydrates) were monitored
free-living, perennial form of P. littoralis have been at the same stations in lamina tissue of Laminaria
fouling Nahant Bay beaches, often accumulating at digitata and L. saccharina. Growth rates were
low tide in amounts of hundreds of tons. In recent considerably enhanced at the eutrophicated stations
years, there have been sightings of large casts of P. both in spring, when exogenous nutrients were at
littoralis on beaches south of Nahant Bay, raising peak values at all stations, and during the summer
concerns that the distribution of the free-living when exogenous nutrients were very low at all
form may be spreading. Recently, my laboratory in stations. Enhanced summer growth rates were
collaboration with a fisheries engineer at MIT has correlated with the increased reserves of N and P
embarked upon a project aimed at developing the accumulated during winter and spring, and
necessary harvesting equipment and strategy for particularly with soluble organic nitrogen reserves.
effecting a solution to Nahant's beach-fouling Accumulation of storage carbohydrates was
problem. This project will examine the problem inversely correlated with growth rate and tissue N
from a population biology as well as engineering and Preserves, presumably because fixed carbon
viewpoint. The current distribution, quantity and in could be incorporated into new protein and thence
situ growth rate of Pilayella will be evaluated, in new tissue only if internal non-protein N reserves
order to determine if harvesting will prove were available.
economically-practical. Molecular techniques will
be used to determine the genetic relationship Copeland, B. J. and D. E. Washlschlag (1968)
between presumed mutant and attached forms. A Biological responses to nutrients -
prototype harvesting system has already been eutrophication: Saline water
developed and is being evaluated. considerations. Advances in water quality
improvement. Water Resources Symposium,
Cho, C.-H. (1991) Mariculture and Austin, Texas.
eutrophication in Jinhae Bay, Korea. Jecosystem; nutrient; production; zooplankton;
Marine Pollution Bulletin 23:275-279. community; fish}
fred tide; mortality; diatom; flagellates) 1) The addition of a known nutrient source to a
The contents of organic matters and sulfide in the marine community resulted in alteration of the
superficial bottom muds, and mass mortality of metabolic patterns of the ecosystem. An increase in
shellfish mainly due to red tides in the Jinhae Bay the amount of nutrient material caused an increase
are reviewed. COD, ignition loss, and sulfide in photosynthetic production. 2) Species diversity
contents increased for the last decade from 1972 to of zooplankton is reduced in marine environments
1982. Causative organisms of red tides also receiving various types of organic wastes, which
changed from diatom to flagellates. A decrease of indicates an interruption of normal community
shellfish production or mass mortalities of this area structure by the addition of new nutrient materials.
would be caused by a high level of eutrophication, 3) A theoretical case was presented to demonstrate
which resulted from both waste discharge from an that slight toxic effects are pronounced although
industrial complex and organic deposits from the not detectable by conventional means of
aquaculture. measurement. With just a slight increase in
mortality rate the biomass of the fishes affected
Colombo, G., I. Ferrari, V. U. Ceccherelli, and R. would be decreased greatly. 4) Experiments on the
Rossi (ed.) (1992) Marine Eutrophication metabolic'rate of fishes revealed that slight
and Population Dynamics. Olsen & Olsen, pollution stresses tend to lower the metabolic rates
Fredensborg. considerably. The depression was greatest when the
Contains eighteen papers on various aspects of fish were already subjected to regular environmental
eutrophication. stress, such as low temperature. 5) Most of the
effects of waste materials in the marine
Conolly, N. J. and E. A. Drew (1985). environment are subtle. It appears that the
Physiology of Laminaria. 111. Effects of community approach rather than the organismic
a coastal eutrophication gradient on approach will be more fruitful in evaluating the

impact of man-made changes in inland saline water blooms. In: E. M. Cosper, V. M. Brice1j and
ecosystems, notwithstanding the fact that the E. J. Carpenter (ed.) Novel Phytoplankton Blooms,
effects of stresses on individual populations can be Causes and Impacts of Recurrent Brown Tides and
physically quantifiable. other UnusualBlooms. Springer-Verlag, Berlin.
lbrown tide; bloom; Narragansett Bay; Long
Correll, D. L. (1981) Eutrophication trends Islandl
in the water quality of the Rhode River No abstract
(1971-1978). In: B. J. Neilson and L. E.
Cronin (ed.) Estuaries and Nutrients. Humana Cosper, E. M., C. Lee, and E. J. Carpenter (1990)
Press, Clifton, New Jersey. Novel "brown tide" blooms in Long
(nutrient; phosphorus) ' Island embayments: A search for the
Five to eight-year data sets on turbidity and a causes. In: E. Gran6li, B. Sunstr6m, L. Edler
series of nutrient parameters have been taken in the and D. M. Anderson (ed.) Toxic Marine
Rhode River, a small tidal river tributary to Phytoplankton. Elsevier, New York.
Chesapeake Bay. Trends of change at headwater jAurococcus; brown tide; bloom; phytoplankton;
stations indicate changes due primarily to local Long Island; organic; micronutrients; nutrients;
watershed runoff, while changes at stations near the precipitation; heterotrophic uptake}
river's mouth are indicative of changes in Unusual blooms of a previously unidentified
Chesapeake Bay. chrysophyte, Aureococcus anophaefferens; have
The data have been analyzed first by summarizing occurred in several coastal embayments along the
monthly and seasonal means and variances, then by northeast coast of the USA. The monospecific
looking for year-to-year trends. Thus, for example, blooms were termed the "brown tide" due to the
linear regressions of summer and fall turbidity resulting water color. The first appearance of the
versus time for 1971-1978 had very low slopes and "brown tide" occurred early in the summer of 1985
low coefficients of determination. An interesting over a wide geographic range in non-contiguous
finding was the pattern or total phosphorus in bodies of water. The bloom did not return to some
surface waters. At the mouth of Rhode River, bays but, on Long Island, New York they recurred
seasonal mean concentrations increased steadily for during the summer of 1986 and in diminishing
each season each year from the fall of 1971 to the densities during the summers of 1987 and 1988.
fall of 1976. The most dramatic increases (four- Historically, a diverse group of small microalgal
fold) were observed in the summer and fall. A much species dominate the phytoplankton biomass and
smaller increase occurred in winter and spring productivity in Long Island bays during the
values. In contrast, year-to-year concentrations of summer. The continued dominance through several
total phosphorus in surface waters in upstream months at high cell densities (> 109 cells 1- 1) of A.
stations over the same time period showed less anophaefferens was the distinctive feature of
clear-cut trends. Also, phosphorus loadings from consequence during these blooms.
local watershed runoff fluctuated widely, but had no Environmental variables which may contribute to
steady rapid rise with time. Although total the occurrence of the "brown tide" include elevated
phosphorus increased dramatically at the mouth of salinities due to drought conditions, pulses or
Rhode River, dissolved orthophosphate, nitrate, and rainfall delivering organic and/or micronutrients to
dissolved ammonia remained essentially constant, bay waters, reduced grazing and restricted flushing
especially in the summer and fall. These data could of bays. The "brown tide" species appears to be
indicate an increasing impact of summertime closely related to an open ocean chrysophyte,
anoxic bottom waters on the phosphorus dynamics Pelagococcus subviridis, and possibly was seeded
of the upper western shore of the Bay. into northeast coastal bays from offshore when
conditions during 1985 were particularly favorable
Cosper, E. M., V. M. Brice1j, and E. J. Carpenter for its growth. The ability of this species to
(ed.) (1989) Novel Phytoplankton Blooms: maintain at least minimal populations during the
Causes and Impacts of Recurrent Brown winter months seems to allow for its recurrence
Tides and other Unusual Blooms (Papers during subsequent summers. Culture studies have
from a conference at Stony Brook, NY, shown thatt his species has growth requirements for
27-28 October, 1988). Springer-Verlag, trace elements, chelators and organic nutrients some
Berlin. of which are different from many common estuarine
No abstract and coastal phytoplankton species. The competitive
advantage of A. anophaefferens over other
Cosper, E. M., W. Dennison, A. Milligan, E. J. potentially co-occurring species probably related to
Carpenter, C. Lee, J. Holzapfel, and L. Milanese its heterotrophic and photoadaptive capabilities.
(1989) An examination of the
environmental factors important to Costa, J. E. (1988) Eelgrass (Zostera marina
initiating and sustaining "brown tide" L.) in Buzzards Bay: Distribution,

production, and historical changes in macrofaunal communities in northern
abundance. Dissertation for Ph.D., Boston Adriatic Sea. Marine Pollution Bulletin
University. 22:503-508.
feelgrass; periphyton; abundance; growth; jAdriatic Sea; macrobenthos; Po River;
recolonization; Buzzard Bay) community structure; dystrophic; Italyl
The past and present-day distribution of eelgrass The macrozoobenthic community in the northern
(Zostera marina L.) was documented using aerial Adriatic Sea, south of the Po river, along the
photographs, field surveys, nautical charts, Emilia-Romagna region coast, was sampled in
sediment cores, and first-had accounts. Eelgrass 1985. Sampling site was central to a highly
growth correlates with local temperature and eutrophicated area with greatly increased intensity
insolation, and annual production is =350 g C m- and frequency of dystrophic events over recent
2yr-1. In Buzzards Bay, eelgrass beds cover 41 decades. The sampled community differs from all
km2 of substrate and account for 12% of primary those described in the same area in a period (1934-
production; in shallow bays, eelgrass equals 40% of 1936) previous to the actual degree of
production. eutrophication. Large abundances of species
Prior to the "wasting disease" of 1931-32, indicative of unstable bottoms, such as the bivalve
eelgrass populations equaled or exceeded present-day Corbula gibba and the polychaete Lumbrineris
abundance. Six to 10 years after the disease, latreilli were recorded. Moreover, the community
eelgrass covered less than 10% of the present-day structural features indicate a state of immaturity,
habitat area. The process of recolonization was such as in early successional stage communities.
similar in many areas: new beds initially appeared The increased frequency of acute dytrophic events
on bare substrates, beds expanded, new beds and consequent shortening of the time between
appeared, and some beds were removed by successive disturbances is proposed as the cause of
disturbance. A computer simulation modeled these bicenosis modification and its current structure and
events, and showed that rapid recolonization of composition.
eelgrass populations is highly dependent on new Crowder, A. and D. S. Painter (1991)
bed recruitment, which in nature depends on seed Submerged macrophytes in Lake Ontario:
dispersal. High disturbance rates slow eelgrass Current knowledge, importance, threats
colonization and lower peak cover. to stability, and needed studies. Canadian
Local changes in eelgrass abundance are driven by Journal of Fisheries and Aquatic Science 48:1539-
anthropogenic and natural disturbances which are 1545.
superimposed on the regional pattern of Isubmerged vegetation; stability; shading; Lake
catastrophic decline and gradual recovery. Ontario)
Hurricanes, ice scour, and freezing periodically The submerged limnetic community in Lake
destroyed eelgrass beds in some areas. Eelgrass Ontario includes algae, bryophytes, and about 30
populations in poorly flushed, developed bays, with species of angiosperms. Their distribution is
declining water quality, never recovered from the accurately known in some areas but not lake-wide,
wasting disease or showed new declines in recent and a whole-lake survey is recommended. In
years. nutrient cycling, submerged vegetation acts as a
The distribution of eelgrass is light limited, and sinks during the summer; metals and metalloids
eelgrass beds may disappear in enriched areas occur in high concentrations in tissues from some
because of increases in algal epiphytes and areas. Known herbivores include invertebrates, fish,
phytoplankton. To identify what levels of nutrient and waterfowl. Stands are necessary for many fish
loading cause these changes, concentrations and taxa as breeding or nursery habitats, and for
inputs of dissolved inorganic nitrogen (DIN) in waterfowl, but may be damaged by carp (Cyprinus
Buttermilk Bay were measured. Periphyton on carpio). Stability has been affected by water levels,
eelgrass leaves and plastic screen strips on floats sedimentation, wave and ice movement, invasive
correlated well to mean DIN. Experimental floats species, herbivory, eutrophication and turbidity, and
released nutrients and demonstrated that small contaminants. Recovery after control of P loading
increases in DIN significantly increased periphyton has occurred in Irondequoit Bay but is delayed by
abundance. The depth of eelgrass growth in turbidity in the Bay of Quinte.
Buttermilk Bay decreased by 9 cm for every I uM
increase in DIN. Periphyton abundance is more Cruz, A. A. d. I., C. T. Hackney, and J.P.Stout
important than phytoplankton concentrations in (1981) Aboveground net primary
limiting eelgrass growth in Buttermilk Bay, productivity of three Gulf Coast marsh
because water in this bay has short residence time, macrophytes in artificially fertilized
and phytoplankton gradients are less important. plots. In: B. J. Neilson and L. E. Cronin (ed.)
Estuaries and Nutrients. Humana Press, Clifton,
Crema, R., A. Castelli, and D. -Prevedelli (199 1) New Jersey.
Long term eutrophication effects on

(nitrogen; enrichment; tidal marsh; macrophytes} indefinite future. Knowledge of nutrient enrichment
Plots (100 M2) of four tidal marsh communities in freshwater systems is reviewed as a point of
(Juncus roemerianus and Spartina alterniflora in departure for understanding the response of estuaries
Alabama, J. roemerianus and Spartina cynosuroides to enrichment. Mixed and stratified estuaries
in Mississippi) common in the Gulf Coast were respond in somewhat different fashions. Major
enriched with commercial NH4NO3 (23 percent N). changes associated with enrichment are changes in
The fertilizer was applied once at the beginning of species succession and oxygen depletion in areas of
1978 growing season to stimulate a farm-plantation organic accumulation, low mixing, and poor
operation at a dosage (136 g/m2) estimated to flushing. If of temporary and local chronic nature or
return to the soil approximately the same amount of widespread occurrence, they could lead to
of nitrogen contained in the plants. Six 0.25 m2 irreversible loss of species and genetic stocks.
Measures of ecosystem health are discussed, and
quadrants were harvested monthly from each pertinent management recommendations are put
community from April to November. The annual forth. Of especial importance are the needs to
net productivity was estimated with a maximum establish local species reserves, to manage total
minus minimum standing crop technique based on system integrity, and to develop better lines of
a predictive periodic model (PPM). A correction for communication between scientists and managers.
plant mortality during the sampling period is
provided in the PPM technique. Annual Dauer, D. M. and W. G. Conner (1980) Effects
aboveground net primary productivity increased by of moderate sewage input on benthic
59 percent in the Alabama J. roemerianus, 84 polychaete populations. Estuarine and
percent in the Mississippi J. roemerianus, 82 Coastal Marine Science 10:335-346.
percent in the S. alterniflora and 26 percent in the [benthic; sewage; polychaete; abundance; biomass;
S. cynosuroides. It appears that short form or high productivity)
marsh macrophytes responded more to nitrogen The polychaete fauna of a sandy intertidal habitat
enrichment than tall form or low marsh plant. receiving effluent from a sewage treatment plant
was compared to that of a control site. The total
Cruz-Kaegi, M. E. and G. T. Rowe (1992) number of individuals, total biomass, and average
Benthic biomass gradients on the Texas- species numbers were significantly greater at the
Louisiana shelf. In: (ed.) Nutrient Enhanced sewage-affected site. The response of individual
Coastal Ocean Productivity, NECOP Workshop species to nutrient enrichment varied. Some species
Proceedings, October 1991. Texas A&M showed significantly greater numbers of individuals
University Sea Grant Program, TAMU-SG-92-109, at the sewage-affected site while other species'
(benthic abundance) densities showed no difference. All species
Macrofauna and bacterial biomass were studied in maintained greater biomass at the enriched site,
relation to depth, distance from the Mississippi most biomass differences were significant. Those
River, primary productivity, and concentration of species which did not show density differences
organic carbon in sediments. Preliminary results between sites tended to have the greater difference
suggest that combined benthic biomass was in average weight per individual. In general, species
relatively low in the Gulf of Mexico compared to with either benthic development or some form of
that off New England. Low benthic biomass is brooding behavior were best able to exploit the
thought to be a function of sediment load, enriched condition by increased densities, or greater
unpredictable hypoxic events and trawling pressure, average biomass per individual. It was concluded
counteracting the positive effects of high primary that for moderate increasee in nutrient
productivity, fine grain size and relatively high concentrations, benthic productivity will be most
concentrations of organic matter in the sediments. enhanced in those sediments with very low
Darnell, R. M. and T. M. Soniat (1981) concentrations of fine sized particles.
Nutrient enrichment and estuarine Dauvin, J. C. and F. Gentil (1989) Long-term
health. In: B. J. Neilson and L. E. Cronin (ed.) changes in populations of subtidal
Estuaries and Nutrients. Hamana Press, Clifton, bivalves (Abra alba and A. prismatica)
New Jersey. from the Bay of Morlax (Western English
Inutrient; estuary; cycle; ecosystem health) Channel). Marine Biology 103:63-73.
Ecosystem health may be defined in terms of (Abra; bivalve; organic; population; reproduction;
system norms or in terms of human utility. It is growth; abundance; life history)
here defined as that state in which the components Two populations of Abra alba (Wood) and one of
and processes remain well within specified limits of A. prismatica (Montagu) (Mollusca: Bivalvia) were
system integrity selected to assure that there is no studied over a 10 year period (1977-1987) in two
diminution in the capacity of the system to render muddy fine-sand subtidal communities of the Bay
its basic services to society throughout the of Morlax, France. The survey provided an example

30
of long-term changes in the three Abra spp. phytoplankton blooms in the western part of the
populations, which displayed synchronized changes, northern Adriatic which is under direct influence of
with a regular annual cycle and increasing densities Po River discharge. Surface salinity decreased to 16
during 1979-1980 related to the higher X 10-3, and concentration of chlorophyll a
concentration of organic matter resulting from the increased up to 17 mg m-3, with oxygen
"Amoco Cadiz" oil spill in March 1978. A. alba supersaturation up to 17 203 X 10-2.
rapidly adapts its demographic strategy to eutrophic Nanoplankton dominated the blooms (up to 76 X
conditions by increasing its reproductive potential, 109 cells m-3
growth, and abundance. During times of , but diatoms, particularly
eutrophication, A. alba has three spawning periods Chaetoceros spp. (up to 17 X 109 cells M-3), also
and two recruitments per year as opposed to two occurred in large numbers. Large 'organic'
spawning periods and two recruitments per year aggregates were formed in the water column with
during oligotrophic conditions. Growth of the dimensions (up to 2 m long) and in quantifies never
juveniles of this species is insignificant until previously observed. During late summer and early
spring for individuals recruited in the autumn, fall aggregates were distributed over the entire
whereas individuals which settle during spring or region, and a general contamination of the coasts
summer display immediate rapid growth. A. occurred. Oxygen concentration was dramatically
prismatica has a low capacity to adapt to eutrophic reduced in the bottom layer of the entire region, an
conditions. It has one annual period of sexual event not observed to such a degree since 1977.
maturation at the end of the summer, with During that year, in contrast to 1988, large
spawning in September-October and settlement quantities of freshwater spread over the entire
beginning in mid-November. Growth of the northern Adriatic causing extended phytoplankton
juveniles after settlement is also insignificant until blooms. In 1988 extremely low horizontal
April. These results enable comparison of the advection (current velocities usually below 20 cm
demographic strategies of these two sympatric S-1) primarily favoured an accumulation and high
species. aggregation of detritus, and greatly reduced oxygen
concentration in the bottom layer.
DeBoer, J. A., H. J. Guigli, T. L. Israel, and C. F.
D'Elia (1978) Nutritional studies of two red DeGroodt, E. G., H. F. J. Los, T. A. Nauta, A. A.
algae. 1. Growth rate as a function of Markus, and I. deVries (1992) Modelling
nitrogen source and concentration. Journal cause-effect relationships in
of Phycology 14:261-266. eutrophication of marine systems: an
fred algae; growth; nitrogen; culture) integral approach. In: G. Colombo, 1.
Gracilaria foliifera (Forsskal) Borgesen and Ferrari, V. U. Ceccherelli and R. Rossi (ed.)
Neoagardhiella baileyi (Harvey ex KUtzing) Wynne Marine Eutrophication and Population Dynamics.
& Taylor were grown in continuous-flow culture Olsen & Olsen, Fredensborg.
under controlled environmental conditions in 15 (modelling; nutrients; phytoplankton species
liter experimental chambers. Growth rate was composition; North Sea)
related to the source and concentration of nitrogen There is evidence that the elevated anthropogenic
enrichment supplied to the plants. Growth rate nutrient input into the North Sea is related to
appeared to follow saturation-type nutrient uptake observed changes in primary production,
kinetics for plants receiving ammonium, nitrate, phytoplankton biomass as well as species
urea or sewage effluent enrichments. Ammonium composition. Mathematical modelling techniques
enrichment produced higher growth rates than were used in order to analyze the influence of these
nitrate or sewage enrichment. The lowest growth inputs. The overall partitioning of gaines and
rates occurred in the chambers receiving unenriched losses has been simulated with a dynamic global
seawater or urea. Half saturation controls (K) for model. The model simulates the spatial and
growth were in the range of 0.2 - 0.4 uM N for all temporal concentrations of nutrients and
N enrichments examined. 'Me low estimated values phytoplankton in the entire North Sea.
of K compared closely with those found for A general, more detailed ecological model has
microalgae and indicate that both species posses the been developed for the representation of vertical
ability to utilize very low concentrations of N. inhomogeneities of nutrients and phytoplankton
species composition. Nutrient and phytoplankton
Degobbis, D . (1989) Increased levels were computed for different nutrient inputs
eutrophication of the Northern Adriatic and varying meterological conditions. The model
Sea. Marine Pollution Bulletin 20:452-457. results show an interdependence of phytoplankton
lphytoplankton; Adriatic Sea; oxygen; mucous species composition and nutrient input. However,
aggregates) there is little effect of nutrient inputs on the
Very calm and warm weather in late spring and calculated chlorophyll-a concentrations at deep
early summer 1988 supported intense non-seasonal (stratified) areas. These results indicate that

31
phytoplankton species composition may be more chlorophyll a biomass. Enhanced growth of
sensitive to changes in the nutrient inputs than the Atlantic menhaden Brevoortia tyrannus in the low
chlorophyll-a concentration. N:Si ration treatment suggested that diatoms
promote transfer of energy to higher trophic levels.
Dennis, L. and H. H. Seliger (ed.) (1979) Toxic Removal of the benthic community and associated
Dinoflagellate Blooms (Proceedings of sediments had a dramatic effect on pelagic
the Second International Conference on community structure. Zooplankton were
Toxic Dinoflagellate Blooms, Key holoplanktonic in character and higher trophic
Biscayne, Florida, 31 October - 5 levels were dominated by ctenophores, medusae,
November, 1978). Elsevier, North Holland. chaetognaths and fish. In the presence of an intact
No abstract benthos the zooplankton had a larger
meroplanktonic component and higher trophic
Dethlefsen, V. and H. V. Westernhagen (1983) levels were dominated by larvae of benthic adults
Oxygen deficiency and effects on bottom (anemones, shrimp) or temporary planktonic adults
fauna in the eastern German Bight in (mysids).
1982. Meeres/orschung 30:42-53.
lbenthos; fish; oxygen} Doi, T. and A. Nitta (1991) Ecological
In August and September 1982 two cruises were modelling at Osaka Bay related to long-
carried out to investigate the extent and effects of term eutrophication. Marine Pollution
02-deficiencies in German and Danish coastal Bulletin 23:247-252.
bottom waters. In two thirds of the more than 15 Imodelling; biomass; faunal
000 km2 covered by the survey 02-content was Based upon the ecological model presented by
less than 4 ml/l (60% saturation); lowest values Anderson & Ursin (1977), we constructed an
were near I ml/l. adequate model which will quantitatively evaluate
In bottom waters with low 02-contents, pH the overall impact of industrial large-scale
development on fisheries at Osaka Bay. It is
values were generally low (7.7). During the first expected that the patterns of changes in biomass of
cruise water in the area investigated was stratified, a dominant species, which caused by eutrophication,
thermocline was between 10-15 m above the will be explained by the model we have established.
bottom. In 02-deficiency areas (02-saturation The simulated results for two years, 1956-57, are as
around 10%) fish catches were low containing dead a whole comparable to the actual variations.
Agonus cataphractus, Pleuronectes platessa and However, the results for several species, including
Limanda limanda. When operating underwater TV sea bass, flat fish, crab, mantis shrimp, and
dead fish (Agonus cataphractus, Callionymus lara, cuttlefish, differ significantly from observed data. It
Ammodytes sp. and flatfish) were detected lying was attributed to the lack of accurate parameters of
dead on the bottom. anabolism and catabolism for younger-aged stages
Evaluation of underwater photography revealed the of these species.
occurrence of dead benthic organisms in areas with
low oxygen. The species were Ophiura albida and Dortch, Q., D. Milstead, N. N. Rabalais, S. E.
Venus stritula (paired empty valves). Possible Lohrenz, D. G. Redalje, M. J. Dagg, M. E. Turner,
causes for the occurrence of oxygen deficiencies are and T. E. Whitledge (1992) Role of silicate
discussed and it is concluded that eutrophication availablilty in phytoplankton species
processes in combinations have triggered low composition and the fate of carbon. In:
oxygen conditions in bottom waters. (ed.) Nutrient Enhanced Coastal Ocean
Productivity, NECOP Workshop Proceedings,
Doering, P. H., C. A. Oviatt, L. L. Beatty, V. F. October 1991. Texas A&M University Sea Grant
Banzon, R. Rice, S. P. Kelly, B. K. Sullivan, and Program, TAMU-SG-92-109, fcomposition,
J. B. Frithsen (1989) Structure and function nutrient ratios, silicate, hypoxia}
in a model coastal ecosystem: silicon, It has been hypothesized that hypoxia and the fate
the benthos and eutrophication. Marine of carbon are determined in part by pytoplankton
Ecology Progress Series 52:287-299. size and species composition, which are influenced
Lowering the N:Si ratio (14:1 to 1:1) in nutrient by many environmental factors, but especially by
(N, P, Si) additions to large (13 m3) outdoor the supply of N and Si. In particular when N and
mesocosms with and without an intact benthic Si concentrations are high, the predominant algal
community generally resulted in higher standing species are large, heavily to moderately silicified
stocks of diatoms. An enriched diatom flora during diatoms. While these may be grazed at rapid rates
the year-long experiment was not accompanied by a and fuel productivity at higher pelagic trophic
concomitant reduction in the extent of levels, a large portion either sinks below the
eutrophication caused by nutrient loading, as judged pycnocline directly or in fecal pellets, perhaps
by oxygen concentration and metabolism or by leading to hypoxia. When N concentrations are not

32
limiting and Si concentrations are low, as often sanguineum Hirasaka and Protogonnyaulax
happens on the Louisiana continental shelf, either tamarensis (Lebour) Taylor. The influence of
lightly silicified diatoms or non-diatoms, usually nitrogen source (i.e. N03 or NH4) on certain iron
small, flagellated algal species, predominate. These stress-mediated effects was studied, and in some
organisms yield less vertical flux of organic matter comparisons were made with nitrogen stress-
to the bottom, because most sinking mediated changes in biogeochemical composition.
phytoplankton is made up of only a few heavily or The half-saturation constant for iron-limited growth
moderately silicified diatom species. Thus, silicate (KU = 1.7-1()-20 M) of G. sanguineum was
availability determines the vertical flux of directly estimated to be 10-1000 times greater than for other
sinking phytoplankton and influences the severity neritic species investigated previously. Also, the
and extent of hypoxia. Any changes in riverine iron requirement of this dinoflagellate, -in terms of
silicate input will affect hypoxia by this Fe/C ratios, exceeded those of certain coastal
mechanism. diatoms by one to two orders of magnitude. Fe/N
Dortch, Q. and T. E. Whitledge (1992) Does ratios demonstrated a larger (1.5-fold) minimum
nitrogen or silicon limit phytoplankton iron content of N03- than NH4-grown cells, likely
production in the Mississippi River reflecting the iron content of N03 assimilatory
Plume and nearby regions? Continental enzymes. Acquisition of nitrogen by Fe-deplete,
She@f Research 12:12293-1309. N03-grown cells was sufficiently inhibited to yield
Inutrient limitation) symptoms of N deficiency, revealed by decreased
The Mississippi River carries very high (ca. 1.4-fold) N quotas and free amino acid/protein
concentrations of nutrients into the otherwise ratios compared to Fe-deplete, NH4-grown cells.
oligotrophic Gulf of Mexico, resulting in high Reductions in chlorophyll a (chl a) quotas (Qchl)
primary production and hypoxia along the and photosynthetic electron transport (PET)
Louisiana continental shelf. The hypothesis that efficiency (as measured by in vivo fluorescence
nitrogen availability controls and ultimately limits indices) occurred under Fe depletion, and are
phytoplankton production ont he shelf was tested consistent with the essential role of iron and chl a
by measuring an indicator of nitrogen deficiency, and PET component (i.e.) cytochrome and Fe-S
the ratio of intracellular free amino acids/particulate proteins) biosynthesis. Nitrogen depletion affected
protien (AA/Pr) in the area of the Mississippi
River plume on a spring ans summer cruise. Qchl similarly, but altered PET efficiency to a
Neither AA/Pr ratios or nutrients in the water markedly lesser extent than did Fe depletion. Iron-
showed nitrogen limitation to be widespread. depleted G. sanguineum exhibited an enhanced iron
Ammonium concentrations were generally quite transport capacity, which failed to be manifested
high, so the lack of phytoplankton nitrogen following a transition from NH4 to N03 nutrition.
deficiency can be explained by rapid regeneration This suppression may result from concurrent iron
rates. Nitrogen limitation was most likely in the and nitrogen stress, due to the inability of Fe-
summer at high salinities. However, ratios of deplete, NH4-grown cells to rapidly assimilate
dissolved nutrient concentrations suggested that N03. The complete initial inhibition of N03
silicate was as likely, or sometimes more likely, to uptake when Fe-deplete, NH4-grown cells were
be limiting than nitrogen. Although silicate given saturating iron additions supports this idea.
depletion -may not cause a decrease in productivity, Iron stress caused reductions in chloroplast number
it could result in major changes in phytoplanakton and some degeneration of lamellar organization in
size and species composition, and ultimately this species. For P. tamarensis, iron limitation
influence tropodynamics, regeneration, the fate of induced the formation of temporary (=pellicle) and
carbon, and severity and extent of hypoxia. resting (=hypnozygotes) cysts. Degenerative
changes in organelle (i.e. chloroplasts,
Doucette, G. J. (1989) Aspects of iron and mitochondira and chromosomes) ultrastructure were
nitrogen nutrition in two red tide largely restricted to pellicular cysts., consistent
dinoflagellates, G y m n o d i n i u in with their hypothesized role of maintaining
s a n g u i n e u m Hirasaka and variability over brief, rather than extended (c.f.
Protogonnyaulax tamarensis (Lebour) hypozygotes) exposure to adverse conditions.
Taylor. Dissertation for Ph.D., The Univeristy of
British Columbia. Dougherrty, J. E. and M. D. Morgan (1991)
(red tide; iron; nitrogen; dinoflagellate; Benthic community response (primarily
phytoplankton; ultrastructurel Chironomidae) to nutrient enrichment and
Iron stress-mediated effects on growth, alkalinization in shallow, soft water
biogeochernical composition, iron and nitrogen humic lakes. Hydrobiologia 215:73-82.
uptake, and ultrastructure have been examined in (benthic; community; nutrients; lake; pH;
the red tide dinoflagellates Gymnodinium chironomonids)

A comparison of the benthic fauna found in two of this increase appears to be of marine origin. At
shallow lakes in the New Jersey Pinelands (USA) two sites within the plume/hypoxia region,
illustrated the impact of elevated pH and nutrients preliminary estimates are that 50 to 70 percent
caused by residential and agricultural disturbance on more organic carbon is presently accumulating than
a naturally acidic, poorly buffered aquatic at the turn of the century. These preliminary
environment. Detailed community analysis interpretations provide strong support for the
suggested that change in community composition central themes of the NECOP program. Analysis
was a better indicator of response to disturbance and interpretation of further supporting information
than biological diversity indices. Chironomonidae is continuing.
(insecta) were the predominant components of the
benthic macroinvertebrate assemblage of both Edgar, R. K. (1970) The effect of nitrogen
undisturbed Oswego Lake (low pH, low nutrients) and phosphorus eutrophication on diatom
and Nescochague Lake (fluctuating pH, elevated community structure. Dissertation for Ph.D.,
nutrients). The genera Procladius, Tribelos, and Rutgers University, The State University.
Pagastiella dominated Oswego Lake, where as Idiatom; community; nitrogen; phosphorus;
Zalutschia zalutschicola, Procladius, Dicrotendipes, bacteria; phytoplankton)
and Tanytarsus dominated Nescochague Lake. An experimental, field test of the hypothesis that
Glyptotendipes was a common and unique member diatom community structure changes with changes
of the Nescochague Lake assemblage. Cluster in water quality was conducted by partitioning a
analysis indicated that the chemical differences brook with Plexiglass cylinders and maintaining in
between lakes were the most important community each different concentrations of nitrate (0. 1, 1.0 and
determinants, although, within each lake, depth and 10.0 mg N03/1) and phosphate (0.3 and 3.0 mg
substrate affected the local communities. Oswego P04/1) for 18 days. The conclusive stability with
Lake exhibited a departure nonchironomonid respect to treatments of all community structural
benthic fauna typical of low nutrient, acid lakes. In parameters - biotic composition, density, species
turn, Nescochague Lake exhibited an enriched diversity (H) and redundancy - strongly suggests
nonchironomonid fauna including mollusks and that this enrichment had no significant effect upon
planaria which were not found in Oswego Lake. the functional interactions occurring among the
species composing the community. It is suggested
Dzurica, S., C. Lee, E. M. Cosper, and E. J. that diatom communities are not as sensitive
Carpenter (1989) Role of environmental functions of water quality as they are purported to
variables, specifically organic compounds be in that their sensitivity is conditional upon the
and micronutrients on the growth of the probabilities of environmental changes affecting the
chrysophyte A u r e o c o c c u s functional status of the community and our ability
anophagefferens. In: E. M. Cosper, V. M. to detect such changes. Additionally, the results
Bricelj and E. J. Carpenter (ed.) Novel indirectly support the idea that products of bacterial
Phytoplankton Blooms, Causes and Impacts of metabolism other than nitrates and phosphates may
Recurrent Brown Tides and other Unusual Blooms. exert a controlling influence over algal
Springer-Verlag, Berlin. communities.
lbrown tide; bloom; growth}
No abstract EI-Sayed, M. K. (1982) Effect of sewage
effluent on the sediment of Northisvatnet
Eadie, B. J., J. A. Robins, P. Blackwelder, S. (a land-locked fjord), Norway. Marine
Metz, J. H. Trefy, B. McKee, and T. A. Nelson Pollution Bulletin 13:85-88.
(1992) A retrospective analysis of [sewage; fjord; Norway; sedimentsl
nutrient enhanced coastal ocean NordAsvanet is a land-locked anoxic fjord along
productivity in sediments from the the west coast of Norway which is used as a natural
Louisiana continental shelf. In: (ed.) recipient of untreated domestic sewage. The study
Nutrient Enhanced Coastal Ocean Productivity' of four core sediments collected from the heavily
NECOP Workshop Proceedings, October 1991 ' polluted basin of the fjord reflects the enrichment of
Texas A&M University Sea Grant Program, surface sediments by Fe. Mn, Cu, Zn and organic
TAMU-SG-92-109, (production, sediments) matter. This surface enrichment is entirely
Sediments have been collected and analyzed to attributed to anthropogenic input.
obtain evidence in support of the argument that
anthropogenic nutrient loading has led to changes Elkins, J. W., S. C. Wofsy, M. B. McElroy, and
in coastal water quality and increased productivity. W. A. Kaplan (1981) Nitrification and
Cores representing approximately 100 years of production of N20 in the Potomac:
input show unmistakable signs of increased evidence for variability. In: B. J. Neilson
accumulation of organic carbon beginning early in and L. E. Cronin (ed.) Estuaries and Nutrients.
the 1900s. Organic tracers show that virtually all Humana Press, Clifton, New Jersey.

34
f nitrification; ammonia; sewage} Egypt. In: R. A. Vollenweider, R. Marchetti
Extensive measurements were carried out during and R. Viviani (ed.) Marine Coastal
the summer of 1977 and 1978 to define Eutrophication. Elsevier, Amsterdam.
concentrations of inorganic nitrogen, 02 and N20 [oxygen, chlorophyll}
in the Potomac River. The chemistry of the river Sea water samples were collected from the Eastern
varied significantly between 1977 and 1978, with Harbour and El-Mex Bay during the period from
nitrification rates slower near the city of November 1987 to January 1989. The
Washington D.C. by more than a factor of 10 in concentrations of dissolved oxygen, hydrogen
1978. The nitrification rate was inversely correlated sulphide, inorganic phosphae, nitrate, nitrite,
with the rate of fresh water flow into the estuary. It ammonia, chlorophyll a and oxidizable organic
appears that production of N20 in 1978 occurred matter are scattered in the ranges (0.00-5.11 ml V
mainly as a by-product of nitrification. The 1), (0.00-2.94 mg 1-1), (0.35-17.6 ug at 1-1)
quantity of N20 released to the atmosphere (0.00-103.5 ug at 1-1), (0.00-21.6 ug at 1-1),
represented approximately 0.3 percent of sewage (0.10-88.0 ug at 1-1), (0.00-5.56 mg m-3) and
nitrogen. Conversion was more efficient in the (0. 10- 13.8 mg 02 1 - 1) respectively.
summer of 1971, about 1-5 percent, reflecting In general, El-Mex Bay exhibited higher
either additional mechanisms for production of N20 concentrations of nutrients and oxidizable organic
or larger yields for gas production in nitrification. matter, which reveal higher eutrophication levels
than the Eastern Harbour. Chlorophyll values were
Elingren, R. (1989) Man's impact on the low and similar for the surface water of both areas,
ecosystem of the Baltic Sea: Energy while the bottom water of El-Mex Bay reflected
flows today and at the turn of the slightly higher levels. Hydrogen sulphide
century. Ambio 18:326-332. concentrations showed wide variations, reflecting
fenergy flow; eutrophication; phytoplankton; higher values for the Eastern Harbour (1.33 mg V
benthic; primary; secondary; fish; production;
ecosystem) 1) compared with El-Mex Bay (0.98 mg 1-1).
The impact of man on the Baltic ecosystem in the Hydrogren sulphide-arnmonia concentrations
20th century is summarized by estimating changes showed an impressive relationship, with estimated
in major energy flows, expressed as organic carbon. ratios of H2SINHc of 1.7 and 1.03 for the surface
Recent eutrophication has increased pelagic primary water of the Eastern Harbour and El-Mex Bay
production by an estimated 30-70% and respectively.
sedimentation of organic carbon by 70-190%. The N/P ratio reflects higher values for the surface
Above the primary halocline, biomass and and bottom waters of El-Mex Bay (10.2, 13.2) in
production of benthos have increased, while 'in the comparison with the Eastern Harbour (7.0, 7.4),
deep waters of the Baltic proper, oxygen deficiency and the majority of the investigated areas seems to
has led to the loss of formerly important food be P-limited. The estimated amount of oxygen
chains over nearly 100 000 km3 of bottom. The present after complete oxidation of organic matter
net . result is an approximate doubling of ranged from 50 to 55% throughout 1988, which is
macrobenthic production, but little increase in about 14% lower than the values recorded during
meiobenthic production. Zooplankton production is the last ten years in El-Mex Bay.
thought to have increased less than primary
production, perhaps by about a quarter. Fish catches Eng, C. T., J. N. Paw, and F. Y. Guarin (1989)
in the Baltic have increased more than tenfold, but The environmental impact of aquaculture
this is considered to be due only partly to increased and the effects of pollution on coastal
fishing effort and the near extermination of fish- aquaculture development in Southeast
eating Baltic seals, though hunting and pollution, Asia. Marine Pollution Bulletin 20:335-343.
are likely to be about equally important. Today, the laquaculture; Asia}
Baltic fishery requires about 10% of the primary Coastal aquaculture is a traditional practice in
production in the Baltic to produce its'catches, Southeast Asia. Accelerated development in the last
whereas the remnant seal population needs less than three decades has created negative environmental
0. 1%. At the beginning of this century the fishery impacts, such as extensive mangrove conversion to
required about 1% and the marine mammals an ponds, changes in hydrologic regimes in enclosed
estimated 5% of the primary production. waters due to proliferation of aquaculture structures,
and discharge of high levels of organic matter into
Emara, H. I., M. A. Shriadah, T. H. Moustafa, and coastal waters. Similarly, the increasing
M. S. El-Deek (1992) Effects of sewage and deterioration of coastal water quality resulting from
industrial wastes on the chemical the discharge of domestic, agricultural and industrial
characteristics of the Eastern Harbour and wastes into coastal waters has affected aquaculture
El-Mex Bay waters of Alexandria, production and profitability. Furthermore, the

35
increased frequency of red tides in the region has relationship between these two variables was
posed serious threats to coastal aquaculture, altered.
especially to mollusc cultivation. The introduction
of management measures to mitigate deteriorating Faust, M. A. and R. J. Chrost (1981)
coastal water quality and the adverse environmental Phytosynthesis, extracellular release, and
impacts of aquaculture development has now heterotrophy of dissolved organic matter
become a matter of urgency to the region. in Rhode' River estuarine plankton. In:
B. J. Neilson and L. E. Cronin (ed.) Estuaries and
Fabrikant, R. (1984) The effect of sewage Nutrients. Humana Press, Clifton, New Jersey.
effluent on the population density and (excretion; DOM; phytoplankton; primary
size of the clam Parvilucina tenuisculpta. production; bacterial
Marine Pollution Bulletin 15:249-253. Rates of phytoplankton photosynthesis,
fsewage; density; size; bivalves; Parvilucina; extracellular release of dissolved organic matter
sediments; nitrogen; California) (DOM), and the assimilation of DOM by
Changes in the population density and size of the bacterioplankton during in situ incubation were
clam Parvilucina tenuisculpta were related to measured in the Rhode River estuary.
municipal waste effluent (in particular, organic Phytoplankton photosynthesis was in the range of
nitrogen concentration) off the coast of Palos 3 to 147 ug C I- I h- 1. The release of DOM
Verdes, California. Average clam size increased as app,@,ared to be inversely related to the
sediment organic nitrogen concentration increased, photosynthetic activity of phytoplankton. A large
but population density increased until organic percentage of the total carbon (C) fixed was released
nitrogen concentration reached a critical level when as 14C-DOM, ranging from 4 to 66 percent of the
population density decreased dramatically. These total. Ultrafiltration was used to fractionate the
effects were explained in terms of increased 14C-DOM into size classes from less than 500 to
available nutrition ( in the form of sediment 300,000 molecular weight (MW). Bacterioplankton
organic nitrogen pollutants) enhancing both
population and individual size growth. However, readily assimilated a wide range of 14C-DOM MW
when a critical concentration of organic size fractions. Both photosynthesis and phosphorus
decomposition products was reached, population uptake were higher in nanoplankton. A positive
density decreased due to the toxicity of the released linear relationship existed between log surface to
compounds. It is suggested that Parvilucina volume ratio and log phosphorus uptake per
tenuisculpta would make a good bioindicator of biomass of phytoplankton. Bacterioplankton
organic enrichment pollution. assimilated phosphorus more rapidly than
phytoplankton. The species composition of
Faganeli, J. (1982) Nutrient dynamics in the plankton populations also constantly changed. The
seawater column in the vicinity of Piran 14C-DOM represented only 0.4 to 2.3 percent of
submarine sewage outfall. Marine Pollution the total DOM in the Rhode River. Its quantity and
Bulletin 13:61-66. quality appear to be determined by the changing
Inutrient; sewage; Adriatic; nitrogen; phosphorus) phytoplantkon population with produces it, and the
Variations of nitrogen and phosphorus changing bacterioplankton population which uses
concentrations in the vicinity of Piran (North it. Preliminary data such as these may yield
Adriatic) submarine sewage outfall were studies as important insight into changes occurring in
well as the treatment plant operation. On the basis metabolic patterns of brackish ecosystems due to
of the data it was concluded that the influence of the biological enrichment with nutrients.
outfall to the nutrient level in that area is
negligible. In comparison the data from the shallow Ferraro, S. P., R. C. Swartz, F. A. Cole, and D.
organically polluted water in the inner part of the W. Schults (1991) Temporal changes in the
Bay of Kopper are reported. benthos along a pollution gradient:
Discriminating the effects of natural
Fanuko, N. (1984) The influence of phenomena from sewage-industrial
experimental sewage pollution on lagoon wastewater effects. Estuarine, Coastal and
phytoplankton. Marine Pollution Bulletin Shelf Science 33:383-407.
15:195-198. fbenthos; pollution; sewage; toxic; California;
(sewage; Yugoslavia; lagoon; phytoplankton; wastewater; sediment}
abundance; Adriatic; blooml As pollution from the Los Angeles County
In a sewage pollution experiment in the lagoon Sanitation Districts (LACSD) outfalls decreased
system at Strunjan, Yugoslavia, an artificially between 1980 and 1983, the macrobenthic
polluted lagoon showed significantly lower community partially recovered and surficial (0-2 cm
chlorophyll a biomass and cell density, and the deep) sediment contamination and toxicity decreased
at 60 m water depth along a pollution gradient from

36
the outfalls. Pollution from the LACSD outfalls Laholm Bay. Possible measures to decrease
continued to decrease but macrobenthic conditions nitrogen loading in the coastal waters, such as
and surficial sediment quality deteriorated I km, increased cultivation of catch crops and increased
was unchanged 3 km, and improved 5-15 km from spreading of manure in the springtime, are
the LACSD outfalls between 1983 and 1986. The presented. A holistic approach includes measures
net effect of natural phenomena is indicated when that may also prove beneficial to a number of
ecosystem changes occur in the opposite directions. related agricultural problems, e.g. monotonous
Our data suggest that the net effect of natural landscape (very few types of crops, decreasing
phenomena (e.g. winter storms, El Nifio) on the grass- and wet-land areas), sensitivity to
benthos was greater than LACSD wastewater environmental catastrophes (atmospheric loading by
effects I km, about equal to LACSD wastewater radioactive materials or by chemicals), surplus
effect 3 km, and less than the LASCD wastewater production, and the threat to almost 200 wild plant
effects 5-15 km from the outfalls at the LASCD species. In addition to direct agricultural measures,
1983-86 mass emission rate. Surficial sediment restoration of wetlands and streams is also
samples collected beyond the I km station from the necessary to reduce nitrogen losses to the coastal
LASCD outfalls probably represented >> 3 years of environment.
natural+effluent particulates accumulation, and they
were, therefore, better suited for detecting long-terni Fletcher, R. L., V. Cuomo, and 1. Palomba (1990)
trends than for testing short-term temporal The "Green Tide " problem, with
variability in surficial sediments significantly particular reference to the Venice
increased between 1983 and 1986, probably Lagoon. British Phycological Journal 25:87.
primarily reflecting renewed wastewater effects near 1green tide; macroalgae; Venice Lagoon; sewage;
the outfalls and the effect of natural phenomena nutrient)
(e.g. storm-induced sediment transport or erosion) The excessive growth of macroalgae is a well
further from the outfalls. Since natural phenomena known consequence of the nutrient enrichment of
may have an effect on the benthos > 3 years of coastal waters by increased levels of sewage
LASCD wastewater effects, short-term benthic discharge and agricultural "run-off'. These growths
changes must be interpreted cautiously at the study are characteristically dominated by green algae, in
site. particular species of Enteromorpha and Ulva, and
are appropriately termed "Green Tides". Such
Fiedler, P. C. (1982) Zooplankton avoidance growth weas particularly evident during the warm
and reduced grazing responses to summer of 1989 in the shallow, muddy harbours of
Gymnodinium splendens (Dinophyceae). the Solent (south coast of England). Problems
Limnology and Oceanography 27:961-965. caused by these algal growths include the fouling of
Izooplankton; red tide; production; grazing) nets and dredges, changes in the community
A dense subsurface layer of Gymnodinium structure of sediment dwelling invertebrates and the
splendens, a feature often observed in coastal waters excessive deposits of rotting driftweed on amenity
off southern California was actively avoided by beaches, requiring removal and disposal by local
macrozooplankton. Filtration rates, measured by Councils. Nowhere is the problem of "Green Tides"
uptake of trace amounts of radiocarbon-labeled more dramatically illustrated than in the Venice
diatoms, and ingestion rates, measured by gut Lagoon, Northern Adriatic. The present paper
fullness, of some herbivorous species were considers the causes and consequences of these
significantly reduced within the layer. These luxuriant macroalgal growths in the Lagoon, the
behavioral responses may help exp ,lain the biological characteristics of the dominant
formation and persistence of dinoflagellate blooms contributors and the presently used methods of
such as red tides in coastal waters often dominated removal, treatment and utilization.
by diatoms with higher maximum growth rates.
Foster, P. (1984) Nutrient distributions in
Fleischer, S., S. Hamrin, T. Kindt, L. Rydberg, the winter regime of the Northern Irish
and L. Stibe (1987) Coastal eutrophication Sea. Marine Environmental Research 13:81-95.
in Sweden: Reducing nitrogen in land Inutrient; Irish Sea; nitrogenj
runoff. Ambio 16:246-25 1. Quasi-synoptic distribution of salinity and the
(nitrogen; Sweden; runoffl concentrations of dissolved inorganic nutrients in
Eutrophication caused by excess nitrogen and the the winter regime of the Northern Irish Sea are
resulting plankton blooms in Kattegat have led to presented. Salinity to nutrient regression analyses
oxygen deficits in the bottom waters. Regional and the distributions of nutrient ratios show that
input of nitrogen affects southern Kattegat. The the characteristics of Atlantic water are variously
interdisciplinary project "Land Use-Water Quality" modified during their northward passage through the
has identified the main source and transport routes Irish Sea. Inherently different chemical
of nitrogen in the drainage basin (10 100 km2) of characteristics of the fresh water sources discharging

37
into the Northern Irish Sea generate a conceivable that the abundance of the specific
conglomeration of water types in the area. Marine flagellate species is closely related to the riverborne
waters along the Irish coast are relatively enriched P- and N-loads.
in silicon, while waters adjacent to the eastern The research as described in this paper intends to
coastal boundary are relatively enriched by simulate the phytoplankton biomass and
anthropogenic nitrogen sources. Possible composition in relation to meteorological and
implications of the spatial dichotomy in nutrient hydrophysical conditions, nutrient loads and
status for the seasonal production cycle in the zooplankton biomass. It can be considered as a
northern Irish Sea are considered. theoretical contribution towards a better
understanding of the production and eutrophication
Fransz, H. G. and J. H. G. Verhagen (1985) processes in this polluted and exploited part of
Modelling research on the production North Sea.
cycle of phytoplankton in the southern
bight of the North Sea in relation to Fraser, T. H. and W. H. Wilcox (1981)
riverborne nutrient loads. Netherlands Enrichment of a subtropical estuary with
Journal of Sea Research 19:241-250. nitrogen, phosphorus and silica. In: B. J.
Imodelling; phytoplankton; production; nutrient; Neilson and L. E. Cronin (ed.) Estuaries and
North Sea) Nutrients. Humana Press, Clifton, New Jersey.
The physio-chemical and biological composition (enrichment; nitrogen; phosphorus; silica; estuary;
of North Sea water off the Dutch coast is influenced phytoplankton)
by various factors including meteorological Seasonal pulses of nutrients are delivered to
conditions, wind and tide induced flows and mixing, Charlotte Harbor from the Peace River and other
river pollution loads, waste loads and various tributaries. Greatest loads occur during the summer
physical, chemical and biological processes. The wet season. Phosphate and nitrate values are much
long term variations (> - I month) of the higher in the Peace River than in adjacent
biological composition are mainly caused by tributaries while ammonium and silica are similar
fluctuations in meteorological conditions and among the streams.
concentrations of nutrients for algae. Phosphate dilution curves in the estuary suggest
The loads of phosphorus and nitrogen compounds that changing concentrations along the chloride
on the North Sea have increased significantly since gradient are the result of mixing processes.
the early 20th century. Especially the discharge of Inorganic nitrogen and silica dilution curves
phosphorus (P) and nitrogen (N) transported by the suggest non-conservative processes are occurring,
river Rhine has increased about sixfold since 1930. since their concentrations decrease faster along the
The nutrient rich river water originating from the chloride gradient than can be explained by dilution
rivers Scheldt, Meuse and Rhine, mixes with sea alone. Thus, both inorganic nitrogen and silica
water in such a way that 50% of the river discharge could become limiting factors in different parts of
is transported in a 15 km wide strip along the Charlotte Harbor during portions of the annual
Dutch and Belgian coast. Especially in this coastal cycle, while phosphorus is always abundant.
area, the increase in nutrient loads has had a marked Phytoplankton populations respond positively to
influence on the nutrient concentrations. In fact, the seasonal pulses of nutrients with higher
increase in N and P and the essentially constant productivity occurring during or just after high river
discharge of reactive silicate (Si) caused a change in flow. Productivity increases from the lower harbor
nutrient, first depleted by phytoplankton, from N or to near the river mouth in all seasons. General
P some 50 years ago to Si nowadays. population levels and productivity are similar to
The question, to what extent the increase in other Florida estuaries, with diatoms dominating
nutrient enrichment has given rise to enhanced the phytoplankton (blue-green algae less than I
phytoplankton growth, or to changes in species percent) community even with the increased
composition, is difficult to answer because of a nutrient levels.
lack of reliable data from the past. Nevertheless it
is conceivable that at present primary productivity Frid, C. L. J. and T. S. Mercer (1989)
may occasionally be higher than in the past. The Environmental monitoring of caged fish
upper limit to the recurrent diatom blooms in farming in macrotidal environments.
Dutch coastal waters is set by the amount of Marine Pollution Bulletin 20:379-383.
available silicate. An increase in the upper limit is Ifish farming; benthos; sedimentation)
not to be expected due to the constancy of the Si- Recent years have seen considerable expansion in
load during the last 50 years. Some flagellate fish farming using cages. These have until recently
species, however, appear to be phosphorus limited been located in the sheltered and low current
in spring and locally dominate diatoms. Nitrogen environments of sea lochs and fjords. Significant
compounds maintain low concentrations benthic enrichment is known to occur at these
throughout the summer. Therefore, it is sites, but the effects are highly localized. Fish farm

38
development is now occurring in areas with strong This paper investigates the relationship between
tidal flows. The sitting of caged fish farming in nutrients, chlorophyll a and physical variables in
macro-tidal environments may reduce the the upper Saronikos Gulf, an oligotrophic marine
environmental impact of the industry on benthic environment south of the Greater Athens
communities at the farm site. However there exists Metropolitan Area. Phosphate, silicate, ammonia,
the potential for accumulation of farm wastes in nitrite, nitrate, salinity and dissolved oxygen were
nearby sedimentary sinks. Additionally the longer determined at eight stations on 9 occasions during
residence time of the wastes in the water column summer 1982. A thermocline led to the
has the potential, especially in regions with already stratification of the water column and the
high nutrient loadings and/or long flushing times, pycnocline was related to the thermocline. The
to lead to stimulation of phytoplankton blooms. values of oxygen were more or less normal. The
Environmental monitoring of such developments eutrophication of seawater in the vicinity of the
therefore needs to consider these addition effects. sewage outfall was demonstrated by surface levels
of chlorophyll a being forty to two hundred times
Friligos, N. (1981) Enrichment by inorganic above background. This parameter provides
nutrients and oxygen utilization rates in evidence for a high phytoplankton standing stock.
Elefsis Bay (1973-1976). Marine Pollution However, there were no appreciable differences
Bulletin 12:431-436. between the nutrients in the outfall area and those
[nutrients; oxygen; Elefsis Bay; anoxicl in the background. This suggests rapid uptake of
This paper concerns the enrichment by nutrients nutrients and/or effective dispersal from the outfall.
and the oxygen utilization rates of an intermittently
anoxic basin. The concentration of nutrients and Friligos, N. (1985) Impact on phytoplankton
dissolved oxygen have been determined during populations of sewage discharges in the
seasonal surveys of water characteristics. The Saronikos Gulf (West Aegean). Water
oxygen utilization rates ranged between 1.3 and 1.8 Research 19:1107-1118.
ml V I month- I from March to June and were [sewage; nutrients; phytoplankton; standing stock)
considerably higher than those reported for oceanic Temperature, salinity and concentrations of
waters. The nutrients enrichment appears to be dissolved oxygen, nutrients, chlorophyll a,
considerably influenced by eutrophication and particulate carbon and nitrogen concentrations in
human activity. During anoxic conditions in sea water near the Keratsini sewage outfall are
summer, the mineralization of organic matter was reported. Suggestions are made on the proposed
associated with consequent accumulation of length of the outfall diffuse and the extent of the
enormous amounts of nitrogen (ammonia), sewage treatment. Eutrophication in the area around
phosphates and silicates. Nutrient ratios underwent the outfall is demonstrated by enhanced chlorophyll
great fluctuation mainly due to anoxic conditions. a, particulate carbon and particulate nitrogen
concentrations, for they provided evidence for high
Friligos, N. (1982) Enrichment of inorganic phytoplantkon standing stock. However no
nutrients in the inner Saronikos Gulf appreciable differences in the inorganic nutrient
(1973-1976). Marine Pollution Bulletin levels between the outfall area and the normal
13:154-158. background were observed. This suggests rapid
Inutrients; Saronikos Gulf; Greece; nitrogen; uptake of nutrients and/or effective dispersal from
phosphorus) the outfall, The study indicates that in the case of
This paper is concerned with the enrichment of oligotrophic waters and in the absence of the
inorganic nutrients in the Inner Saronikos Gulf near dissolved oxygen depletion, primary treatment may
the Athens sewage outfall. This Inner Gulf tends to be preferable to secondary treatment.
accumulate nutrients above the background level,
expecially ammonia which was about four times Friligos, N. (1989) Nutrient status on a
more than the background value. Comparison of silled embayment in the Ionian Sea (Gulf
nutrient enrichment was made between the Inner of Patras). Toxicological and Environmental
Gulf and Elefsis Bay. It was found that Elefsis Bay Chemistry 20-21:21-27.
contained considerably more nutrients than the Inutrient; Gulf of Patras; Mediterranean; nitrate;
Inner Gulf. This was mainly due to the different silicatel
sources of nutrients as well as morphology of each . This paper is concerned with the enrichment of
area and the circulation of the waters. inorganic nutrients in the Patraikos Fulg. The Gulf
of Patras is a silled embayment opening into the
Friligos, N. (1985) Compensation effect of Ionian Sea on the west, and through the straits of
phytoplankton on nutrients from a Rio onto the Gulf of Corinth, on the east. The
sewage outfall in summer. Hydrobiologia latter communicates though the Corinth canal with
126:53-58. the Aegean Sea. Results of the Patraikos Gulf
(sewage; nutrients; phytoplankton; abundancel show only slighter greater concentrations of

39
nutrients than background, in particular nitrate and data from 1980-1984 is 1.94 mmol/m3. Mean
silicate. Also a comparison is made with the nutrient concentrations in the area between the
nutrient concentrations in polluted coastal gulfs of southern entrance of the Great Belt and Darss Sill
the Aegean. The different nutrient levels are due to increased between 1975 and 1984 from 0.8 to 1.6
the different sources of nutrients as well as the mmol/m3 total phosphorus and from about 6 to 10
morphology and the circulation of the waters. mmol/m3 inorganic nitrogen. Comparatively
Gast, V. (1985) Bacteria as a food source higher nutrient concentrations at Bokniseck station
for microzooplankton in the Schlei Fjord are correlated to higher salinity. However, no data
and Baltic Sea with special reference to are available on nutrient imports into Kieler Bucht
ciliates. Marine Ecology Progress Series with saline deep water. Unchanged nutrient
22:107-120. concentrations at Bokniseck prior to 1975 are an
(bacteria; micro zoopl ankton; Baltic; ciliates-, argument against any dominant influence of
Uronema} anthropogenic inputs which until 1974 increased to
In situ vestigations revealed that number and annually 1500 t of total phosphorus and 12 700 t
biomass of micro zoopl ankto n increased with of total nitrogen, plus 5000 t of nitrogen from the
eutrophication along the length of the Schlei Fjord atmosphere. Higher phosphorus concentrations
(FRG). The same observation was made for total 1980-1984 are correlated to severe oxygen
bacterial numbers and biomass. Microzooplankton deficiency in the deep water. Mobilization of
of the Schlei and total bacterial numbers showed a phosphorus from sediments becoming anoxic could
minimum in winter and major periods of be the reason. Biota could be such sources and
development in late summer/autumn and spring. sinks of nutrients that influence the nutrient
Usually the microzooplankton biomass in the balance of Kieler Bucht. To sum up, there are
Schlei was greater than the bacterial biomass. In several processes besides anthropogenic inputs that
contrast, the bacterial biomass for 5 of 6 stations in influence nutrient concentrations in water. An
the Baltic Sea surpassed that of the assessment of winter water nutrient concentrations
microzooplankton during the summer. Number and alone is no adequate tool for the evaluation of the
biomass of microzooplankton in both bodies of effects of anthropogenic nutrient inputs via rivers,
water can mostly be attributed to protozoans of the sewage, land runoff, groundwater and from the
3 to 30 um fraction. Determined with the aid of atmosphere.
radioactivity labeled bacterial cultures, the filtration Gerlach, S. A. (1990) Nitrogen, phosphorus,
rate of 'natural' micro zooplankton populations plankton and oxygen deficiency in the
exhibited a distinct dependency on German Bight and in the Kiel Bay. Kieler
microzooplanktonic biomass and water temperature. Meeresforschungen, Sonderheft 7:1-34 1.
In March 1982 microzooplankton populations in JGerman Bight; Kiel Bay; Baltic; phosphate;
the eutrophic Schlei Fjord filtered 5 to 58 % of the nitrate; nitrite; ammonium; nitrogen; phosphorus;
water per day. In the central Baltic Sea in August nutrients; oxygen; hydrogen sulphide; bloom;
1982 the rate was 70 % d- I during the late stage of phytoplankton; biomass; abundanc6; sediments;
a decaying blue-green algae bloom. Laboratory turbidity; atmosphere; review; assemblage}
experiments with Uronema marinum clearly No abstract
showed that bacteria concentration exert a
considerable imjluence on the development of Giesen, W. B. J. T., M. M. v. Katwijk, and C. d.
ciliates. Only when a limiting concentration of Hartog (1990) Eelgrass condition and
about 1 x 106 bacteria ml- I is attained does a turbidity in the Dutch Wadden Sea.
proliferation of ciliates commence. Hence, bacteria Aquatic Botany 37:71-85.
ca.n represent an important food source for feelgrass; Zostera; Dutch Wadden Sea; turbidity;
microzooplankton, especially in biotopes with a submerged vegetation; fight}
high bacterial number and biomass. Populations of eelgrass (Zostera marina L.) in the
Dutch Wadden Sea have witnessed two major
Gerlach, S. A. (1988) Eutrophication of phases of decline this century. The first was the
Kieler Bucht. Kieler Meeresforschungen, total disappearance of sublittoral beds during the
Sonderheft 6:54-63. wasting disease epidemic of the early 1930s, and
JBaltic; nitrogen; phosphorus; nutrients} their subsequent failure to recover. The second was
Prior to 1975, average winter water nutrient the gradual disappearace of littoral eelgrass after the
concentrations at station Bokniseck in Kieler Bucht mid-1960s. It is argued here that both the lack of
(Western Baltic) were 1.23 mmol/m3 total recovery of the sublittoral beds, and the
phosphorus and 12.7 mmol/m3 dissolved inorganic disappearance of the littoral population, may have
nitrogen. Nitrogen concentrations did not change been effected, to a large extent, by changes in
until 1984. The mean of a few total phosphorus turbidity. The increasing turbidity can be linked

with progressively increasing eutrophication as application of this model is quite useful for testing
well as deposit extracting and dredging activities. the environmental consequences depending on
different interventions for phosphorus loading
Gieskes, W. W. C. and G. W. Kraay (1977) reductions.
Continuous plankton records: Changes in
the plankton of the North Sea and its Goldman, C. R. (1988) Primary productivity,
eutrophic southern bight from 1948 to nutrients, and transparency during the
1975. Netherlands Journal of Sea Research early onset of eutrophication in ultra-
11:334-364. oligotrophic Lake Tahoe, California-
(plankton; eutrophicationj Nevada. Limnology and Oceanography 33:1321-
No abstract 1333.
1primary productivity; transparency; turbidity}
Gilmartin, M. and N. Revelante (1980) Nutrient For more than half a century, the trophic status of
input and the summer nanoplankton water bodies has been of interest to limnologists
bloom in the Northern Adriatic Sea. and oceanographers alike. This report demonstrates
P.S.ZN.I.- Marine Ecology 1:169-180. the close, inverse relationship between 14C_
(nutrient; phytoplankton; nanoplankton; estimated primary productivity and transparency
microplankton; eutrophicated) during the earliest stages of cultural eutrophication.
A four-ye@r oceanographic data set from the As the population in the Tahoe basin has rapidly
Northern Adriatic Sea identified the factors favoring increased, Lake Tahoe has been characterized by an
development of the nanoplankton or microplankton increase in primary productivity that has averaged
components of the phytoplankton crop. 5.6% yr- I for the last 28 yr. There has been a
Latitudinally atypical summer nanoplankton concomitant decline in transparency of 0. 37 m yr- 1.
blooms, perhaps of recent origin, develop under During winter months when transparency is
vertically stratified conditions in the presence of highest, the average annual loss has been slightly
high nutrient concentrations (including silicates).
Including the summer blooms, the nanoplankton greater (0.40 m yr-1). The average annual Secchi
were responsible for more than half the total depth has decreased by 7 m during the last 19 yr of
phytoplankton production about 90% of the time. intensive monitoring. There has also been a
No single mechanism governed phytoplankton cell significant increase in the light extinction
size or size succession, but rather a number of coefficient. Photosynthetic efficiency has increased
mechanisms which were not mutually exclusive. while there has been a gradual shrinkage of the
euphotic zone. During the same period the total
Giovanardi, F. and E. Tromellini (1992) N03--N content of the lake has increased
Statistical assessment of trophic significantly, but total P content has not. Lake
conditions. Application of the OECD Tahoe primary production has become increasingly
methodology t o the marine P sensitive during the last decade as N has
environment. In: R. A. Vollenweider, R. accumulated in the system. A gradual increase in
Marchetti and R. Viviani (ed.) Marine Coastal the N:P may prove to be a general evolutionary
Eutrophication. Elsevier, Amsterdam. characteristic of oligotrophic lakes during the
Ichlorophyll, phosphorus, Adriatic Sea} earliest stages of eutrophication. The importance of
In order to study the time-space variability of the variability is particularly evident from this study.
Po-Adriatic system, a multivariate statistical
method has been used. This approach allows the Goldman, J. C. (1976) Identification of
determination of the source of variability and the nitrogen as a growth-limiting nutrient in
parameters that mainly maintain the system in a wastewaters and coastal marine waters
perturbed state. The OECD classification of through continuous culture algal assays.
trophic state for inland waters has therefore been Water Research 10:97-104.
applied to the marine environment. This trophic (wastewater; nitrogen; nutrients; phytoplankton;
classification criterion has been interpreted in terms assay; N/P ratio)
of probability of assignment to the different trophic Two series of continuous culture algal assays
categories. Functional inter-rel ation ships between conducted in 1973 and 1974, totaling 63
total phosphorus (TP) and trophic indicators experiments, were performed on mixtures of
(chlorophyll and peak chlorophyll) have been seawater and wastewater - both treated and untreated
determined. Based on the regression of TP versus - from five locations along the coasts of MA and
chlorophyll and on the log-normal distribution of RI. The results conclusively show that nitrogen
the chlorophyll data, an empirical model (Monte was the growth-limiting nutrient in these
Carlo method) has been developed, thus determing wastewaters and in coastal marine environments
the probabilities associated with the trophic level in receiving such wastes. There was a linear
the Emilia-Romagna coastal waters. The relationship between total inorganic nitrogen (IN)

41
in the influent and particulate nitrogen (PN) competition. Journal of Experimental Marine
representing algal biomass up to a IN Biology and Ecology 57:15-24.
concentration of about 10 mg 1-1. In addition, the JpH; compsitition; Phaeodactylum; Dunaliella;
N:P ratios in the test alga, Phaeodactylum phytoplankton)
tricornutum, varied between 10 and 20 (by atoms), The results of a series of competition experiments
whereas the N:P ratios in the wastewater-seawter between the chlorophyte Dunaliella tertiolecta
mixtures were between 4 and 12, thus providing (Dun) Butcher and the diatom Phaeodactylum
additional evidence that nitrogen was limiting. tricornutum (TFX-1) Bohlin demonstrate
Enrichment with nitrogen and/or phosphorus to conclusively that Phaeodactylum tricornutum
span a range of N:P ratios from 3.9 to 20 resulted dominates in intensive marine cultures when the
in a linear response in PN concentration to (sum)N pH rises above =10. This dominance results
additions. Additions of P had no effect on algal because of the diatom's unique ability among
growth. Removal of detergent-P in communities marine species to tolerate alkaline conditions.
practicing marine waste disposal will hence have When the pH is regulated both freshwater and
little impact on the control of eutrophication in marine algae, once firmly established in culture at
coastal waters. their respective pH optima, can resist invasion
from competing species. Hence, pH control may be
Goldman, J. C., Y. Azov, C. B. Riley, and M. D. a method for maintaining species other than P.
Dennett (1982) The effect of pH in tricornutum in mass culture. When Dunaliella
intensive microalgal cultures. 1. Biomass tertiolecta, however, is grown under even slight pH
regulation. Journal of Experimental Marine stress it becomes susceptible to invasion by
Biology and Ecology 57:1-13. Phaeodactylum tricornutum. This susceptibility to
JpH; phytoplankton; biomass; Phaeodactylum; takeover by P. tricornutum increases with
Dunaliella; carbon) increasing pH. In contrast, the freshwater
Two freshwater and two marine algal species were chlorophyte.Chlorella vulgaris Beji., which also is
grown in intensive continuous cultures at a fixed sensitive to increasing pH, is capable of remaining
dilution rate of 0.5 day-1, but a varying pH levels dominant at any pH within its tolerance range when
in the range 7.6 to 10.6. Both freshwater species invaded by the pH-insensitive chlorophyte,
Scenedesmus obliquus (Turp.) Kutz. and Chlorella Scenedesmus obliquus (Turp.) Kurz. Although
vulgaris Beij., grew up in pH 10.6 although C. allelopathic interactions may be responsible for the
vulgaris was more adversely affected by alkaline pH success of Chlorella vulgaris under seemingly
than was Scenedesmus obliquus . Of the marine stressful conditions, the success of Phaeodactylum
species, Phaeodactylum tricornutum (TFX-1) tricornutum at increasingly higher pH seems to be
Bohlin was hardly affected by varying pH up to its related primarily to the alga's pH tolerance
maximum tolerable level of 10.3, whereas growth characteristics an'd not to any chemical interactions
of Dunaliella tertiolecta (Dun) Butcher was with competing species.
adversely affected by increasing pH and ceased when Gowen, R. J. and N. B. Bradbury (1987) The
the pH exceeded 9.3. These results are consistent ecological impact of salmonid farming in
with the general observations that many marine coastal waters: A review. Oceano raphy and
species cannot tolerate alkaline pH values much 9
above 9.5. Moreover, the unique ability of Marine Biology: Annual Review 25:563-575.
Phaeodactylum tricornutum to grow at pH > 10 (fish farming; organic enrichment; sediments;
probably is a major factor contributing to its well review)
documented success in large-scale outdoor cultures No abstract
that are poorly buffered. It is difficult to separate Goldman, C. R. (1988) Primary productivity,
metabolic from purely chemical factors that nutrients, and transparency during the
influence the pH tolerance limits of the individual early onset of eutrophication in ultra-
species. The lower pH limits were, however, oligotrophic Lake Tahoe, California-
distinctly controlled by the production of alkalinity Nevada. Limnology and Oceanography 33:1321-
concomitant with N03- uptake, whereas the upper 1333.
pH limits in the case of Scenedesmus obliquus and [primary productivity; transparency; turbidity}
Phaeodactylum tricornutum seemed to be regulated For more than half a century, the trophic status of
primarily by metabolic control. In no case was the water bodies has been of interest to limnologists
availability of inorganic carbon an influencing and oceanographers alike. This report demonstrates
factor in setting the maximum attained pH. the close, inverse relationship between 14C_
Goldman, J. C., C. B. Riley, and M. R. Dennett estimated primary productivity and transparency
(1982) The effect of pH in intensive during the earliest stages of cultural eutrophication.
microalgal cultures. 11. Species As the population in the Tahoe basin has rapidly
increased, Lake Tahoe has been characterized by an

increase in primary productivity that has averaged suggested that an appropriate value of q can be used
5.6% yr- I for the last 28 yr. There has been a to predict the potential maximum increase in
concomitant decline in transparency of 0. 37 m yr- 1. phytoplankton which would result from a given
During winter months when transparency is anthropogenic nitrogen discharge. The sensitivity
highest, the average annual loss has been slightly of such predictions to error in determination of q is
greater (0.40 m yr-1). The average annual Secchi discussed, and the values obtained for q compared
depth has decreased by 7 m during the last 19 yr of with observations in algal culture and mesocosms.
intensive monitoring. There has also been a Grandli, E. (1981) Bioassay experiments in
significant increase in the light extinction the Falsterbo Channel- nutrients added
coefficient. Photosynthetic efficiency has increased daily. Kieler Meeresforschungen, Sonderheft
while there has been a gradual shrinkage of the 5:82-90.
euphotic zone. During the same period the total jbioassay; Baltic; nutrients; phytoplankton;
N03--N content of the lake has increased primary production; compositionj
significantly, but total P content has not. Lake I In situ enrichment bioassays were performed
Tahoe primary production has become increasingly during the summers of 1978 and 1979 in the
P sensitive during the last decade as N has Falsterbo Channel, south Baltic Sea. Phosphorus
accumulated in the system. A gradual increase in and/or nitrogen was added daily for up to two weeks
the N:P may prove to be a general evolutionary to 200 1 polyethylene bags with unfiltered surface
characteristic of oligotrophic lakes during the water. Additions of nitrogen or nitrogen plus
earliest stages of eutrophication. The importance of phosphorus invariably increased the biomass and
variability is particularly evident from this study. 14C fixation of phytoplankton. Phosphorus
additions had no such effects. Phytoplankton
Gowen, R. J., P. Tett, and K. J. Jones (1992) species which reacted most strongly to the
Predicting marine eutrophication: the enrichment were Aphanotheca sp., Nodularia
yield of chlorophyll from nitrogen in spumigena, Skeletonema costatum, Nitzschia
Scottish coastal waters. Marine Ecology closterium, Chaetoceros w'ighami and Oocystis sp.
Progress Series 85:153-161. The mean C/Chl a quotient was around 70 for
lphytoplankton; chlorophyll; prediction; model; chlorophyll a values below 6 mg m-3 but decreased
Scottish water; nitrogen; nitrite; nitrate) to about 30 for chlorophyll a values above 10.
There is perceived to be a problem of
eutrophication in European marine coastal waters Gran6h, E., P. Carlsson, P. Olsson, B. Sundstr6m,
and hence a need to predict the response in terms of W. Gran6li, and 0. Lindahl (1989) From
enhanced biomass of phytoplankton resulting from anoxia to fish poisoning: The last ten
the input of anthropogenic nitrogen. This response years of phytoplankton blooms in
was investigated indirectly by studying the Swedish marine waters. In: E. M. Cosper,
relationship between nitrate and phytoplankton V. M. Brice1j and E. J. Carpenter (ed.) Novel
chlorophyll concentrations in waters of the Scottish Phytoplankton Blooms; Causes and Impacts of
west coast. Two-thirds of the 60 data sets analyzed Recurrent Brown Tides and Other Unusual Blooms.
gave significant inverse regression of chlorophyll Springer-Verlag, Berlin.
on nitrate concentration. This result is explained by (phytoplankton; bloom; Sweden; effects)
interpreting synoptic data as representing variation No abtract
in time, with nitrate decreasing as a result of its
assimilation by phytoplankton and conversion into Gran6li, E., W. Graneli, and L. Rydberg (1986)
chlorophyll-containing biomass. Thus the absolute Nutrient limitation at the ecosystem and
value of the slope of each significant regression the phytoplankton community level in
estimates the yield (q) of chlorophyll from nitrate the Laholm Bay, south-east Kattegat.
and, indeed, from any form of nitrogen assimilable Ophelia 26:181-194.
by microalgae. The median value for q was 1.05 [nutrient; phytoplankton; abundance; nitrogen;
mg chl (mmol N)-1; the range from 0.25 to 4.4 phosphorus)
encompassed 95% of values. Some, but not all, of Nutrient enrichment experiments in the laboratory
the variation in q could be explained by error in with surface waters from the Laholm Bay, indicate
individual estimates due to chemical-analytical and that phytoplankton is nitrogen-limited. Phosphorus
sampling errors or to inhomogencities in each addition to phytoplankton samples rarely caused a
sampled phytoplankton population. The remaining biomass increase in aggreement with negligible
variation in q included a seasonal trend, which alkaline phosphatases activity. Usually the
might have resulted from changes in phytoplankton inorganic-N/P quotient in surface waters was also
species composition, nutrient limitation status, or below the Redfield ratio (16:1, by atoms),
the balance between anthropogenic and indicating potential nitrogen limitation. The supply
heterotrophic partitioning of nitrogen. It is ratio (runoff and deep water) of inorganic nutrients

43
to the surface water of the bay was 23N: 1P for scientists from the German Democratic Republic,
March-October and thus indicates a mean shortage Poland and Sweden. The water used for the
of phosphate. During summer when river runoff is experiments was collected outside Warnemiinde
small, the low N/P-ratio in the deep water (10:1) (GDR); Rostock (GDR), Sopot (P) and Falsterbo
which is mixed into surface water suggests nitrogen (S). The bioassays were performed under similar
limitation, in aggreement with the enrichment laboratory conditions. Nitrogen was the potentially
experiments. Benthic denitrification and the low most limiting nutrient for phytoplankton biomass
N/P-ratio of zooplankton excretion may make formation in coastal water from Rostock, Sopot
regenerated primary production nitrogen-limited. and Falsterbo. During winter, before the spring
The recent suggestion by Smith (1984), that phytoplankton bloom had started, phosphorus was
marine ecosystems, like lakes, are phosphorus- the "most limiting" nutrient in the Rostock area.
limited because any nitrogen shortage is alleviated For the Warnemiinde area, no clear limiting
through nitrogen fixation, does not apply to the nutrient was found, except during July 1984, when
bay itself, where water exchange is too rapid to nitrogen addition doubled the phytoplankton
allow for an adjustment of the N/P-supply quotient. biomass. The lowest algal standing stocks were
found for Warnerniinde and Flasterbo, followed by
Gran6li, E., H. Persson, and L. Edler (1986) Sopot. Extremely high chlorophyll-a values were
Connection between trace metals, found outside Rostock. Our results show that these
chelators and red tide blooms in the coastal areas in the southern Baltic do not differ
Laholm Bay, SE Kattegat - An essentially from each other with respect to the most
experimental approach. Marine limiting nutrient (sensu Liebig), although their
Environmental Research 18:61 - 78. nutrient levels are quite different. Nutrient
[copper; dinoflagellates; species composition) limitation experiments are discussed in the context
The influence of the trace metals, copper, of assumed eutrophication of the Baltic.
aluminum and iron, and of the strong complexing
agents, EDTA and NTA, on phytoplankton growth Grandli, E. and K. Sundback (1985) The
in water from a brackish water bay was investigated response of planktonic and microbenthic
through bioassay experiments. A diatom algal assemblages to nutrient enrichment
(Skeletonema costatum (Grev.) Clev) and a in shallow coastal waters, Southwest
dinoflagellate (Prorocentrum minimum (Pav.) J. Sweden. Journal of Experimental Marine
Schiller) were used as test organisms. The growth Biology and Ecology 85:253-268.
of both phytoplankton species was strongly Inutrient; nitrogen; phosphorus; microalgae;
inhibited by copper. This inhibition was generally phytoplankton; production; biomass)
inhibited by EDTA and NTA. Both phytoplankton Field and laboratory nutrient (nitrogen and
species were considerably less inhibited by phosphorus) enrichment studies were performed
aluminum than by copper at the same metal among natural phytoplankton and
concentration. While S. costatum responded to microphytobenthic assemblages from the brackish
copper and chelator additions in the same way in water Oresund, S.W. Sweden. The response of
sea water samples from different seasons, the algae from a low-nutrient area (Falsterbo Canal)
growth of P. minimum exhibited pronounced was compared to that of algae from a polluted,
seasonal variation. Other parameters than the nutrient-rich area (Lomma Bay).
values of pCu must be considered in order to The biomass (measured as chlorophyll a) of both
account for the experimental results. This work phytoplankton and microphytobenthos from the
supports the theory that alterations in contents of Falsterbo Canal increased after the addition of
trace metals and natural chelators in seawater are nitrogen. Phytoplankton growth was stimulated by
important factors behind shifts in phytoplankton the addition of phosphorus to the nitrogen-rich
species composition. water of the polluted Lomma Bay. Sediment
chlorophyll a showed no significant increase after
Gran6li, E., S. Schulz, U. Schiewer, D. the addition of nutrients in the Lomma Bay. In
Gedziorwska, W. Kaiser, and M. Plinski (1988) Is containers without sediment, phytoplankton uptake
the same nutrient limiting potential was calculated to account for = 90 % of the
phytoplankton biomass formation in disappearance of inorganic fixed nitrogen from the
different coastal areas of the Southern water. In the sediment containers the
Baltic? Kieler Meeresforschungen, Sonderheft microphytobenthos was estimated to account for =
6:191-202. 20% of the nitrogen uptake. The rest was
[phytoplankton; biomass; Baltic; nutrients; presumably lost mainly through denitrification.
chlorophyll} When containers with microphytobenthos from
During the winter 1984 and summer 1985, a Lomma Bay were kept in the dark, phosphorus was
series of limiting nutrient experiments (enrichment released at a rate of up to = 180 uM m-2 day- 1. We
tests) were performed with coastal Baltic water by suggest that by producing oxygen microbenthic

algae keep the sediment surface oxygenated thereby Bay. Field and mesocosm studies of the
decreasing phosphorus transport from the sediment effects of eutrophication and organic
to the overlying water. input on benthic population. In: J. S.
Gray and M. E. Christiansen (ed.) Marine Biology
Gran6li, E., B. Sundstr6m, L. Edler, and D. M. of Polar Regions and Effects of Stress on Marine
Anderson (ed.) (1990) Toxic Marine Organisms. John Wiley & Sons, New York.
Phytoplankton (Proceedings of the (MERL; benthic; macrofauna; Narragansett Bay;
Fourth International Conference on Toxic organic; eutrophication; populationj
Marine Phytoplankton, Lund, Sweden, Benthic macrofaunal population in Narragansett
26-30 June, 1989). Elsevier, New York. Bay and replicate mesocosms representative of
No abstract portions of Narragansett Bay respond rapidly to
variation in food supply. During five years of
Gran6li, E., K. Wallstr6m, U. Larsson, W. sampling sharp increases in August mortalities
Gran6li, and R. Elffigren (1990) Nutrient occur when estimated carbon demands for benthic
limitation of primary production in the respiration exceed the amount of organic carbon
Baltic Sea area. Ambio 19:142-151. reaching the bottom. Rapid recovery following
JBaltic; enrichment; primary production; nitrogen; these periods of high mortality indicate a very
phosphorus; nutrient; limitation) resilient community. Benthic population densities
In the Baltic Sea area, including the Kattegat, the in mesocosms rapidly increase in response to
external N/P loading ratios are generally well above external additions of plant material, and to within
the 16:1 Redfield ratio for all subareas (the system increases in productivity from nutrient
Bothnian Bay, The Bothnian Sea, the Baltic proper additions. Larval immigration and predation are also
and the Kattegat). During winter, the inorganic N/P important in determining population densities in
ratio in surface waters varies. Appreciably higher the experimental systems but these variables are
values than the loading ratio are found for the less easy to control and the effects are less obvious.
northernmost basin, the low-saline Bothnian Bay,
while lower values than the loading ratio are found Grassle, J. P. and J. F. Grassle (1984) The
for the Baltic proper and the Kattegat. Nutrient utility of studying the effects of
enrichment tests indicate general N limitation in pollutants on single species populations
the Baltic proper and the Kattegat, although in benthos of mesocosms and coastal
stimulation of algal growth after P enrichment has ecosystems. In: H. H. White (ed.) Concepts in
been found in the Baltic proper during summer Marine Pollution Measurements. Maryland Sea
blooms of blue-green algae. Blooms of blue-green Grant, College Park.
algae are common in the Baltic proper but hardly 1polychaete; bivalve; amphipod; nutrients;
ever occur in the Bothnian Bay and the Kattegat. abundance; Mediornastus ambiseta; Nucula
This has been the case for the last century, annulata; Polydora ligni; Ampelisca abdita)
indicating natural summer N limitation. Full-scale No abstract
experimental manipulation of the external N/P
loading ratio has been carried out in the Gray, J. S. (1979) Pollution -induced
HimmerfJdrdan basin, south of Stockholm. Results changes in populations. Philosophical of the
suggest nitrogen as the most limiting nutrient in Transactions of the Royal Society of London, Part
coastal areas of Baltic proper, uninfluenced by direct B 286:545-561.
nutrient discharges. The knowledge of the effects of 1population; community; pollution; organic
altered external nutrient supplies for nutrient matter; abundance; benthic fauna; strategy; log-
limitation in the Baltic Sea system as a whole is normal; indicatorl
too limited to allow for reliable predictions. The effects of pollution by organic matter, oil or
However, the Baltic Sea may have developed industrial waste on marine communities are
towards a more pronounced N limitation due to a remarkably similar. Diversity values fall, biomass
twofold historic increase in P supply relative to N and numbers of organism initially rise and then fall
supply. At present, the situation may be reversed as as the pollution load is increased. Diversity indices
N supply is probably increasing more rapidly than are, however, insensitive to pollution-induced
P supply. Management of the Baltic Sea area changes and have to be assessed subjectively.
cannot be based on removal of either N or P in Departure from a log-normal distribution of
sewage, but must take both elements into individuals among species offers a sensitive and
consideration, as well as differences between sub- objective method of assessing perturbation effects
basin and between polluted and offshore areas. on communities.
Under severe pollution stress, the dominant
Grassle, J. F., J. P. Grassle, L. S. Brown-Leger, species are those which have a flexible life-history
R. F. Petrecca, and N. J. Copley (1985) ranging from direct development to a planktonic
Subtidal macrobenthos of Narragansett larva and the ability to undergo short-term genetic

45
selection. Species having a somewhat less flexible Gray, J. S. and E. Paasche (1988) On marine
life-history strategy show increased abundance under eutrophication. Marine Pollution Bulletin
conditions of slight pollution. The increase in 19:349-350.
abundance of seven or eight neither rare nor (coast; review)
common species, which gives the departure from No abstract
log-normal distribution, is suggested as being the
most significant and the earliest detectable change Griggs, G. B. and T. S. Hopkins (1976) The
caused by pollution in a community. Thus the delineation and growth of a sludge field.
presence of a species in a polluted area may be Water Research 10:501 to 506.
more a question of life-history strategy than the (sewage; organic carbon; methane; oxygen;
tolerance of adverse environmental conditions. If sediment; anoxic)
this hypothesis is correct, considerable doubt must The color, odor, organic carbon, and acoustic
be placed on the ecological relevance of data from reflection properties of bottom sediment can be
toxicity tests. used to delineate the area affected by the Athens
sewage outfall. Both color and organic carbon
Gray, J. S. (1992) Eutrophication in the content clearly define the region of high organic
sea. In: G. Colombo, 1. Ferrari, V. U. buildup around the discharge point. Maximum
Ceccherelli and R. Rossi (ed.) Marine organic carbon values of over 6% were determined
Eutrophication and Population Dynamics. Olsen & compared to normal regional values of less than
Olsen, Fredensborg. 0.5%. The bottom area covered by black anaerobic
(review; benthic communities; general model; sediments has increased 7 fold to nearly 9 km2 in
remedial action; research needs) the past 2.5 yr. A smaller core area around the
Eutrophication begins with incresed growth rates outfall can be easily distinguished by its strong
of a range of organisms from phytoplankton and seismic reflection characteristics believed to
macroalgae to benthic animals and fish. This is coincide with the production of methane gas.
followed by a change in the species composition of
communities and then by a reduction in number of Haas, L. W., S. J. Hastings, and K. L. Webb
species as oxygen concentrationns of water begin to (1981) Phytoplankton response to a
fall. Finally, anoxia ensues and few organisms stratification-mixing cycle in the York
survive and only bacterial mats remain. River estuary during late summer. In: B.
Most data show only correlative relationships J. Neilson and L. E. Cronin (ed.) Estuaries and
between nutrient input and effects on biological Nutrients. Humana Press, Clifton, New Jersey.
organisms. For Example, there is no established iphytoplankton; hydrographic dynamics; nuisance
relationship between nutrient inputs and primary bloom)
production; nor has a quantative relationship As part of a larger multidisciplinary study of the
between organic enrichment and changes in benthic lower York River estuary, phytoplankton response
communities been established. to a tidally related cycle of stratification-
The Vollenweider nutrient loading concept has destratification was examined during August 1978.
been widely applied to the management of A "red water bloom" dominated by the
freshwater lakes. It is shown that the parameters of dinoflagellate Cocchlodinium heterolobatum'
the model cannot be estimated in marine contexts. initially observed in the lower York River
In the marine environment the most widely coincident with the spring tide-induced water
remedial action for eutrophication is for areas that column destratifiction event. It is proposed that the
are suspected to be affected to prepare budgets of dinoflagellates initiating the red tide were advected
external nutrient loadings and then to find cost- into the estuary in deep water during the preceding
effective ways of reducing the amounts of nutrients. period of stratification or were derived from cysts in
Yet examples from Norway show that such budgets the sediments and that des trati fi cation provided
can be widely inaccurate and costly political access to the surface waters. The extent of the red
decisions can be taken based on extremly poor data. water increased during the ensuing restratifiled period
In a research context, for benthic communities in the York River, and several lines of evidence
subject to eutrophication, two models are proposed; indicated that C. heterolobatum migrated diurnally
one for responses to oxygen saturation and the between ammonium enriched waters below the
other, a more speculative one for changes in C?N halocline (8-10 in) and the relatively nutrient-poor
in sediments. It is suggested that, if verified, the surface waters. Other estuarine systems in which
latter will allow quantative dose-response phytoplankton blooms associated with alternating
predictions to be made, rather than the present state- periods of stratification-destratification have been
of-the-art of simply simply making correlations observed are noted. The results illustrate the close
between nutrient loads and various properties of relationship between phytoplankton and
benthic communities. hydrographic dynamics in this estuarine system and
emphasize the necessity to include the study of

hydrographic processes in the study of Ifish; Baltic;increase; decrease)
phytoplankton dynamics. The discharge of nutrients to the Baltic Sea has
increased considerably during the 20th century.
Hagmeier, E. (1978) Variations in There are several indications that this
phytoplankton near Helgoland. Rappots et eutrophication has affected the fish community.
Proc@s-verbaux des Riunions Changes in species composition has been reported
Counseil International pour LExploration de la from coastal areas, e.g. local increases in the
Mer 172:361-363. abundance of pikeperch and decreased in whitefish.
lphytoplankton; abundance; nutrients; phosphate) Total fish catches, dominated by herring, sprat, and
A summary is given of the observations on cod, have increased tenfold in the past fifty years
Helgoland Roads between 1962 and 1974. There and doubled in the last twenty-five years. This
was a tendency of phytoplankton stocks to increase increase is mainly due to intensified fishing, but
during this period, although this was not possibly also to eutrophication and decreased fish
statistically significant. The increase was probably predation by seals.
related to the significantly rising amounts of
nutrients available. Hardy, J. T. and Z. Zubayli (1976)
Phytoplankton standing crop and sewage
Hallegraeff, G. M. (1993) A review of nutrient enrichment along the central
harmful algal blooms and their apparent coast of Lebanon. Environmental Pollution
global increase. Phycologia32:79-99. 11:195-202.
[red tides; nuisance blooms; review) (phytoplankton; Lebanon; nutrients; chlorophyll a;
no abstract sewage)
Surface nearshore water samples from seven
Hallegraeff, M. and J. L. Maclean (ed.) (1989) stations along the central coast of Lebanon were
Biology Epidemology and Management of analyzed at different seasons for concentrations for
Pyrodinium Red Tides (Proceedings of nitrate, nitrite, ammonia, phosphate and
the Management and Training Workshop, chlorophyll-a standing crop.
Bandar Seri Begawan, Brunei Darussalam, Concentrations of nutrients and chlorophyll-a are
Philippines, 23-30 May, 1989). Fisheries higher within 10 km along coastal zone south of
Department, Ministry of Development, Manila. Beirut and increase with decreasing distance from
No abstract two major untreated sewage outfall. In some cases,
chlorophyll-a standing crop near outfalls (e.g. 2.18
Han, M.-S., K. Furuya, and T. Nemoto (1992) ing m-3) are about 10 times greater than unenriched
Species-specific productivity of stations further south or than values reported from
S k e I e t o n e m a c o s t a t u m elsewhere in the eastern Mediterranean. Near
(Bacillariophyceae) in the inner part of outfalls, concentrations of phosphate, ammonia and
Tokyo Bay. Marine Ecology Progress Series chlorophyll-a tend to be greatest in summer, while
79:267-273. nitrate is greatest in winter.
lphytoplankton; productivity; Skeletonema; Tokyo Preliminary measurements by carbon-14
Bay; photosynthetic rate} assimilation indicate that the enriched coastal
The composition of red tides in Tokyo Bay varied region has a higher rate of primary production than
with season; those during summer 1986 and 1987 the unenriched area more than 10 km south of
were comprised almost entirely of Skeletonema Beriut.
costatum. Primary productivity by S. costatum
ranged from 5.2 to 70.4 % of the total productivity; Harlin, M. M. and B. Throne-Miller (1981)
on an annual basis, its contribution was 18.8 %, as Nutrient enrichment of seagrass beds in a
revealed by species- specific photosynthetic rate Rhode Island coastal lagoon. Marine
(SSP), determined by the micromanipulation of Biology 65:221-229.
14C-labelled cells under simulated in situ (lagoon; seagrass; vegetation; algae; seaweeds;
conditions. SSP of S. costatum normalized with nutrients; growth}
cell volume, and indicator of growth activity, Seagrass and algal beds showed a variety of
showed temporal variations as the species responses when the water column was treated with
composition of the red tides changed. The volume- low level additions of ammonium, nitrate and
specific SSP was high in the initial phase of the phosphate. The nutrients were added separately to 3
bloom. However, the high volume-specific SSP uniform seagrass beds of a temperate coastal lagoon
was rather short-lived. during 1979 and 1980. (1) Ammonium caused the
production of dense mats of free-floating green
Hansson, S. and L. G. Rudstani (1990) algae Enteromorpha plumosa and Ulva lactuca. It
Eutrophication and Baltic Sea fish also stimulated growth in both the leaf and root-
communities. Ambio 19:123-125. rhizome fractions of Zostera marina. This growth

response in Z. marina was greater in the area where water quality and fisheries agencies must coordinate
current reached 12 cm s- I than in the area with monitoring activities, standardize techniques, and
little or no current. The concentration of nitrogen establish and maintain long-term data sets to
in the tissue did not change. In contrast, where evaluate the effects of water quality and fisheries
current was lacking, Z. marina growth increase programs separately and together; (2) controlled,
with ammonium was small, but the concentration mesoscale, whole system experiments should be
of nitrogen in the tissue doubled over that in performed to quantify rates (e.g. growth, predation,
control plots. The growth of Ruppia maritima was etc.) of food web interactions; and (3) the scientific
inversely related to the growth of green algae in the community should promote research which
same plots. The red alga Gracilaria tilvahiae did not quantifies the impact of changes in food web
grow better in ammonium, but its tissue reddened. dynamics on changes in toxic substance levels in
(2) Nitrate additions enhanced the growth of the Great Lakes fishes.
green seaweeds Enteromorpha spp. and U. lactuca,
but not Z. marina or R. maritima. G. tilvahiae, Hein5nen, A. P. (1991) Bacterial numbers,
when fertilized in isolation from other plants, biomass and productivity in the Baltic
showed a marginal response to this nutrient, and Sea: A cruise study. Marine Ecology Progress
the tissue always reddened. (3) Phosphate enhanced Series 70:283-290.
growth in Z marina and R. maritima exposed to I bacteria; abundance; biomass; Baltic;
moderate current. G. tilvahiae growing alone productivityl
showed a small growth response to phosphate. The Bacterial numbers, biomass and productivity were
phosphate made no difference in the growth of the studied during 2 cruises in the Baltic Proper, Gulf
green seaweeds. (4) None of the nutrient of Finland and the Bothanian Sea. Bacterial
supplements noticeably altered the species population in the open sea area had longer turnover
composition of either epiphytic or planktonic algae times and lower productivity than in coastal areas,
associated with the beds, although we did detect comparable to those found in the open ocean, but
small increases in their numbers. The rapid and their biomass and abundance were as high as in the
dense growth of green algae in nitrogen -enriched coastal areas of the Baltic Sea. High bacterial
water probably limited growth of adjacent productivity in the aphotic water layer in the Gulf
seagrasses and red algae. Because these seaweeds did of Finland is suggested to indicate eutrophication of
not use the phosphate, it became available to other this area.
plant components. The overall floral response to
nutrient addition in seagrass communities depends, Herbland, A., A. L. Bouteiller, and P. Raimbault
therefore, upon the particular nutrient supplied, the (1987) Does the nutrient enrichment of
ability of alterate species in the area to compete for the equatorial upwelling influence the
that nutrient and the velocity of current in the size structure of phytoplankton in the
specific area. Atlantic Ocean? Oceanologica Acta Special
Volume No. 6:115-120.
Hartig, J. H., J. F. Kitchell, D. Scavia, and S. B. [nutrients; phytoplankton; size structure;
Brandt (1991) Rehabilitation of Lake upwellingl
Ontario: The role of nutrient reduction A study of the size structure of chlorophyll a
and food wed dynamics. Canadian Journal of (Chla) covering the major part of the equatorial
Fisheries and Aquatic Science 48:1574-1580. Atlantic Ocean from 5-N to 5-S leads to the
fLake Ontario; nutrient; food web; ecosystem) conclusion that seasonal nutrient enrichment in the
The Laurentian Lakes have a complex history of open Eastern equatorial Atlantic does not drastically
changes due to eutrophication, invasion of exotic affect the size distribution of the primary producers:
species, and fisheries and phosphorus management 90% of the total Chla is everywhere contained in
practices. Remedial actions have reduced nutrient the <10um fraction on the average. In the coastal
loadings and enhanced the role of food web upwelling near Dakar, this percentage is less than
interactions in improving water quality. Workshops 60%, for the same range of Chla concentrations. In
sponsored through the United States - Canada the equatorial region the percentage of <1 urn Chla
International Joint Commission have addressed the is the same in the deep chlorophyll maximum
relative importance of nutrient abatement and/or located at the top of the nitracline during the warm
food web manipulation in affecting water quality season and in the subsuperficial maximum of the
trends. Both controls have combined to enhance upwelling. Therefore, from an ecological point of
water clarity in Lake Michigan. Lake Ontario has view, the term "upwelling" is misleading: the
already exhibited the effects of nutrient controls and seasonal equatorial upwelling seems to be nothing
may be on the verge of manifesting food web other than the movement towards the surface of the
controls. Research and monitoring deep chlorophyll maximum, with no appreciable
recommendations to elucidate the effects of nutrient increase of its value and some slight modification
and food web controls include the following: (1) of its trophic organization.

phosphate cannot be recycled in the Wadden Sea
Herdendorf, C. E. (1986) Rebirth of Lake itself, but is rather transported out of the system,
Erie: Recovering from phosphorus thereby enhancing again bloom formation at the
enrichment. Ohio Sea Grant, Ohio Sea Grant frontal region of the Elbe River.
Series, OHSU-RS-51.
f lake; Erie; phosphorus; recovery; algae; Hesthagen, 1. H. (1977) Migrations,
phytoplankton; oxygen; benthos; fish) breeding, and growth in Pomatoschistus
No abstract minitus (Pallas) (Pisces, Gobiidae) in
Oslofjorden, Norway. Sarsia63:17-26.
Hesse, K., U. Hentschke, and U. Brockman (1992) 1growth; sand goby; Pomatoschistus; OslofJord;
A synoptic study of nutrient and length)
phytoplankton characteristics in the Aspects of the biology of the sand goby were
German Wadden Sea with respect to studied over a period of 21 months. The goby is
coastal eutrophication. In: G. Colombo, I. common in the littoral from April (at temperatures
Ferrari, V. U. Ceccherelli and R. Rossi (ed.) above 3'C) to July, when breeding takes place, and
Marine Eutrophication and Population Dynamics. from September to December, when the somatic
Olsen & Olsen, Fredensborg. growth occurs. At temperatures above 19'C and
lphytoplankton; nutrients; Wadden Sea) below 4-5'C the goby is rare in the littoral.
For the first time seasonal patterns of nutrient and The growth is very rapid, the 0-group fish
phytoplankton distribution were evaluated in an reaching almost the length of the 1-group fish
interdisciplinary synoptic study covering the whole during their first growing season. Growth is faster
area (2,500 km2) of the northern German Wadden than in other populations from similar latitudes,
Sea. Besides the large impact of the river Elbe, suggested to be due to eutrophication. Spawning
nutrient patterns revealed the importance of local occurs even in less than I m depth. There is an
sewage plumes originating from agricultural indication that light intensity, together with
drainage and coastal purification plants. N to P temperature, is important for breeding to start.
ratios were relatively high over all the area in the The food consists of any organism of appropriate
winter and spring, indicating potential phosphorus size and texture. Polychaetes are a conspicuous
limitation for phytoplankton. growth. During the component. Feeding occurs intensively all through
spring bloom of Phaeocystis globosa, which the stay in the littoral, but its lower during the
culminated in biomass concentration sof 0.5 mg C breeding season. Various features of the biology of
per liter, phosphorus levels decreased to limiting the sand goby in OslofJorden appear to be simple
concentrations in some of the outer regions of the and more clearcut when compared to other
Wadden Sea. In general, however, nutrient stocks populations studied, probably due to the stable and
were never exhausted, emphasizing the importance predictable environmental conditions.
of turbidity as a major factor controlling primary
production in this area. Hinga, K. R. (1990) Alteration of
In summer high levels of inorganic phosphate and phosphorus dynamics during experimental
low N/P ratios were observed, reflecting rapid eutrophication of enclosed marine
remineralization of organic phosphorus ecosystems. Marine Pollution Bulletin 21:275-
constituents. It is suggested that the bulk of 280.
inorganic phosphate in summer results from the (phosphorus; MERL)
breakdown of allochthonous phytoplankton blooms A 28 month eutrophication experiment was
being advected into the region from sites of intense conducted in marine mesocosms at Marine
growth such as the frontal region of the Ebe river Ecosystems Research Laboratory of the University
plume and semi-enclosed coastal basins. Red tides of Rhode Island. Each mesocosm contained 13 m3
of decomposing Noctiluca scintillans were observed of seawater and a 'layer of benthic sediments
in the Wadden Sea and massive blooms of transferred from adjacent Narragansett Bay.
Glenodinium foliaceum in the lagoons, amounting Nitrogen, phosphorus, and silica were added daily to
to concentrations of 13 mg C per liter and 19 mg C the mesocosms.
per liter, respectively. Mass occurrences of toxic The paper examines net exchange of phosphorus
forms were not recorded, although several species of between benthic sediments and water column during
Dinophysis as well as Alexandrium sp. occurred in the experiment. At low loading rates the regular
low cell numbers in spring and summer. annual pattern of phosphate concentrations is still
Allochthonus POC input was estimated to be evident but the amplitude of the pattern is
approximately 85 g C per m2 per year, equalling an magnified. At higher loading rates the annual
excess remineralization of about 13 tons of pattern is lost and the effectiveness of the sediments
inorganic phosphate per day for the whole area. to act as a 'buffer' to water column concentrations
Owing to inadequate light conditions in the Wadden is reduced. In some cases the nutrient loading
Sea water, it is assumed that this high amount of caused a release of phosphorus from the sediments.

sites. Close correlation was found between the
Hinga, K. R. (1992) Co-occurrence of logarithmic concentration of seawater inorganic
dinoflagellate blooms and high pH in nitrogen and that of tissue nitrogen (r = 0.949).
marine enclosures. Marine Ecology Progress This also applied to seawater phosphate and tissue
Series 86:181-187. phosphorus (r = 0.927). It is conclude that Ulva is
I dinoflagellates; diatoms; blooms; pH; a good bioindicator of seawater eutrophication.
phytoplanktonj
High abundances of dinoflagellates in mixed Hobbie, J. E. and J. J. Cole (1984) Response
phytoplankton populations in marine enclosures of a detrital foodweb to eutrophication.
were strongly correlated with high pH during 23 Bulletin of Marine Science 35:357-363.
enclosure-years of weekly samples. Diatom blooms (bacteria; abundance; activity)
were not similarly correlated with high pH. The The response off planktonic bacteria and micro-
correlation with high pH was not the result of protozoa to an experimental eutrophication gradient
dinoflagellate blooms themselves drawing down the in coastal marine waters was investigated. Bacterial
C02 and driving up the seawater pH. Examination and microflagellate numbers, as well as glutamic
of individual blooms of >500 cells ml- I indicates acid mineralization and 3H-thymidine
that dinoflagellate cell counts increased only after incorporation, were measured in a series of 13 m3
the pH was driven high (i.e. >8.5). High pH mesocosms for 6 months. The systems received a
occurred either by natural processes (diatom daily addition of sea water (3.7% of total) and
blooms) or, in one case, by an artificial nutrients so that the nutrients added were 0, 2, 4,
manipulation of the pH in the enclosure. There 16 and 32 times the anthropogenic nitrogen,
were 9 periods in which the seawater pH exceeded phosphorus and silica loading of Narragansett Bay,
8.5. Dinoflagellate blooms occurred during 7 of Rhode Island.
those events. A high pH affinity for dinoflagellates Both the biomass and primary production of the
could help explain reported successional sequences planktonic algae increased in direct response to
of diatom blooms followed by dinoflagellate nutrient loading through the 16X treatment; the
blooms and the association of dinoflagellate response in the 32X treatment equaled that of the
blooms with eutrophication. Seawater pH should 16X treatment. In general, bacterial numbers and
probably be included with other environmental growth (calculated from thymidine uptake) were
factors in studies of the mechanisms that control higher in the more eutrophic tanks but there were
the occurrence of field dinoflagellate blooms. lags in the microbial response to the peaks and
valleys of the algal growth. These lags caused poor
Ho, Y. B. (1987) Ulva Lactuca correlation between instantaneous measurements of
(Chlorophyta, Ulvales) in Hong Kong microbial and algal parameters such as numbers of
intertidal waters - Its nitrogen and bacteria and algal production. However, when daily
phosphorus contents and its use as a production was summed for the entire 6 months of
bioindicator of eutrophication. Asian the study there was excellent correlation between
Marine Biology 4:97-102. the total algal production and the total bacterial
(green algae; Ulva; sewage; nutrients; tissue; production (r=0.89) for six tanks. It is difficult to
nitrogen; phosphorus; bioindicator; Hong Kong} convert from 3H-thymidine incorporation to
The cosmopolitan green alga Ulva lactuca L. is absolute quantities of bacterial production. We used
often found growing in areas contaminated by a conservative conversion factor derived from
domestic sewage thus reflecting its ability to thrive laboratory values for the thtmidine content of
under such conditions. In order to study the effects bacterial cells to calculate bacterial production.
of eutrophic waters on the nutrient content of the Bacterial production (respiration plus growth) was
alga and its potential as an indicator species, both nearly a constant fraction of planktonic primary
seawater and Ulva samples were collected from 10 production at all levels of eutrophication and
rural and I I urban intertidal sites around the Hong averaged 31%. Our results are in good agreement
Kong Island over a period of two years. Analysis of with other estimates of the carbon flux through the
the water samples showed that the mean dissolved detrital component of pelagic ecosystems.
inorganic nitrogen (N03 and NH4) and phosphate
W04) levels in the rural sites were respectively Hodgkiss, I. J. and B. S. S. Chan (1987)
49.7 ug N I- I and 6.7 ug P 1- 1. Water samples Phytoplankton dynamics in Tolo
from the urban sites contained very high mean Harbour. Asian Marine Biology 4:103-112.
levels of ammonia (222.4 ug N 1-1) and phosphate (Tolo harbour-, phytoplankton; density; abundance;
composition; nutrients; nitrogen; phosphorus; red
(45.3 ug P 1-1) reflecting the extent of sewage tide; diversity)
contamination. The amount of tissue nitrogen and An intensive study of the Tolo Harbour
phosphorus in Ulva from urban areas were phytoplankton was carried out between January
respectively 71 and 93% more than those in rural 1983 and February 1985. The results presented in

50
this paper refer to the dynamics of the total the normoxic group (P<0.05 in both cases). A net
phytoplankton population at two stations, one in uptake of ammonia occurred in ammonia-enriched
the inner harbour and the other in the outer harbour, conditions - probably along a reversed NH4 +
for comparative purposes. gradient, as downhill pNH3 gradients were
Results are presented in terms of the total maintained in all groups - and may represent the
phytoplankton cell density in the entire water only means by which some brachial efflux of
column at each station; the abundance of the 5 ammonia could proceed.
major species of Bacillariophyceae and Dinophyceae
at each station; and the relative contributions of the Howells, G. P. (1972) The estuary of the
6 classes of phytoplankton at each station in 1983 Hudson River, U.S.A. Proceedings of the
and 1984. Royal Society of London, Part B 180:521-534.
Increases in total phytoplankton standing crop and JHudson River estuary; phytoplankton; nutrients;
changes in the percentage composition of the bloom)
component species are related to increasing nutrient The Hudson River is a major waterway of the
levels in Tolo Harbour, The increased contribution eastern seaboard of the United States. Its two major
of the Dinophyceae (also reflected in the increase in tributaries, the Mohawk and Hudson, combine to
red tide incidents) is particularly stressed, because of form the long narrow estuary of the Lower Hudson.
the implications which such algal blooms have for The area is populated and industrially developed,
the Tolo Harbour ecosystem. and it is predicted that both domestic and industrial
Hognestad, P. T. (1987) Assessment of the demands for water will increase, as well as the
environmental conditions in the demand for waste disposal and for the maintanence
Skagerrak and Kattegat. ICES, ICES of its recreational resources.
Cooperative Research Report, (nitrogen; Recent studies of the ecology of the Estuary,
phosphorus; Kattegat; Skagerrak; priarnry survey of the levels of pollutants in water, and their
production; oxygenj distribution between water, muds and the biota
No abstract indicate that water quality may need control if
problems of maintenance of fisheries,
Hosie, D. A., R. F. Uglow, L. Hagerman, T. eutrophication, and drinking water quality are to be
Sondergaard, and W. Weile (1991) Some effects avoided. 'Mere is clearly a case for relevant research
of . hypoxia and medium ammonia into the processes and pathways of pollutants
enrichment on efflux rates and circulating within this aquatic ecosystem, and for the study of
effects of pollutants on biological communities in
levels of ammonia in Nephrops relation to concentration and time of exposure.
norvegicus. Marine Biology 110:273-279. Estuaries are areas of great natural variations in
(hypoxia; ammonia; Kattegat; Skagerrak; decapod; environmental conditions and their flora and fauna
Nephrops norvegicus) is selected and restricted in its variety by the need to
Eutrophication has been reported for autumn tolerate or regulate in the variable conditions. For
months in regions of the Kattegat/Skagerrak, this reason, the capacity of estuaries to accept
causing stress to bottom-living organisms, The pollutants which enhance natural variations is
present studies, undertaken in April (1989), relatively great. The limit of environmental
investigated the effects of hypoxia and high acceptance need to be determined by achieving the
ammonia levels in the burrowing decapod Nephrops best reconciliation between industrial development
norvegicus (L.). The net ammonia efflux rates and and the maintenance of amenity.
circulating ammonia levels at 6 and 12 'C, at
normoxia [partial] pressure Of 02 in the water Hull, S. C. (1987) Macroalgal mats and
(toff), PW02 = 155 torr)] and hypoxia Pw02 = 24 species abundance: A field experiment.
torr) in normal seawater and ammonia-enriched (300 Estuarine, Coastal and Shelf Science 25:519-532.
umol ammonia 1-1) seawater were examined. The Imacroalgae; mats; benthos; infauna; abundance;
hourly weight-specific efflux rates were very Ythan estuaryl
variable and in all groups included some individuals A field experiment was carried out whereby the
which showed periods of no net efflux, or even a density of macroalgae (Enteromorpha spp.) was
net uptake of ammonia. At each temperature, net manipulated and the resultant changes in sedimeni
efflux-rate differences due to treatments were not infaunal density were monitored. Four densities of
significant (P>0.05; ANOVA, in all cases) and Enteromorpha spp. were used: 0, 0.3, 1 and 3 kg
only the differences between the net efflux rates of FW m-2, corresponding to control, low-, medium-,
the normoxic groups were significantly affected by and high-density plots. The experiment ran from
temperature (P<0.05; ANOVA). Circulating May to October 1985 and was sampled on three
ammonia levels were also variable, and at 6'C the occasions. By July, the density of Corophium
ammonia-enriched groups had significantly higher volutator was reduced at all weed levels when
weight-specific blood ammonia content values than

51
compared to control plots, whereas densities of principal factor affecting the physiology of cysts of
Hydrobia ulvae, Macoma balthica, Nereis Chattonella. No cysts germinate at 10 -C.
diversicolor, and Capitella capitata, all increased. Optimum temperature range is between 20 and 25
Samples taken in October when the weed mats were -C for germination. For maturation (acquisition of
buried in the sediment showed fewer differences germinability) of the cysts, low storage temperature
than in July. Macoma, Nereis, and Capitella were of I I -C or below for more than four months is
still significantly more abundant at medium and essential, whereas no significant maturation is
high weed densities. Corophuim showed no observed at 20 -C or more. In freshly collected
significant treatment effect. There was, however, a sediments, marked seasonality of germinability was
highly significant difference in population size confirmed in Suo-Nada. The cysts have
structure for Corophium. Measurements of germinability between spring and early summer.
sediment redox potential and silt content under The cysts spend a period of spontaneous dormancy
medium- and high-density plots revealed rapid between autumn and the next spring, and they
anoxia with a significant increase in siltation. mature during the winter season. The life cycle of
Chattonella is therefore well adapted to seasonal
Hungspreugs, M., W. Utoomprkporn, S. temperature fluctuation in the Seto Inland Sea. And
Dharmvanij, and P. Sompongchaiyakul (1989) further, alteration between benthic and planktonic
The present status of the aquatic stage is presumably unconstrained by virtue of
environment of Thailand. Marine Pollution shallowness of the Seto Inland Sea. The life cycle
Bulletin 20:327-3332. of Chattonella can be regarded as superior strategies
[Thailand; bloom; red tide) in conclusion for the occurrences of the red tides in
The rapidly increasing population and developing temperate waters such as the Seto Inland Sea.
agro-industrial activities in Thailand exert
considerable stress on the aquatic environment. Jaworski, N. A. (1981) Sources of nutrients
Untreated municipal wastes are discharged directly and the scale of eutrophication problems
or indirectly to canals and rivers, causing high in estuaries. In: B. J. Neilson and L. E.
BOD5 values and bacterial contamination close to Cronin (ed.) Estuaries and Nutrients. Humana
populated areas. Improvements in domestic sewage Press, Clifton, New Jersey.
treatment are likely to be prohibitively expensive [estuary, freshwater, nutrient, budget, comparison}
in the near term at least. Eutrophication of coastal A comprehensive analysis of external sources of
waters is an emerging problem. By contrast, few nutrients is presented including an impact,
problems have been documented from trace metals comparison of external loadings, and the resulting
discharged by industries, and the public health scale of eutrophication. The major emphasis of the
threat from seafood contamination does not appear analysis is on nitrogen and phosphorus.
to be significant. Uses of chlorinated hydrocarbon The relative contribution of various external
pesticides are declining sharply, these being sources for five major ecosystems is delineated.
replaced by organophosphate compounds which Discussion of seasonal and long-term trends of
degrade rapidly in aquatic environments. Oil external sources is presented. The impact of
pollution has not been a problem, although fears of external sources on the eutrophication process is
a major spill exists. evaluated. A scale of eutrophication for estuarine
ecosystem is suggested for comparing the impact of
Imai, I., S. Itakura, and K. Itoh (1991) Life nutrient enrichment.
cycle strategies of the red tide causing Comparisons of external nutrient loadings and the
flagellates Chattonella (Raphidophyceae) scale of eutrophication for 13 estuarine and
in the Seto Inland Sea. Marine Pollution freshwater ecosystems are made. A detailed
Bulletin 23:165-170. comparison of the five estuaries of the Chesapeake
(red tide; flagellates; cyst; life cycle; blooml Bay is presented from which relationships between
Chattonella antiqua (Hada) Ono and Chattonella external nutrient loadings (g/m2/yr) and eutrophic
marina (Subrahmanyan) Hara et Chihara are the conditions are suggested.
most noxious red tide flagellates which cause Analysis of the relationship between external
serious damage to fish farming, especially to nutrient loadings, nitrogen/phosphorus ratio, and
yellowtail culture, in Japanese coastal waters such eutrophic state suggests that, for the East Coast
as the Seto Inland Sea during summer. Cysts of estuarine ecosystems, if the phosphorus loading is
Chattonella were identified from sediments of the 1.0 g/m2/yr or less, excessive eutrophic conditions
Seto Inland Sea. These cysts overwinter in sea can be prevented. The favorable response of the
bottom and play an important role in initiating the Potomac estuary to phosphorus control
summer red tides. Most of the cysts adhere to solid demonstrates that excessive eutrophic conditions
surfaces such as diatom frustules and sand grains, may be alleviated with advanced wastewater
which may aid in keeping the cyst populations treatment, depending on many factors including
within seed beds for red tides. Temperature is a nutrient loadings.

form of the silicoflagellate Dictyocha speculum.
Jaworski, N. A. and J. O.Villa (1981) A This special form does not develop the siliceous
suggested approach for developing skeletons which normally characterize the genus
estuarine water quality criteria for Dictyocha, its growth is apparently independent of
management of eutrophication. In: B. J. the availability of dissolved silicate. The first
Neilson and L. E. Cronin (ed.) Estuaries and massive bloom of this organism was observed in
Nutrients. Humana Press, Clifton, New Jersey. the Kiel Bight area, FRG, in May 1983. We
(criteria; management; frameworkl suggest that the occurrence of this new type of
A conceptual approach for developing water bloom is related to changes in nutrient conditions
quality criteria for eutrophication management is due to increased eutrophication.
suggested. The three basic components of the
framework include source ambient relationships, Johansson, J. 0. R. and I. R.R. Lewis (1992)
effects, and impact analyses. The approach focuses Recent improvements of water quality
on a conceptual method for developing decision- and biological indicators in Hillsborough
making criteria as opposed to the classical water Bay, a highly impacted subdivision of
quality criteria for a single value of limitation. The Tampa Bay, Florida, USA. In: R. A.
approach to developing framework of response Vollenweider, R. Marchetti and R. Viviani (ed.)
relationships which can be readily incorporated into Marine Coastal Eutrophication. Elsevier,
water quality standard-setting processes that include Amsterdam.
environmental considerations and technological and Ichlorophyll, seagrass, blue-green algae,
economic factors. transparencyl
Hillsborough Bay, the eastern uppermost section
Jingzhong, Z., D. Liping, and Q. Baoping (1985) of the Tampa Bay system, is surrounded by a large
Preliminary studies on eutrophication and metropolitan complex, supports extensive
red tide problems in Bohai Bay. industrial activity, and serves as a major shipping
Hydrobiologia 127:27-30. port of fertilizer products. The Bay was determined
(red tide; Bohai Bay; nutrients; phytoplantkon; to be highly eutrophic during the late 1960's. The
community; composition; productivity; sewage; City of Tampa's primary sewage treatment plant
trophic state index} and runoff from fertilizer industry activities were
A study was carried out in 1978-1981 on the considered as the major sources of excessive
relationship between eutrophication and the nutrient loading. Loadings from both these sources
occurrence of red tides in Bohai Bay. have been reduced during the last 10 years, which
An assessment method using a single parameter has probably been the leading cause of improved
as well as the multiparameter trophic State Index water quality parameters such as water clarity,
was adopted. DIN, DIP, COD loading and dissolved oxygen, and chlorophyll. The
concentrations and phytoplankton community improvements recorded in these parameters may, in
(species composition, diversity index, standing part, be related to large biomass reductions of a
crop, chlorophyll a and primary productivity) were planktonic blue-green alga, which used to dominate
used as parameters. The western part of the bay and the fall and early winter phytoplankton population.
two sewage outfall sites showed the highest values Coincidental with improved water quality, seagrass
of chlorophyll a, nitrate, phosphate, standing crop, and an attached macro-alga have vegetated shallow
primary productivity and the greatest population areas around the bay, which had been barren of
instability. Therefore, Bohai Bay, can be regarded as attached vegetation for several decades.
mesotrophic, but its estuary eutrophic.
Anthropogenic factors are mainly responsible for Jones, R. A. and G. F. Lee (1986)
its eutrophication. A relationship between Eutrophication modeling for water
environmental factors and red tide species was quality management: An update of the
found. Noctiluca miliaris, Prorocentrum minimum Vollenweider OECD model. Water Quality
and Skeletonema coastatum have their bloom Bulletin 11: 118-.
periods from July to September and are the most [model; fish; yield; phosphorus; biomass;
significant species. production)
No abstract
Jochem, F. and B. Babenerd (1989) Naked
Dictyocha speculum - a new type of Jonge, V. N. d. and H. Postma (1974)
phytoplankton bloom in the Western Phosphorus compounds in the Dutch
Baltic. Marine Biology 103:373-379. Wadden Sea. Netherlands Journal of Sea
IDictyocha; phytoplankton; bloom; Research 8:139-153.
silicoflagellate; Baltic} (phosphorus; Dutch Wadden Sea; Rhine}
We report on the bloom incidence of a curious During the period of July 1970 to July 1971
new type of phytoplankton organism: the naked phosphorus concentrations (phosphate, dissolved

organic phosphorus and particulate phosphorus) organic matter over 3 yr changed in a way that
have been measured over the whole Dutch Wadden would be predicted from the faunal changes and
Sea including the Ems-Dollard area and the northern consistent with the hypothesis that sedimentation
part of the Usselmeer. had caused the variation. Increased abundance were
Phosphate values were in general higher than in accompanied by increased organic content.
the North Sea. Highest concentrations occur in
summer in the inner parts as a result of Josefson, A. B. and R. Rosenberg (1988) Long-
accumulation and mineralization of organic term soft-bottom faunal changes in three
material. In this way steep gradients are built up shallow fjords, West Sweden. Netherlands
between these inner parts and the tidal inlets, with Journal of Sea Research 22:149-159.
maxima of 2.5 ugst P/l. In winter differences are (benthic; fauna; abundance; biomass; composition;
small and the phosphate distribution is more diversity)
homogeneous. Very low phosphate values were Three West Swedish fjords were investigated at
only found in spring and summer in the tidal inlet the same 14 stations in 1976 and 1986, and 8 of
between the islands Vlieland and Terschelling where these had been investigated in the 1920's. The
phosphorus probably becomes limiting for stations are situated at 7 to 27 in water depths in
plankton growth. protected areas without any significant local
Since 1950 phosphate concentrations increased pollution input, but now with organically enriched
three times (0.5 to 1.7 ugat P/I average). Over the sediments. Comparisons of benthic fauna between
same period the river Rhine shows a similar 1976 and 1986 showed the following significant
increase. However, particulate phosphorus reductions: in total mean abundance and biomass
concentrations only doubled in the Wadden Sea (2.4 (excluding some large and rare species); in
to 5.1 ugat P/100 mg carbonate-free suspended abundance and biomass of molluscs; in abundance
matter, on the average). It has been assumed that of suspension feeders and carnivores. Similarity
the difference between the two factors is due to a indices gave significant differences between 1976
considerable increase in microbial activity. and 1986. The greatest changes had occurred in the
two fjords with the most restricted water
Josefson, A. B. (1987) Large-scale patterns circulation. Although these faunal changes may be
of dynamics in subtidal macrozoobenthic attributed to several factors, recently increased
assemblages in the Skagerrak: Effects of periods of hypoxia in the bottom water, which
a production-related factor. Marine Ecology could be a result of large-scale eutrophication, are
Progress Series 38:13-23. suggested as the main impact on the benthos.
Imacrobenthos; assemblage; organic; sediment;
abundance) Justi'c, D. (1987) Long-term eutrophication
Macrobenthic infauna at water depths between 10 of the Northern Adriatic Sea. Marine
and 300 in was monitored over 5 yr in an 80 km Pollution Bulletin 18:281-284.
wide area off the Swedish west coast. Year-to-year JAdriatic Sea; anoxia; oxygen; primary production;
variations in number of individuals were analyzed nutrients)
for total fauna, trophic and reproductive groups and In the Northern Adriatic Sea, the occurrence of
dominating species. Results demonstrated anoxic events near the bottom has considerably
significant variations in total abundance increased in frequency during the last 15 years.
synchronized between stations suggesting the Although it has been suggested that increasing
importance of some extrinsic factor. Relative nutrient inflow from land-based sources has caused
positions of dominants seemed largely unaltered, the phenomenon, there has been no direct evidence
suggesting a similar response among species to the for this. An analysis of the oxygen distribution in
common factor. Much of the variability was the Northern Adriatic Sea between 1911 and 1982
accounted for by short-lived predatory and surface- clearly demonstrates that the probability of anoxic
deposit-feeding polychaetes and surface- depo sit- events near the bottom has increased in time. This
feeding cumaceans. A comparison of the variability process is related to the increasing primary
between species with planktotrophic and production near the surface. There is strong
lecithotrophic larval development showed no evidence that the long-term nutrient enrichment of
difference between the categories, suggesting that the Northern Adriatic Sea forces the above changes.
the common factor operated on the bottom.
Temperature and salinity data suggest only small Justi'c, D. (1988) Trend in the transparency
differences between years. It is argued that the of the Northern Adriatic Sea 1911-1982.
major cause of the common variability pattern is a Marine Pollution Bulletin 19:32-35.
factor related to production in the sea such as Itransparency; Adriatic; primary production;
sedimentation of organic matter, which affects nutrientsi
either of, or a combination of, settlement, somatic In the northern Adriatic Sea, the Secchi disk has
growth and survival of the bottom. Sediment been used in oceanographic studies since 1911. An

analysis of the data collected during summers Kat, M. (1987) Toxic and non-toxic
between 1911 and 1982 demonstrates that, on the dinoflagellate blooms on the Dutch
average, the Secchi disk depth has decreased in coast. In: T. Okaichi, D. M. Anderson and T.
time. It is likely that a decrease in light penetration Nemoto (ed.) Red Tides: Biology, Environmental
has reduced the benthic primary production and thus Science, and Toxicology. Elsevier, New York.
has increased the probability of an occurrence of [bloom; dino; abundance; toxic}
anoxic events near the bottom. The long-term The water of the Dutch coastal area is a mixture
nutrient enrichment of the freshwaters discharging of water entering the North Sea in the south
into the northern Adriatic Sea appears to be the through the Straits of Dover, fresh water from the
main factor which has caused the above changes. rivers Rhine, Meuse and Scheldt and British coastal
water (figure 2.). The seasonal occurrence and year-
Justi'c, D. (1991) Hypoxic conditions in to-year frequency for the period 1973-1984 will be
the northern Adriatic Sea: Historical described of 9 dinoflagellates occurring in a 70 km
development and ecological wide area along the coast. Special attention has
significance. In: R. V. Tyson and T. H. been paid to the presence of toxin producing
Pearson (ed.) Modern and Ancient Continental dinoflagellates and the results of this program for
Shelf Anoxia. The Geological Society, London. 1986 and 1987.
(oxygen; Adriatic; Po River; nitrogen; phosphorus;
hydromedusae; meroplankton) Kautsky, N., H. Kautsky, U. Kautsky, and M.
In the northern Adriatic Sea the occurrence of Waern (1986) Decreased depth penetration
anoxic events and related benthic mortalities has of Fucus vesiculosus (L.) since the
considerably increased in frequency during the past 1940's indicates eutrophication of the
15 years. This paper summarizes recent findings Baltic Sea. Marine Ecology Progress Series
and presents new evidence on the causes and 28:1-8.
consequences of this phenomenon. Trends in the flight; depth penetration; seaweeds; chlorophyll;
oxygen content 1911-1984, trends in the Secchi Baltic)
disk depth 1911-1982 and long-term changes in the Although nutrient inputs to the Baltic Sea have
communities of benthos and meroplankton are increased drastically since the end of the last
described. The relative importance of a set of factors century, there is still little hard biological evidence
which create favorable conditions for the of a general eutrophication of the Baltic Sea outside
development of hypoxia is discussed. locally polluted areas. A revisit after 40 yr to some
well-documented diving stations in the outer
Kaiser, W., A. Irmisch, D. Nehring, F. Georgi, and archipelago of the Aland Sea gave us an
G. Bruel (1990) Ecological investigation in opportunity to register any changes in benthic
the onshore pelagic zone near vegetation that could be linked to eutrophication.
Warnemiinde from March 1985 to March By mapping the vertical distribution in coverage of
1986. Limnologica 20:33-36. bladder wrack Fucas vesiculosus (L.) at I I stations
[nutrients; chlorophyll; pelagic} we observed that the lower limit of this alga had
From March 1985 to March 1986 ecological moved upwards at 10 stations from maximally 11.5
investigations were carried out in the pelagic zone m in 1943/44 to 8.5 m in 1984. Also the depth of
near Warnemiinde with the aim of analyzing the maximum development had withdrawn from 5 to 6
annual variations in some oceanological parameters m in 1943/44 to 3 to 4 m in 1984, while coverage
and ascertaining the productivity and eutrophication at these depths was about the same: 58% and 51 %
of this region. Main results are as follows: respectively. The deeper specimens today at 8.5 m
- All parameters studied show annual variations, had the same dwarfed appearance as those found at
which are greatest for temperature, nutrients and 11.5 m in the 1940's; at that time growth at 8.5 m
biological parameters. was luxuriant. During both studies the decrease in
- Short term variations (few days) were strong, F.vesiculosus coverage with depth towards the
except in the case of temperature. lower limit could be approximately fitted to an
- Close connections exist between the biological exponentially decreasing light attenuation curve.
and chemical parameters. Since the structure of Baltic hardbottom
- The influence of the discharge from the River communities is almost totally governed by abiotic
Warnow on the ecological conditions could not be factors, the changes in depth penetration are
clarified. probably caused by decreased transparency of the
- As comparisons of chlorophyll and seston water column due to eutrophication. Results
concentrations with those of the Arkona Basin and indicate that the decreased water transparency arises
the Lijbeck Bay show, the area studied can be from a 40 to 50% increase in summer values of
characterized as a slightly eutrophic region of the chlorophyll a and nutrients in the offshore surface
Mecklenburg Bay. water of the Baltic Sea since the 1940's.

55
Keller, A. A. (1988) An empirical model of the same environmental variables as the major
primary productivity (14 C) using limiting factors. Both models provided good fits to
mesocosm data along a nutrient gradient. the data (explaining 77-82% of the total variance).
Journal of Plankton Research 10:813-834. The predictive ability of the mechanistic model was
1primary production; MERL) significantly greater (p<0.05) than the principal
The two parameters of the hyperbolic tangent component regression when tested on two
equation, Pm and a, were estimated from in situ independent data sets. Predictive ability between the
vertical profiles of primary production using two models was statistically compared by F - tests
mesocosm data along a nutrient gradient. The on the residual variability from each method.
parameters, derived from 4-h (around noon) 14C Keller, A. A., P. H. Doering, S. P. Kelly, and B.
incubations, were used together with the K. Sullivan (1990) Growth of juvenile
photosynthesis-light curve and hourly solar Atlantic menhaden, Brevoortia tyrannus
radiation data to calculate daily primary production (Pisces: Clupeidae) in MERL
rates (Pd). Approximately 40% of the daily mesocosms: Effects of eutrophication.
production occurred in the 4h around noon. Limnology and Oceanography 35:109-122.
Considering parameter uncertainty, there was no [secondary production; growth; Atlantic menhaden;
indication of an increase in variation in production nutrient; MERL)
with increased nutrient loading, nor did biomass- Growth and mortality of recently hatched Atlantic
specific P-I parameters increase. Annual production menhaden (Brevoortia tyrannus ) were examined for
ranged from 82 to 901 gCm-2 year-1 and was 7 months under two levels of nutrient enrichment
highest in the highest nutrient treatment tank, (with and without silicate enhancement) and control
Daily productivity ranged from 0.02 to 9.1 gCm- conditions in experimental marine mesocosms (13
2day-1 and was significantly correlated, in all m3). Results from six plankton-dominated systems
treatments, with a composite parameter BI01k confirmed the often hypothesized but rarely
(where B is phytoplankton biomass; I0 is daily demonstrated link between nutrient loading, food
radiation and k is the extinction coefficient). Linear quantity, and growth in marine fish. Weekly
regression of Pd against BIOlk indicated that much abundance and mortality were not significantly
different (P @t 0.05) between controls and treatment
of the variability (86%) in productivity was mesocosms. On the other hand, von Bertalanffy's
explained by light availability and phytoplankton growth coefficients and mean asymptotic sizes of
biomass. Two approaches for predicting juvenile Atlantic menhaden were significantly
productivity were compared: (i) predicting different (P < 0.05) and positively correlated (r2 =
production directly from environmental variables 0.98) with mean food availability (measured as the
(i.e. B101k ) and (ii) predicting the parameter of the sum of phytoplankton plus zooplankton carbon),
P-I curve from environmental variables and using which in turn was significantly (P < 0.05) and
these to calculate daily production. positively correlated (r2 = 0.94) with nutrient
Keller, A. A. (1989) Modeling the effects of loading (measured as the input of N+P+Si).
temperature, light, and nutrients on Keller, A. A. and R. L. Rice (1989) Effects of
primary productivity: An empirical and a nutrient enrichment on natural
mechanistic approach compared. populations of the brown tide
Limnology and Oceanography 43:82-95. phytoplankton A u r e o c o c c u s
Ichl a; 14C production; primary productivity; anophagefferens (Chrysophyceae). Journal
MERL) of Phycology 25:636-646.
The prospect of reliably predicting production (brown tide bloom; nutrient; picoalgae; MERLI
from environmental variables in heterogeneous The brown tide picoalgae Aureococcus
estuarine environments offers clear logistic anophagefferens Hargraves et Sieburth was present
advantages over measurement with the 14C in approximately equal numbers in 12 large scale
technique, particularly if detailed coverage (in space (13000 L) mesocosms at the start of a nutrient
and time) is desired. To that end, two different addition experiment in June 1985. Increases in
techniques for predicting phytoplankton production abundance in untreated systems mimicked the
were tested and compared using data from pattern of bloom development in Narragansett Bay,
Narragansett Bay, R.I., and experimental estuarine Rhode Island, the seawater source for the
ecosystems. Primary productivity was first experiment. Aureococcus increased to maximal
estimated via an empirical technique (principal values of 2.6 X 109cells L- I and persisted at high
component regression), which incorporated numbers (> 108 cells L- 1) for 7-8 weeks. In nutrient
temperature, light, and nutrients as independent addition tanks, the picoalgae bloomed briefly (1-3
variables. This approach was compared with a
traditional mechanistic formulation which included weeks) but rapidly declined to the usual level (_107

56
cells L-1 for eukaryotic algae in Narragansett Bay). loading. Average summer fluxes for treatments
The decline in picoalgae abundance was followed by ranged from about 150 to 1200 umols NH4 + m_2
an increase in total diatoms in all nutrient treated 0, with highest rates being recorded at the 2
tanks. Mean picoalgae abundance in the mesocosms highest loading levels. Benthic nutrient
and the bay was significantly (P<0.05) and regeneration did not increase in direct proportion to
inversely correlated (r=-0.93) with mean loading, but appeared strongly related to net
concentration of dissolved inorganic nitrogen. The primary production rates across the enrichment
persistence of the brown tide species in control gradient. Comparison, across the experimental
mesocosms and Narragansett Bay appears related to gradient, of the external nutrient supply rate with
its ability to grow at very low concentrations of the benthic feedback of dissolved nutrients to the
dissolved inorganic nitrogen, levels previously water column indicated that beyond a loading of
shown to limit diatom growth. about 2000 to 5000 mmols N m-2 yr- I the water
Keller, A. A. and R. L. Rice (1990) Variation column nutrient dynamics became dominated by the
in DCMU-enhanced fluorescence relative external supply. Results suggest that the
to chlorophyll a : Correlation with the importance of benthic-pelagic interactions to
brown tide bloom. Journal of Phycology biogeochernical and ecological cycles of coastal
26:202-205. areas may decrease markedly with nutrient
[brown tide blooms; fluorescence; nutrients; enrichments which exceed the needs of the
MERL) autotrophic components of the ecosystem.
DCMU-enhanced fluorescence and extracted Kelly, M. and M. Naguib (1984)
chlorophyll a were simultaneously measured in Eutrophication in Coastal Marine Areas
Narragansett Bay, Rhode Island and the MERL and Lagoons: A Case Study of 'Lac de
(Marine Ecosystems Research Laboratory) Tuins'. UNESCO, UNESCO reports in marine
mesocosms during the 1985 brown tide bloom. science, 29.
Marked differences in the relationship between these 1coast; lagoon; review; concept; processes)
variables were observed as the phytoplankton During the period from September 6 through 18,
community shifted from dominance by picoalgae to 1982 a workshop was conducted in Tunis, Tunisia
diatoms. The fluorescence to chlorophyll a ration to discuss and teach about problems of
was significantly (P<0.05) higher in the eutrophication in coastal marine areas. The history
mesocosms and the bay when the brown tide of the concept of eutrophication, as developed for
species (Aureococcus anophagefferens Hargraves et north-temperate lakes, was discussed. The major
Sieburth) dominated the phytoplankton community processes involved, including nutrient enrichment,
compared with other taxa. Although several factors excess algal growth, oxygen depletion, and the
could have affected the relationship we believe the microbial processes in anaerobic environments were
high ratios are related to the pigment composition described. Participants described eutrophic
and / or small size of the brown tide organisms. conditions found in coastal areas of their respective
Kelly, J. R., V. M. Berounsky, S. W. Nixon, and coUntries (all bordering the Mediterranean). The
C. A. Oviatt (1985) Benthic-pelagic eutrophication of Lac de Tuins, perhaps the most
coupling and nutrient cycling across an eutrophic embayments in the world, was discussed
experimental eutrophication gradient. and presented as a case study. During the workshop
Marine Ecology Progress Series 26:207-219. we realized that the concepts of cultural
[nutrient; flux; benthic; pelagic; nitrogen; eutrophication as originally described for north-
phosphorus; MERL} temperate lakes apply imperFectly to the marine
Sediment-water exchange rates of dissoloved environment. But the processes in both
environments are similar, with the addition of large
inorganic nitrogen (NH4+, N03-, N02-) and quantities of nutrients, excessive algal growth,
phosphorus (P04 3-), water column concentrations depletion of oxygen with consequent microbial and
of both dissolved and particulate forms of N and P, chemical changes, and the death of large numbers of
and net primary production (14C) were measured animals and complete change of ecosystem
during a summer period in large (13 m3) structure.
experimental mesocosms that had been subjected Kemp, W. M., R. R. Twilley, J. C. Stevenson,
to continuous daily nutrient additions (N, P, Si) for W. R. Boynton, and J. C. Means (1983) The
over I yr. The concentrations of combined decline of submerged vascular plants in
dissolved inorganic plus particulate nutrient forms the upper Chesapeake Bay: Summary of
(N, P) were linearly related to the nutrient input results concerning possible causes.
rate across a loading range from 596 to 34100 Marine Technology Society Journal 17:78-89.
mmols N m-2 yr-1. Benthic nutrient regeneration
generally increased, although not uniformly, with

57
fvascular plants; submerged; Chesapeake Bay; Kennedy, V. S. (ed.) (1984) The Estuary as a
vegeation; nutrient; epiphytic; phytoplankton; Filter. Academic Press, Orlando.
algae; biomassl No abstract
This paper provides a summary and synthesis of
research conducted to investigate possible causes of Kerr, S. R. and R. A. Ryder (1992) Effects of
the decline in abundance of submerged aquatic cultural eutrophication on coastal marine
vegetation (SAV) in the upper Chesapeake Bay fisheries: a comparative approach. In: R.
beginning in the late 1960s. Three factors were A. Vollenweider, R. Marchetti and R. Viviani (ed.)
emphasized in this study; runoff of agricultural Marine Coastal Eutrophication. Elsevier,
herbicides; erosional inputs of fine-grain sediments; Amsterdam, London, New York, Tokyo.
nutrient enrichment and associated algal growth. (fish yield}
Widespread use of herbicides in the estuarine Several generalities are common to the effects of
watershed occurred contemporaneous with the SAV cultural eutrhopication on fisheries in a variety of
loss; however, extensive sampling of estuarine marine and freshwater ecosystems. Foremost is the
water and sediments during 1980-81 revealed that absence of definitive diagnostic features to permit
typical bay concentrations of herbicides (primarily discrimination between direct eutrophication effects,
atrazine) rarely exceeded 2 ppb. On two occasions and those engendered by other factors, often
relatively high values (20-45 ppb) were observed associated with anthropogenic change. It is also
for brief (2-4 h) periods in a small cove following clear that euthrophication may profoundly modify
runoff events. Short (2-6 h) and long (4-6 wk) term the structure, function and species composition of
experiments indicated that ephemeral phytoxic fish production systems, with attendant
effects would be expected in response to these consequences for both ecosystem function and the
highest herbicide concentrations followed by rapid economic value of the yield. Eutrophication and its
recovery. However, normal concentrations (<5 ppb) associated effects fall into one of four categories:
had little measurable effects on plants. Historical direct effects on essential environmental factors;
increases in turbidity have been documented for diminished suitability of the habitat factors
some bay tributaries since the 1940s. During our necessary to sustain fisheries yeirld; transmission
study light (PAR) attenuated by suspended fine- of effects accelerated by the rapid transfer of
grain sediments contributed more to total turbidity inimical biota among affected ecosystems; potential
in bay shallows (< 1.5 in) than did phytoplankton for elaboration of fisheries yield in terms of
chlorophyll a. Diel cycles of PAR available in products that are not acceptable for human
SAV beds indicated that plant photosynthesis was comsumption. Taken together, the patterns of
light-limited for much of the day, and PAR often change accompanying cultural euthrophication of
fell below the compensation level (IC) needed for fish production systems are generally recognizable,
minimal plant growth. Although some SAV but are not always amenable to differentiation from
species exhibited considerable ability to adapt to the effects of associated factors, or to precise
reduce light by such mechanisms as increased quantification.
pigmentation and stem elongation, increased
turbidity has probably reduced overall depth K61iler, A. and F. H61zel (1980) Investigation
distribution of SAV markedly. Effects of the on health conditions of flounder and
continual increase in nutrient enrichment of the bay smelt in the Elbe estuary. Helgoldnder
(documented since 1930) were tested by Meeresuntersuchungen 33:401-414.
experimentally fertilizing pond mesocosms at Idisease; health; flounder; smelt; River Elbe; North
levels common to the upper estuary. Moderate to Sea; oxygen; nitrite)
high nutrient loadings resulted in significant Since 1958 the occurrence of flounder (Platichthys
increases in growth of epiphytic and planktonic flesus L.) and smelt (Osmerus eperlanus L.) in the
algae and decreases in SAV production, as well as lower River Elbe has shifted gradually toward the
premature seasonal senescence of fertilized plant mouth of the river into the North Sea: this may be
populations. Direct measurements demonstrated the due to periodical deprivation of oxygen and
inhibitory effect of epiphytic growth on SAV fluctuating peaks of toxic nitrite concentrations.
photosynthesis, due largely to light attenuation. The interior organs of both fish species caught in
The results of these various experiments were the Elbe estuary were examined. Macroscopic
synthesized into an ecosystem simulation model inspections revealed discolouring of the liver to be
which demonstrated the relative potential most pronounced in specimens 2-3 years of age.
contributions of the 3 factors to SAV declines, Cross-sections of the liver demonstrated
where nutrients > sediments > herbicides. Other histopathological symptoms of oedematous
factors and mechanisms are also discussed along degeneration in smelt, and extreme lipoid
with possible resource management options. vacuolation in flounder increasing with size and age
of the fishes. The intestine of both species
investigated showed massive desquarnation defects

58
of the mucosa, occurring in smelts shortly after high individual numbers per m2 all over the
hatching, and in flounders after immigration into Dogger Bank in contrast to 1951-52, when they
the estuary. The observations made suggest the were found only in few individuals. Extensive
influence of toxic compounds distribution in the patches of the bivalve species Spisula subtruncata
aquatic environment on the health of both species. discovered at the northeastern border of the Dogger
Bank in the fifties were not found. Only small,
Kononen, K. and A. Niemi (1984) Long-term fast-growing species like Abra prismatica, Tellina
variation of the phytoplankton Jabula and Montacuta bidentata occurred in high
composition at the entrance to the Gulf individual numbers per m2. In 1987 the total
of Finland. Ophelia, supplementum3:101-110. biomass shows a reduction of about 30% in the
(phytoplankton abundance; assemblage; area compared with 1950-54.
composition; blue-green algae; N/P ratio} Some hypotheses are given to explain the changes
Quantitative data on the phytoplankton in the sea in macrofauna distribution on the Dogger Bank
zone off Tvdrminne in 1968-1981 have been between 1950-1954 and 1985-1987, taking into
analyzed. The area is characterized by strong account the increasing eutrophication and pollution.
maxima of diatoms and dinoflagellates in spring
and occasional maxima of blue-green algae in late Lam, C. W. Y. and K. C. Ho (1987) Red tides
summer. Rapid changes in hydrography, including in Tolo Harbour, Hong Kong. In: T.
interactions of different water masses, are frequent Okaichi, D. M. Anderson and T. Nernoto (ed.) Red
in the area. The year-to-year fluctuations of different Tides: Biology, Environmental Science and
phytoplankton species and groups are discussed Toxicology. Elsevier, New York.
against the background of the hydrographical and fred tide; oxygen}
chemical changes in the Baltic Proper in the 1970s. Tolo harbour, a 15 km long landlocked inlet in
Rises in the surface salinity and nutrient level were the northeastern part of Hong Kong, has been
reflected by the increased total biomass, especially severely affected by red tides over recent years. The
in summer, during the late seventies, and frequency of occurences has markedly increased
heterocystic blue-green algae became more since 1980 with 69 incidents recorded from 1980-
abundant. The vernal marine diatoms were also 1986. A corresponding increase in dinoflagellate
more abundant in the late seventies than in the late numbers in the inlet was found. More than twenty
sixties, but this was partly caused by variations in causative organisms were identified. Prorocentrum
weather conditions in spring. Diatoma elongatum, tridtinum, P. dentaturn, P. sigmoides
which favours low salinity, was more abundant in Gymnodinium sp., Noctiluca scintillans and a
the late sixties. However, many dominant marine variety of small flagellates occurred most
species showed considerable year-to-year fluctuation frequently. Red tides were found throughout the
not directly connected with the changes observed in whole year with peak occurrence in spring and
the environment during the study period. autumn. The water quality, oceanography and
Kr6ncke, 1. (1990) Macrofauna standing meteorological conditions characterizing red tide
stock of the Dogger Bank. A comparison: occurrence during the period from 1976-1986 were
11. 1951-1952 versus 1985-1987 Are analyzed and reported. 'Me increasing red tides are a
changes in the community of the consequence of accelerated eutrophication in Tolo
northeastern part of the Dogger Bank due Harbour following intensive urban development in
to environmental changes? Netherlands the catchment. The nutrient loadings of nitrogen
Journal of Sea Research 25:189-198. and phosphorus have increased more than two-fold
Imacrofauna; benthic; opportunistic; polychaetes; in the past decade. Concurrent increase in the
bivalves; species composition; abundance; nutrient levels in the inlet water has been found.
biomass; Dogger Bank} The major impact of red tides at present is causing
During April/May 1985-87 some of Ursin's fishkills by anoxia.
(Ursin, 1960) stations on the Dogger Bank from Lara, R. J., E. A. Gomez, and A. E. Pucci (1985)
April/May 1951-52 were revisited. This paper Organic matter, sediment particle size
concentrates on results from the 'Tail End', the and nutrient distributions in a sewage
northeastern part of the Dogger Bank. The affected shallow channel. Marine Pollution
polychaete species Ophelia borealis and Goniada Bulletin 16:360-364.
maculata, which were dominant species in 1951-52, (nitrogen; phosphorus; Blanca Bay; sediment;
showed similar distributions in 1985-87 on the protein)
whole of the Dogger Bank. Changes in the Sediment and seawater samples were taken in a
macrofauna distribution are obvious in the increase sewage affected channel in Blanca Bay, Argentina.
of opportunistic small polychaete species like
Spiophanes bombyx, Scoloplos armiger and Protein content, 'humic substances' and silt-clay
Magelona spp. In 1985-87 these species occurred in percentage were evaluated in the sediments; NH4

59
P04 3+ and 'dissolved organic matter' in seawater. the frequency of dividing cells (FDC) method as
A highly significant correlation (r---0.747, n=59) being 38 g C m-2 yr-1 at the control station and
was obtained for the relation between protein and % 50 g C m-2 yr- I at the eutrophication stations. To
silt-clay. The 'humic substances' distribution estimate the mean in situ bacterial cell volume a
depends more on the distance from the outfall than correlation between FDC and cell volume was used.
does the protein content, probably because of the The increased annual primary production in the
important relation of the latter with the sediment eutrophicated area was due mainly to higher
particle size. The dissolved components showed a production during spring and autumn, largely by
dilution pattern similar to that of a contamination phytoplankton cells (mainly diatoms) retaind by a
plume in a moving water mass, reaching normal 10 um filter. Primary production during summer
values at 1700 m from the outfall. was similar in the two areas, as was the
distribution on different size fractions. This could
Larsson, U., R. Elmgren, and F. Wulff (1985) possibly explain the similar bacterial production in
Eutrophication and the Baltic Sea: the trophic layers at all stations since the bulk of
Causes and consequences. Ambio 14:9-14. bacterial production occurs during summer. It was
(nitrogen; Phosphorus; Baltic) demonstrated that selective filtration does not
New estimates of the total nutrient (N and P) load quantitatively separate photoautotrophs and
to the Baltic Sea are considerably higher than earlier bacteria. A substantial fraction of the primary
calculations. Estimated total nitrogen load is 1.2 X production occurs in the size fraction <3um. The
106 tons/year, total phosphorus load 77 X 103 primary production encountered in the 3-0.2 um
tons/year. These figures may still be too low. The fraction was due to abundant picoplankton (0.5 to
load may have increased about 4x for total nitrogen gX 107 ind l- 1), easily passing 3 um filter. The
and about 8x for total phosphorus, due to man's picoplankton was estimated to constitute up to
activities. 25% of the total phytoplankton biomass in the
Available data indicate that phosphorus is control area and up to 10% in the eutrophication
deposited in large amounts in the sediment, and that area.
nitrogen is lost from the sediment through
denitrification. Laws, E. A. (1983) Man's impact on the
marine nitrogen cycle. In: E. J. Carpenter
Larsson, U. and Hagstr6m (1982) and D. G. Capone (ed.) Nitrogen in the Marine
Fractionated phytoplankton primary Environment. Academic Press, New York.
production, exudate release and bacterial [nitrogen; cycle; budget; New York Bight; sewage;
production in a Baltic eutrophication Kanaohe Bay}
gradient. Marine Biology 67:57-70. No abstract
(phytoplankton; primary production; bacteria;
Baltic) Laws, E. A. and D. G. Reda1je (1979) Effect of
The distribution of phytoplankton primary sewage enrichment on the phytoplankton
production into four size fractions (>10um, 10- population of a subtropical estuary.
3um, 3-0.2 um and <0.2 um), the utilization of Pacific Science 33:129-148.
algal exudates by bacteria and the bacterial Isewage; phytoplankton; estuary; primary
production were studied in a eutrophication gradient production; Kanaeohe Bay; chlorophyll a; biomass;
in the northern Baltic proper. The polluted area nutrients; light; growth ratel
exhibits substantially increased nutrient, especially Phytoplankton primary production; concentrations
nitrogen, levels while only minor differences occur of chlorophyll a, particulate carbon and nitrogen,
in salinity and temperature regimes. Total primary adenosine triphosphate, inorganic nitrogen and
production was 160 g C m-2 yr- I at the control phosphorus; and secchi depths were measured at
station and about 275 g C m-2 yr- I at the four stations in Kanaeohe Bay, Oahu, on a
eutrophicated stations. The estimated total exudate biweekly basis for 20 months prior to diversion of
release was 16% of the totally fixed 14CO2 in the sewage discharges from the bay. Nutrient
enrichment experiments designed to determine
control area and 12% in the eutrophicated area biomass limitation indicated that phytoplankton
(including the estimated bacterial uptake of biomass, as measured by chlorophyll a, was
exudates). The difference in 14CO2 uptake rates nitrogen-limited in all parts of the bay, and that
between incubation of previously filtered water (.6, phosphorus was simultaneously limiting in the
<2, <1 urn) and unfiltered water was used to sector of the bay furthest from the sewer outfalls.
estimated bacterial uptake of phytoplankton Mean light-saturated productivity indices in all
exudated which were found to contribute about half parts of the bay were about 11-12 ing C mg-1 chl a
of the estimated bacterial carbon requirement in hr- 1, values close to the maximum reported for
both areas. Bacterial production was estimated by phytoplankton in eutrophic marine environments.

60
Based on the results of dawn-to dusk C-14 community as a result of the sewage discharges
incubations and an estimated phytoplankton C: chl appear to have resulted from the elevated
a ratio of 50 by weight, phytoplankton growth concentrations of plankton in the water, various
rates were estimated to fall in the range of 4-6 measures of seston concentration appear to be the
percent per hour in all parts of the bay. Such most ecologically significant indicators of nutrient
growth rates are close to the maximum growth enrichment in the system.
rates reported for marine phytoplankton grown on
light-dark cycles in continuous culture, suggesting Lee, G. F. and R. A. Jones (1981) Application
that phytoplankton growth rates (as opposed to of the OECD eutrophication modeling
biomass) were limited primarily by suboptimal or approach to estuaries. In: B. J. Neilson and
supraoptimal light intensities rather than by L. E. Cronin (ed.) Estuaries and Nutrients. Humana
nutrients. Based on these growth rates and an Press, Clifton, New Jersey.
assumed phytoplankton C:N ratio of 5.68 by Imodeling; lake; estuary}
weight, nitrogen recycling was estimated to account Approximately five years ago, the Organization
for 80 percent of phytoplankton nitrogen uptake in for Economic Cooperation and Development
the part of the bay receiving direct sewage inputs, (OECD) initiated a 22 country, 200 lake and
and for over 90 percent of phytoplankton nitrogen impoundment study of nutrient load-eutrophication
uptake in the other sectors of the bay. Estimates of response relationships. Emphasis in the study is
living and detrital particulate carbon were made being given to the evaluation of the Vollenweider
based on an assumed C: ATP ratio in living models for correlating nutrients load with
organisms of 285 by weight. From this eutrophication response. The U.S. part of this
partitioning, living carbon was found to vary by a study included the investigation of about 40
factor of 3-4 between the sewage-enriched and waterbodies or parts thereof. The results of the U.S.
unenriched sectors of the bay. However, estimated and the other studies all show a strong correlation
detrital carbon concentrations were uniform between the phosphorus load to a water body as
throughout the bay, as were the measured normalized by mean depth and hydraulic residence
concentrations of inorganic nitrogen. These results time, and the planktonic algal chlorophyll, Secchi
are consistent with the interpretation that the depth (water clarity), and the hypolinmetic oxygen
population of microorganisms, both bacteria and depletion rate. These relationships have been
phytoplankton, are substrate- limited in all sectors developed to a sufficient degree of sophistication so
of the bay. that they are the method of choice for estimating
the impact of altering the phosphorus load to a P-
Laws, E. A. and D. G. Reda1je (1982) Sewage limited waterbody on eutrophication-related water
diversion effects on the water column of quality.
a subtropical estuary. Marine Environmental While the U.S. OECD waterbodies were primarily
Research 6:265-279. lakes and impoundments, included as part of the
(phytoplankton; Kaneohe Bay; sewage; estuary; data base upon which the statistical correlataions
growth} were made were three parts of the Potomac estuary.
A study of the phytoplankton community and It appears that in general, the OECD eutrophication
water column chemistry in Kaneohe Bay, Oahu, modeling approach is applicable to estuarine
Hawaii, before and after the diversion of secondary systems as well as lakes and impoundments. In
treated sewage from the bay has shown that changes addition to reviewing the U.S. OECD study results,
in total nutrient concentrations in the water column this paper presents a discussion of the
cannot be accurately predicted without taking into modifications that may need to be made in the
account water column-benthos interactions. During OECD-Vollenweider eutrophication modeling
the first year after sewage diversion, the approach in order to apply it to some estuarine
decomposition of about 400 tonnes of benthic systems.
organisms, primarily filter feeders, resulted in water
column dissolved organic nitrogen and phosphorus Lee, V. and S. Olsen (1985) Eutrophication
concentrations roughly an order of magnitude and management initiatives for the
higher than those expected in the absence of such control of nutrient inputs to Rhode
interactions. The biomass of phytoplankton appears Island coastal lagoons. Estuaries 8:191-202.
to have been N limited both before and after sewage flagoon; management; macroalgae; nutrient; salt
diversion, with internal nutrient recycling pond; seaweeds; macroalgae)
accounting for 70-99% of phytoplankton nutrient An assessment of developing eutrophic conditions
uptake. Both the biomass and growth rate of the in small temperate lagoons along the coast of
phytoplankton declined as a result of the sewage Rhode Island suggests that in such shallow,
diversion, with post-diversion growth rates macrophyte based systems the response to nutrient
evidently well below nutrient-saturated values. enrichment differs from that described for
Since the principal stresses on the bay's coral reef planktonic based systems. The nitrogen loadings

per unit area of the salt ponds are 240-770 mmol N (Webster): A mesocosm study. Biological
per m2 per year. Instead of the high nutrient Bulletin 171:143-160.
concentrations, increased phytoplankton biomass Inutrients; reproduction; polychaete; MERQ
and turbidity, leading to eventual loss of benthic The influence of organic enrichment on growth
macrophytes described for such systems as the and planktontrophic development of the spinoid
Chesapeake, Patuxent and Appalachicola Bay, polychaete Streblospio benedicti Webster was
nutrient enrichment of the Rhode Island lagoons examined in two mesocosm experiments conducted
has led to increased growth of marine macroalgae. at the MERL facility, University of Rhode Island.
The increased macroalgal growth appears to alter Specimens of S. benedicti were collected and their
the benthic habitat and a shift from a grazing to reproductive traits monitored near the conclusion of
detrital food chain appears to be impacting a two-year eutrophication experiment. Nutrient (N,
important shellfisheries. As more extensive areas of P, and Si) enrichment at 8X and 32X the average
organic sediments develop, geochernical cycling aerial input into Narragansett Bay, Rhode Island,
changes, resulting in higher rates of nitrogen resulted in increases in body length, segment
re mineralization and accelerated eutrophication. number, and length per segment, and a doubling of
The major sources of nitrogen inputs to the salt brood size in S. benedicti females. These increases
ponds have been identified and a series of were substantially higher during May (120C) than
management initiatives have been designed to limit August (200C). Enrichment effects were stronger in
inputs from present and potential development the 8X than 32X nutrient treatment. In the sewage
within the watersheds of the lagoons. sludge experiment body size increased 20% over
control values at the highest (8X) sludge treatment
Lenzi, M. (1992) Experiences for the level (nitrogen loading equivalent to the 8X
management of Orbetello Lagoon: nutrient treatment) but no significant increase was
eutrophication and fishing. In: R. A. noted at the 4X sludge level, which received half as
Vollenweider, R. Marchetti and R. Viviani (ed.) much nitrogen as the 8X sludge treatment. Mean
Marine Coastal Eutrophication. Elsevier, brood size increased by a factor of 4.6 over controls
Amsterdam. in the 8X sludge treatment and by a factor of 2.3 in
Iseaweeds, fish yield} the 4X sludge treatment. Within the range of adult
The development of human activity has caused body sizes observed, brood size enhancement
hypertrophic conditions in Orbetello Lagoon. occurred independent of increased length or segment
Decisions are based on increasing water circulation in both nutrient and sludge enrichment treatments.
and on removal of part of the seaweed biomass. The ability to translate elevated food supply directly
Consequently, important changes in the trophic into increased reproductive output may underly
structure of the ecosystem have taken place, with a opportunistic dynamics in macrobenthos. Brood
significant increase in animal bomass. This fact size enhancement of the magnitude observed
has persuaded us that management of a lagoon probably contributes to the high S. benedicti
environment with high production is possible. densities found in polluted or organically enriched
Leppdkoski, E. (1980) Man's impact on the settings.
Baltic ecosystem. Ambio 9:174-18 1. Liaci, L. S. (1980) Pollution effects on
[Baltic; oxygen; review; benthos; abundance; marine biota. Memorie di Biologie Marina e di
compositionj Oceanographia Suppi.10:85-94.
The drainage from about 250 river systems and (organic; colored tides; dinoflagellates}
the municipal and industrial wastes of some 17 Effects of organic substance pollution on marine
million people sooner or later end up in the Baltic, organisms have been studied and related to
which has been described as one of the most eutrophication. The study refers effects of coloured.
severely polluted sea areas in the world. Human tides due to Dinoflagellates on mollusca and fishes.
activities have caused excessive eutrophication and The pattern of uptake and accumulation of some
resulted in damaging oil spills, and the chronic heavy metals was taken into consideration.
discharge of heavy metals and other toxic In fact, some specimens clearly showed that their
substances like PCBs which are seriously affecting uptake was directly proportional to the
the biomass of the Baltic. Even though the Baltic concentration of the heavy metal in the sea. Other
is one of the most thoroughly studied seas, much factor such as salinity, temperature etc. and their
remains to be done and the author concludes that we role on uptake, accumulation and elimination of the
will probably never fully understand it as a "natural metal are reported. Furthermore, the effects of
and man-influenced system". pesticide pollution has been examined with special
reference to the episode occurring in the harbour of
Levin, L. A. (1986) Effects of enrichment Brindsi where methaldehyde was spilt.
on reproduction in the opportunistic As a consequence of pollution, skeletal and gill
polychaete Streblospio benedicti deformities, as also defects of reproduction, growth,

62
number of eggs and epidermal papillomas and
hepatic tumors were observed. Alteration of the Lucht, F. and M. Gillbricht (1978) Long-term
biocenosis and involutions of ecological series due observations on nutrient content near
to organic slimes have been noted. Helgoland in relation to nutrient input of
the River Elbe. Rappots et Proc@s-verbaux des
Likens, G. E. (ed.) (1972) Nutrients and Rgunions
Eutrophication: The Limiting Nutrient Counseil International pour LExploration de la
Controversy (Proceedings of the Mer 172:358-360.
Symposium on Nutrients and [nutrients; Helgoland; River Elbe; phosphate;
Eutrophication: The Limiting Nutrient nitrogen; discharge; freshwater}
Controversy, W.K. Kellog Biological No abstract
Station, Michigan State University, ]I-
12 February, 1971). American Society of Maclean, J. L. (1984) Indo-Pacific toxic red-
Limnology and Oceanography and the Allen Press, tide occurrences, 1972-1984. In: A. W.
Lawrence, Kansas. White, M. Anraku and K.-K. Hooi (ed.) Toxic Red
No abstract Tides and Shellfish Toxicity in Southeast Asia.
Southeast Asian Fisheries Development Center and
LISS (1990) Status report and interim the International Development Research Center,
actions for hypoxia management. Long (toxic; red tide; blooml
Island Sound Study, U.S. Environmental No abstract
Protection Agency, [oxygen; hypoxia; modeling;
impacts; sewage; runoff; management; nitrogen; Maclean, J. L. (1989) Indo-Pacific red tides,
Long Island Soundl 1958-1988. Marine Pollution Bulletin 20:304-
No abstract 310.
(red tides; Indo-Pacific; dinoflagellate; blooml
Littler, M. M., D. S. Littler, and B. E. Lapointe A summary of recent toxic red tides in the
(1986) Baseline studies of herbivory and southwestern Pacific region is given, showing that
eutrophication on dominant reef the dinoflagellate Pytodinium bahamense var.
communities of Lboe Key National compressa remains the major causative organism of
Marine Sanctuary. NOAA, NOAA Technical paralytic shellfish poisoning and is apparently
Memorandum, NOS MEMD 1. continuing to spread around the region. Noticluca
(algae; community; nutrient; limitation) scintillans blooms occur in some countries and
Presently, only limited knowledge is available have been associated with mass mortality of fish.
regarding the dominant algal assemblages at Looe The raphidophyte Chattonella (=Hornellia) has
Key National Marine Sanctuary and the effects of been implicated in both fish and shrimp kills. A
herbivory and nutrient limitation in controlling the relationship with El Nino-Southern Oscillation
structure of these communities. The goals of this (ENSO) events and some similarities with recent
research are to: (1) provide a preliminary inventory plagues of the crown-of-thorns starfish (Acanthaster
and general distributional assessment of dominant planci) are discussed.
algal communities, (2) initiate pilot studies,of the
effects of nutrient enrichment and grazing on Maestrini, S. Y. and E. Gran6li (1991)
structure of the reef communities and (3) contrast Environmental conditions and
findings with other reef habitats for which ecophysiological mechanisms which led
comparable data are available. This research to the 1988 Chrysochromulina polyepis
accomplishes these goals. Increases our bloom: An hypothesis. Oceanologica Acta
understanding of ecological processes that influence 14:397-413.
dominant reef communities and provides data of use Ibloom; nuisance; nitrogen; Scandinavianj
in management decisions directed towards Information available on the 1988
preserving Looe Key National Marine Sanctuary in Chrysochromulina polyepis bloom in Scandinavian
a natural state. coastal waters is evaluated. Special attention is paid
to the early bloom stage and the sequence of both
LoCicero, V. R. (ed.) (1975) Toxic environmental and physiological mechanisms
Dinoflagellate Blooms (Proceedings of leading to the bloom, and a tentative, integrated
the First International Conference on scheme is given of causes which led to the observed
Toxic Dinoflagellate Blooms, Boston, 4- events.
6 November, 1974) (MIT Sea Grant Environmental conditions favourable for
Report MITSG 75-8). Massachusetts Science Chrysochromulina polyepis are considered to have
& Technology Foundation, Wakefield, resulted from: a) an exceptionally high runoff of
Massachusetts. nitrogen-rich water in winter and early spring; b)
No abstract strong mixing of the water column, immediately

63
followed by a long period of vertical stability and production of the toxin seems to be favoured by
stratification; c) a diatom bloom 30% larger than phosphorus deficiency, however, a large input of
usual, peaking around 20 March; and d) a second nitrogen in combination with a reduction in the
diatom bloom in mid April, largely dominated by phosphorus loading is believed to have contributed
Skeletonema costatum. These events combined to indirectly to the bloom, by changing the nutrient
cause the euphotic layer to become silicate- status of the coastal waters from being nitrogen- to
exhausted, phosphorus-poor and fairly nitrogen- phosphorus-limited.
rich. The lack of silicate prevented diatom growth.
Weak turbulence and strong stratification favoured Mdkinen, A., I. Haahtela, H. Ilvessalo, and J.
growth of non-siliceous, motile nanoplankton. Lehto (1984) Changes in the littoral rocky
C polyepis is believed to have outcompeted co- shore vegetation in the Seili area, SW
existing algal species by producing a toxin acting Archipelago of Finland. Ophelia Suppl.
first as a grazer repellent, while its cell density was 3:157-166.
still below 104 individuals 1-1. At the conjectured Imacroalgae; seaweeds; species composition; rocky
location of first appearance of the bloom (or at one shore; Archipelago}
of several locations), in the Skagerrak near Gullmar The littoral macrophyte communities were studied
Fjord, the grazing repellent forced a shift of grazing in 1968-69 by Ravanko at 23 rocky shores around
to other algae around the latter part of April. With Seili, SW archipelago of Finland. Four of these
increased population density of C. polyepis (106 shores were revisited in 1981-82. The most
cells 1-1), the toxin then became sufficiently noticeable finding was the altered species
concentrated to affect directly not only by grazers, composition in the area. The number of green algae
but all other organisms. From late April, the C. and phanerograms had increased and the number of
polyepis population increased free from grazing brown and red algae decreased. The macrophyte
and other losses. In conditions of light and nutrient biomass (fresh and dry weight m-2) had drastically
sufficiency, a high growth rate (corresponding to decreased because of the total lack of Fucus
0.8 div. d-1) was achieved. Actively growing cells vesiculosus.
were buoyant and remained in the upper part of the Mdkinen, A., J. KddriA, and M. Rajasilta (1988)
water column. Factors controlling the occurrence of
With increasing population density (107 cells I- Furcellaria lumbricalis (Huds.) Lamour
1), nutrient became exhausted and self-shading and Phyllophora truncata (Pallas) Zinova
decreased available light. By mid May, the growth in the upper littoral of the Archipelago
rate had decreased to zero, and the population started of SW Finland. Kieler Meeresforschungen,
to become senescent. C. polyepis deteriorated in Sonderheft 6:140-146.
physiological state, perhaps party because of self- (red algae; Furcellaria; Phyllophora; plankton;
poisoning, and this led to sinking. No further light; Mytilus; archipelago}
growth occurred, but cell densities showed higher The influence of eutrophication on the occurrence
values (several 107 cells V 1, with maximum of the red algae Furcellaria lumbricalis and
density between 5 and 10 in) because distribution Phyllophora truncata was studied in the sea area of
became patchy both horizontally and vertically. At Turku. Due to increased sediment load and
the end of May a significant part of the population planktonic production, light penetration in the
had died, and cell leakage produced an increase in water has decreased in the northern parts of the
dissolved organic matter. The resultant lower study area. Furthermore, the quality of the sea
turbidity and regenerated nutrients then allowed bottom has changed. Plots of I m2 (N=100) in the
other flagellates and some diatoms to resume upper littoral zone (0.5-6m) were studied by
growth. The remaining C. polyepis cells sunk to SCUBA diving, in order to investigate the factors
the pycnocline (15-20 in), which acted as a barrier controlling in the occurrence of the red algae. Two
and greatly retarded this sinking, by which means transects were situated in the eutrophicated area, and
the cells became concentrated, and produced the two in the reference area. In the plots, the
highest reported cell densities at the pycnocline. It percentage cover of each plant species and of
is assumed that the bloom later sedimented. Mytilus edulis was documented, and the Secchi disc
Since relevant environmental conditions for the visibility and depth was measured.
initiation of the bloom occurred over a very large The numerical data were analyzed by correlation
area (75.103 km2) and the dominant currents were analysis and stepwise multiple regression analysis
not altogether consistent with the spreading of the (BMDP2R). In the eutrophicated area, abiotic
bloom, polygenesis of the bloom should be factors (percentage cover of plain bottom and
considered a possibility. bottom quality) explained 81.28% of the variation
To what extent long-term global change and of red algae cover. In the reference area, the biotic
eutrophication might have contributed to this factors (total number of species and Mytilus cover
exceptional bloom remains unclear. Since the

in %) were the most important factors, explaining and temporal extent of bottom water anoxia has
66.4% of red algae cover. increased in response to higher phytoplankton
productivity due to anthropogenic nutrient loading.
Malone, T. C. (1984) Anthropogenic However, synoptic observations of environmental
nitrogen loading and assimilation conditions (river flow, temperature, salinity,
capacity of the Hudson River estuarine dissolved oxygen, and nutrients) and phytoplankton
system, USA. In: V. S. Kennedy (ed.) The from 1984 to 1988 indicate that oxygen depletion
Estuary as a Filter. Academic Press, Orlando. is not related to phytoplankton productivity on
fnitrogen; phytoplankton; production; regeneration; seasonal to interannual scales. Rather, nutrient
Hudson River; sewage; capacity} loading is linked to oxygen depletion and summer
The coastal plume of the Hudson River estuary anoxia via the spring accumulation of
receives inputs of new nitrogen of sewage origin phytoplankton biomass which is more than
from the lower estuary and of offshore origin from sufficient to fuel oxygen depletion. Thus,
adjacent coastal water. As a consequence of these interannual variability in the magnitude of summer
inputs, the plume is one of the most productive anoxia is primarily driven by climate factors which
coastal systems in the world's ocean. The extent to govern vertical density stratification (mixing).
which such high production reflects the input of
sewage-nitrogen depends on the capacity of Mambeke, F. v., M. A. Bianchi, and G. Cahet
phytoplankton to assimilate new nitrogen input to (1984) Short-term b "acterial reactivity of
the plume and on the magnitude of sewage-nitrogen nitrogen-enriched seawater of a eutrophic
input relative to coastal inputs, i.e., on the capacity lagoon. Estuarine, Coastal and Shelf Science
of the plume to "filter" inputs of new nitrogen. 19:291-301.
The importance of new nitrogen varies seasonally fbacteria; nitrogen; lagoon; growth; respiration}
relative to nitrogen recycled within the plume. We carried out, in September' 24 h experiment
Regenerated production increases from 10% of total to determine the potential rea-\Lity of bacterial
phytoplankton production during winter to a communities found in the sea wat@r i.n Arcachon
maximum of 80% during summer. Sewage- Basin (France). To create eutro ic systems,
nitrogen supports an average of 54% of new batches of seawater were enriched wit 50 mg I of
production during the spring bloom period (March- either ammonium chloride or amino aci Samples
r
May) compared to 121% during the subsequent were taken every 3 h. Quantitative measrements
period of stratification (June-October) and 221% were made of direct counts (AODC), biomass and
during winter (November-February). On an annual ATP content. The heterotrophic bacterial
t eir
'h,
basis, these estimates indicate that phytoplankton communities were defined in terms of t (
production has increased by ca. 30% in response to catabolic potentialities and 'specific' diversity.
the input of sewage-nitrogen. Bacterial heterotrophic activity was established b'
y
These patterns have important implications in measuring heterotrophic uptake and the
terms of cross-shelf transport and exchange. The mineralization percentage of labelled glucose or
supply of new nitrogen from offshore is highest amino acids. From these data, ratios of AMP/ATP
during March-May and, along with sewage- and P/B were calculated where P is the uptake of
nitrogen, appears to support a large seaward export labelled substrate and B is the biomass.
of nitrogen in the form of phytoplankton biomass. After the nitrogen enrichment an increase of the
In contrast, most new nitrogen is exported via respirat.ion was seen at first. After 6 h, a biomass
plankton fraction of phytoplankton food webs peak appeared associated with a continuous increase
during summer. Consequently, a relatively small in the number of cells. Bacterial growth was
fraction of phytoplankton production is available to concurrent with a qualitative modification of the
fuel oxygen demand below the pycnocline during community: UAI increased, and diversity dropped.
summer when assimilation capacity is greatest. A less diversified community resulted suggesting
an immature ecological state (zymogenous
Malone, T. C. (1991) River flow, bacteria). Low values of respiration percentage
phytoplankton production and oxygen (20%) characterized high growth yields.
depletion in Chesapeake Bay. In: R. V.
Tyson and T. H. Pearson (ed.) Modem and Ancient Marchetti, R. (1992)* The problems of the
Continental Shelf Anoxia. The Geological Society, Emilia Romagna coastal waters: facts and
London, England. interpretations. In: R. A. Vollenweider, R.
(oxygen; phytoplankton; Chesapeake Bay; climate; Marchetti and R. Viviani (ed.) Marine Coastal
anoxia) Eutrophication. Elsevier, Amsterdam.
Oxygen depletion leading to a summer anoxia in [gels, Adraitic Sea}
subpycnocline waters of the mesohaline reach of In this work, the two main problems concerning
Chesapeake Bay has been a annual event for at least the upper Adriatic coastal waters are examined: the
five decades. It is generally assumed that the spatial so-called algal blooms and the gels or excretions.

65
The former comprises the uncontrolled reporduction centimeters of sediment each year, resulting in
of microalgae causing an enormous mass of living production of H2S in the uppermost layer.
matter which is incompatible with bathing, the Originally, the benthic fauna was dominated (as per
decomposition of which causes oxygenc consuption numbers of individuals) by Nucula nitidosa and (as
and the death of the animals living on the sea per biomass) by Echinocardium cordatum and
bottom. This phenomenon is not new and is Ophiura spp. After 6-15 months of mussel
strongly connected with the polluting loads from culturing these and other species disappeared and
the Po and other minor rivers along the coast. The were replaced by opportunistic polychaetes
latter - in respect to the upper Adriatic coatal waters (Capitella capitata, Sclelepis fuliginosa, and
during the summers of 1988 and 1989 - is due to Microphthalmus sczelkowskii). The abundance of
the excretion of products which are normally these species fluctuated due to changes in the
emitted by algal cells with various functions and position of the redoxcline and their annual life
which, under circumstances still not completely cycle. The anaerobic sediments and mass occurrence
understood, can reach enormous amounts as of opportunistic polychaetes were localized to a
occurred in the summer of 1989. In the July of zone under and 5-20m around the cultures. After
that year, it was calculated that about 10,000 km2 harvesting of the mussels only limited recovery
of sea were covered by a layer of gel. The nature of was observed after one and a half year.
the compounds of which the gel is formed hs been
individuated in polysaccharides forming a structure McComb, A. J., R. P. Atkins, P. B. Birch, D. M.
incorporating 98-99% of water and which is Gordon, and R. J. Lukatelich (1981)
completely harmless for humans. The most Eutrophication in the Peel-Harvey
probable cause of the formation of these gels has estuarine system, Western Australia. In:
been identified with the strong meteorological and B. J. Neilson and L. E. Cronin (ed.) Estuaries and
climatic anomalies which characterized that period, Nutrients. Humana Press, Clifton, New Jersey.
traces of which have not been found in the data of 1green algae; Cladophora; phytoplankton; nutrient;
the preceding 100 years. Australia)
The conclusions of this analysis are that, The most obvious symptom of eutrophication in
although for the first of the two events adequate this estuarine system is a green alga, Cladophora,
action on the sources of contamination would have which was sparse in 1966 but now accumulates and
some probability of success, for the second there is rots on the shores. The work is part of a continuing
no reasonable counter-action to be proposed at study designed to asses the relationships between
present. nutrient input and the growth of Cladophora and
phytoplankton.
Matsuoka, K. (1989) Morphological features The system consists of two shallow basins,
of the cyst of Pyrodinium bahamense var. interconnected and linked to the ocean by a narrow
compressum. In: G. M. Hallegraeff and J. L. channel. It is fed by three rivers; 90 percent of river
Maclean (ed.) Biology, Epidemology and flow occurs during four winter months.
Management of Pyrodinium Red Tides. Fisheries Phytoplankton and water nutrient levels are low
Department, Ministry of Development, Manila. in summer, but high during and after an input of
(dino; Pyrodinium; cyst} river nutrients in winter. Nitrogen: phosphorus
Cysts of Pyrodinium bahamense var. compressum ratios, regression analyses, and nutrient limitation
and several similar species are compared with assays suggest that nitrogen is potentially limiting
special attention to differences in process characters. in summer and autumn, phosphorus in winter and
Keys to these species with and without references spring. The Harvey typically supports higher
to archeophyle features are provided. phytoplankton levels that the Peel. Diatom
populations may be replaced by blue-green in
Mattson, J. and 0. Linddn (1983) Benthic summer in the Harvey.
macrofauna succession under mussels, Cladophora forms detached sphere of branched
Mytilus edulis L. (Bivalvia), cultured on filaments and is only prominent in the Peel.
hanging long-lines. Sarsia 68:97-102. Changes in biomass and growth of confined
lbenthic; macrofauna; succession; culture; populations in the field, together with laboratory
sediments; Capitella; Nucula; opportunistic; experiments, show that growth occurs when
abundance; biomass} temperatures and light intensities are high, not in
Cultures of the mussel Mytilus edulis produce winter when water column nutrient levels are high.
accumulations of feces as well as mussels on the Water from between the algal spheres has increased
sea floor below the cultures. Changes in the levels of phosphorus compared with the water
sediment composition and benthic community column above, emphasizing the possible
structure under such cultures were studied over a importance of nutrient release from decaying
three-year period on the Swedish west coast. material below. It is suggested that phytoplankton
Deposition of organic matter created several are important in trapping water-column nutrients

66
during and after river nutrient input, and that aspects clearly explains the Ulva mass blooms by
subsequent Cladophora and phytoplankton growth the dynamic trapping of water masses in bays
depends on nutrient recycling. where tidal residual drift vanishes, and shows the
main responsibility to be the increase of nitrogen
McErlean, A. J. and G. Reed (198 1) Indicators inputs (from agricultural sources) in triggering the
and indices of estuarine overenrichment. recent magnification of a natural phenomenon.
In: B. J. Neilson and L. E. Cronin (ed.) Estuaries
and Nutrients. Humana Press, Clifton, New Jersey. Messner, U. and J.-A. v. Oertzen (1990) Recent
[indicator; index; estuary; eutrophicationj changes in the phytal zone of Greifswald
Indicators and indices of nutrient overenrichment, Bay. Limnologica 20:183-186.
which might be applied to the problems of JGreifswald Bay; submerged vegetation; coverage;
estuarine overenrichment, are reviewed from the plankton; light; red algae}
literature from the time of the national symposium The phytal zone of Greifswald Bay, a coastal
on eutrophication (1968) to the present. While water in the northeast of the GDR was studied from
many existing methods have been applied to spring to autumn in 1985. The submersal
estuarine work and some new ones have been vegetation was mapped, and it was found that the
developed, there do not appear to be a large number coverage of the bottom had decreased from 90%
of usable techniques for estuarine investigation. (Seifert, 1938) to about 10%. The authors consider
increasing plankton production and the deterioration
McLusky, D. S., M. Teare, and P. Phizacklea of light conditions at the bottom owing to
(1980) Effects of domestic and industrial eutrophication are the main causes of this decline.
pollution on distribution and abundance Quantitative studies of the macrofauna of the phytal
of aquatic oligochaetes in the Forth zone show that the zone covered by red algae
estuary. Helgoldnder Meeresuntersuchungen [mainly Furcellaria fastigiata (L.) Lamour in the
33:384-392. investigation area] is populated by a large number
fdistribution; abundance; Oligochaetes; Forth of species in various growth stages. The fauna
estuary; macrofauna; diversity} inhabiting the less ramified Potamogeton
The Forth estuary is a major estuary on the cast pectinatus L. and Zostera marina L. stands was less
coast of Scotland; it receives effluents from diversified. It can be assumed that the decline of the
domestic and industrial (petro-chemical and red algae stocks to insignificant remains has led to
distilling) sources. Following a study on the major changes in the food web and energy flow.
distribution of macrofauna of the intertidal areas in Although no quantitative data can yet be offered, it
relation to pollution (McLusky et al., 1978), this must be feared that loss of this habitat threatens the
paper is concerned with the distribution and continued existence of a herring population there.
abundance of aquatic oligochaetes and the small The dense bluegreen blooms in mid-summer may
polychaete Manayunkia aestuarina in relation to restrict the use of these waters by holiday makers
estuarine salinity, organic enrichment, and for bathing. The changes that have taken place thus
industrial effluent. In the most polluted parts of the seem to jeopardize many of the uses to which
estuary oligochaetes are the sole inhabitants of the Greifwald Bay has been put.
mudflats; in other less polluted flats they are very
abundant. In the least polluted parts the numbers of Millner, R. S. (1980) Pulp and paper mill
oligochaetes diminish as the numbers and diversity waste pollution in the Swale, a tidal
of macrofauna increases. Estimates of the channel on the east coast of England.
production of oligochaetes are given. Helgoldnder Meeresuntersuchungen 33:366-376.
(pulp; paper; oxygen; benthos; abundance)
Menesguen, A. (1992) Modelling coastal The effect of pulp and paper mill waste discharges
eutrophication: the case of French Ulva in the Swale, a tidal channel on the southeast coast
mass blooms. In: R.A.Vollenweider, R. of England, has been studied. A pulsing tidal
Marchetti and R. Viviani (ed.) Marine Coastal movement in the Swale results in effluent being
Eutrophication. Elsevier, Amsterdam. trapped within the estuary for up to 20 days. This
(Ulva, macroalgae) has resulted in adverse effects on the quality of the
During the last decade, the excessive proliferation water and sediment along a substantial part of the
of Ulva sp. (Chlorophyceae) during spring and estuary with the greatest effect occurring east of the
summer has been detrimental to recreational use of mill in the direction of the residual current. A
several beaches of Brittany (France). These reduction in the dissolved oxygen concentration
worldwide mass blooms are well known in occurred along a 13-kin stretch of the estuary with
eutrophicated lagoons (e.g. Venice, Tunis), but are mean values falling to 53% saturation near the
paradoxical along open beaches with a large tidal mill. High levels of organic matter in the sediment
range and low terrestrial nutrient inputs. Numerical reflect the deposition of material from the mill
modelling of the coupled physical and biological effluent stream and loss on ignition values of up to

67
12 % were found east of the mill, falling to 5 % has been found in data sets, indicating phosphorus
within 4 km northwest of the mill. The macrofauna as the primary stimulating algal growth nutrient
of the intertidal mudflats was typical of a stressed while, from a compositional point of view,
environment with a low number of species and diatoms always accounted for most of the
with a few being represented by large numbers of phytoplankton increase. A similar effect of diatom
individuals. The dominant species was the sabellid dominance has also been observed in autumn assays
polychaete Manayunkina aestuarina which reached conducted on a differently shaped community
peak numbers of over 1.5 million m-2 1.5 km east (represented by Chaetoceros spp. and Gymnodinium
of the mill. An increase in the total biomass near sp.) while nitrogen and phosphorus together
the mill is thought to be associated with organic stimulated phytoplankton growth.
enrichment from the mill wastes.
Montgomery, R. T., B. F. McPherson, and E. E.
Mills, C. A., S. I. Heaney, C. Butterwick, J. E. Emmons (1991) Effects of nitrogen and
Corry, and J. M. Elliott (1990) L a k e phosphorus additions on phytoplankton
enrichment and the status of Windermere productivity and chlorophyll a in a
charr, Salvelinus alpinus (L.). Journal of subtropical estuary, Charlotte Harbor,
Fish Biology 37 (Supplement A): 167-174. Florida. U.S. Geological Survey, Water-
Ifish; lake; trophic status; nutrient; enrichment; Resources Investigations Report 91-4077.
deoxygenation; feedingj f nitrogen; phosphorus; phytoplankton;
All English population of Artic charr, Salvelinus productivity; estuary; Charlotte Harbor)
alpinus (L.), are found in the Lake District The response of natural phytoplankton
(northwest England). There are at least four races of assemblages in a subtropical coastal plain estuary,
charr in Windermere, the largest lake in England; Charlotte Harbor, Florida, to inorganic nitrogen and
the North and South basins of the lake each contain phosphorus additions was determined from
two distinct races that spawn in autumn and spring measurements of relative changes in both the
respectively. The spring spawners in both basins uptake of carbon-14 and concentrations of
probably represent less than 15% of the total chlorophyll a. The effects of nitrate plus nitrite
population in the lake. nitrogen, ammonia nitrogen, and orthophorus
Changes in the population density of charr in the additions over a series of concentrations were
lake are described briefly and examined in relation evaluate through in situ experiments conducted
to the trophic status of the lake. Other factors that during periods of seasonally low and high river
could possibly affect the charr population are inflows. The responses to nutrient additions were
reviewed, especially the influence of climate evaluated for three different size fractions of
change. phytoplankton. Relative changes of phytoplankton
carbon uptake and chlorophyll-a concentrations
Mingazzini, M., A. Rinaldi, and G. Montanari were highly variable with regard to season,
(1992) Multi-level nutrient enrichment location, nutrient, and size fractions.
bioassays on Northern Adriatic coastal Within areas of the estuary characterized by lower
waters. In: R. A. Vollenweider, R. Marchetti salinities, phytoplankton exhibited a distinct
and R. Viviani (ed.) Marine Coastal seasonal pattern to additions of inorganic nitrogen.
Eutrophication. Elsevier, Amsterdam. Under seasonally high freshwater inflow,
(species abundance, nutrient enrichment, Adriatic phytoplankton show little response to inorganic
Sea, limitation) nitrogen additions, whereas under seasonally low
In order to evaluate the role of nutrients on inflow, phytoplankton responded to the inorganic
phytoplankton growth inNorthern Adriatic coastal nitrogen additions. The seasonally high freshwater
waters, nutrient enrichment bioassays at three inflow increased ambient inorganic nitrogen
different complexity levels of the experimental concentrations and water color. The high water
system were undertaken. The algal growth color greatly reduced light penetration in the water
responses both to inorganic (P and N) and to column and limited phytoplankton productivity.
organic (group B vitamins) added nutrients were The effect of nutrient additions in the higher
measured on Skeletonema costatum and salinity reaches of the estuary indicates that, under
Prorocentrum micans, which are considereed the normal conditions, these areas are continually
most representative species in local blooms within nitrogen limited. During periods of high freshwater
the two groups of diatoms and dinoflagellates, inflow during the summer months, the estuary can
respectively. The results obtained from the be divided conceptually into a low-salinity zone
simplest experimental level, in which the cultured where phytoplankton production is mediated by
algae were added to filtered sea water, are compared light availability, as determined by high water
with the superior levels in which the same species color, abd a high-salinity zone where
were naturally abundance in phytoplankton phytoplankton production is nitrogen limited.
communities of tested samples. A good agreement Seasonal nutrient concentration data and

comparisons among inorganic nitrogen inputs and the shores of the partially enclosed Tolo Harbour
observed phytoplankton productivity support the have created Hong Kong's first critical marine
experimental bioassay results. Each of these lines environmental problem, secondary sewage
of evidence indicates that, exclusive of seasonal treatment schemes having been inadequate to
riverine influences that affect light penetration of prevent this. Agricultural wastes (notably pig
the water column, nitrogen availability normally manure) compound the problem in Tolo Harbour,
limits phytoplankton production within the but virtually everywhere in the New Territories
Charlotte Harbor estuarine system. lowland water courses are polluted from this source
leading to the degradation of most enclosed bays.
Moore, C. G. and T. H. Pearson (1984) Sufficient evidence is at hand to understand local
Response of a marine benthic copepod long-term trends in environmental degradation.
assemblage to organic enrichment. Hong Kong's coastal water are polluted generally;
Proceedings of the Second International Conference Tolo Harbour, grossly so. Concern exists regarding
on Copepod, Ottawa, Canada. the safety of bathing beach water quality,
(benthic; copepods; assemblage; sewage; Scottish contaminated sea food, and the overall quality of
coast; density} other enclosed bodies of water in Hong Kong.
No abstract
Moshiri, G. A., N. G. Aumen, and W. G.
Mori, K. (1979) Effects of artificial Crumpton (1981) Reversal of the
eutrophication on the metabolism of the eutrophication process: A case study.
Japanese oyster Crassostrea gigas. Marine In: B. J. Neilson and L. E. Cronin (ed.) Estuaries
Biology 53:361-369. and Nutrients. Humana Press, Clifton, New Jersey.
(oyster; Crassostrea; gonads; development; Ibloom; dinoflagellates; fishkill)
spawning; fats; mortality; Matsushima Bay; Japan; Bayou Texar, Pensacola, Florida, is a bayou
growth} estuary which was advancing toward eutrophy, due
Matsushima Bay, one of the richest oyster-culture to nutrient loadings from various sources. The
areas in Japan, is subject to heavy artificial occurrence of characteristic symptoms of
eutrophication, mainly from the inflow of city and eutrophication, coupled with the closing of the
factory sewages. The physiological activity in the bayou for water contact recreation, led to the
oyster Crassostrea gigas in hanging cultures (i.e. initiation of an intensive seven-year study. Initial
suspended in wire-mesh cages) in this embayment results indicated that Carpenter's Creek is a prime
declined markedly with progressive development of source of most nitrogen species and that an
the gonads; this decline in activity coincides with exchange exists between dissolved phosphates and
eutrophic ation -induced accumulation of fatty those adsorbed onto sediments. Carbon fixation
material in the epithelia of the digestive respiratory rates varied, with the stations closest to Carpenters
substrates in order to maintain its increased Creek exhibiting the most productivity. It also
physiological needs. However, the fats are appeared that nitrates may be more important in
inefficient energy sources for the oyster and hence, controlling rates of carbon fixation than
during each spawning season, 50% mortality phosphates. An ensuing study directed at phosphate
occurs. In addition, such phenomena as exchange suggested that reducing conditions in the
overmaturation of the gonad and disturbance of the muds cause substantial release of iron-bound P04
lipid and steroid metabolism seem to accelerate this from the sediment-water interface followed by
mass mortality. On the basis of the results, the adsorption onto particles in the aerobic flocculent
author has compiled a schematic diagram which layer above the interface. This mechanism could
illustrates the possible causes of mass mortality allow sufficiently rapid exchange Of P04 between
among C gigas in an eutrophic environment such sediment and water resulting in a low, stable
as Matsushima Bay. concentration in the water as observed. Other
investigations involving bacterioplankton,
Morton, B. (1989) Pollution of the coastal dissolved glucose, and heterotrophic production
waters of Hong Kong. Marine Pollution indicated that algal primary productivity is the
Bulletin 20:310-318. major source of dissolved glucose. A final study
JHong Kong; Victoria Harbour; Tolo Harbour; demonstrated a paucity of benthic
oxygen; phytoplankton; algae; Ulva; red tides; macroinvertebrates, probably due to a graded
benthos; abundance; benthos; fish} suspension of particles at the sediment-water
I This paper reviews the available literature on interface and the absence of a sharply defined
Hong Kong's marine coastal pollution. The highly bottom. The zooplankton community exhibited a
urbanized and industrial cities of Victoria and low diversity while numbers of individual species
Kowloon pollute Victoria Harbour with untreated were high. The phytoplankton community was
domestic sewage and industrial effluents. The composed primarily of dinoflagellates, with
development of new cities, Shatin and Tai Po, on diatoms, cryptophytes, chlorophytes and

microflagellates occurring in lesser quantities. It monthly during the summers of 1975 and 1976 in
was also suggested that the importance of toxins several shallow north Florida coastal and estuarine
from algal blooms to fish kills may be greater than systems by inorganic carbon-14 uptake and
previously indicated. Specific recommendations phosphorus-32 bioassays. The results of these
made after the fourth year of study were nutrient enrichments experiments suggest that
implemented in most part and led to a substantial phosphorus is frequently more important than
improvement of water quality and the subsequent nitrogen in limiting phytoplankton productivity in
opening of the bayou to the public for recreational nearshore northern Gulf of Mexico. A multiple
use. regression mo 'del was constructed to determine
which combinations of environmental and nutrient
Murakami, A. (1973) Red tide in the Seto variables could explain the most variation of
Inland Sea. Information Bulletin on phytoplankton productivity in these coastal
Planktology in Japan 19:131-132. systems. The final regression model was:
[Japan; nitrogen; phosphorus; red tide; bloom) P.P. = 32.1 + 48.4 S.R.P. = 0.54 Salinity
In the past 10 yr, a serious situation of red tide where: P.P. is phytoplankton productivity in ugCl-
has developed in the Inland Sea. The probable cause 1 hr-1, S.R.P. is soluble reactive phosphate in ug-
is excessive eutrophication. In the Inland Sea, the atom P04 1-1, and salinity is part-per thousand.
concentrations of soluble inorganic N and P in
seawater have increased greatly compared with This model explained 64 percent of the variation in
concentrations found 20 yr ago. The only phytoplantkon productivity.
fundamental countermeasure of such red tide is to Nitrogen has been identified as the primary
eliminate the source of excessive eutrophication. limiting nutrient for phytoplankton in coastal
waters and it has been proposed that the removal of
Musayeva, E. 1. (1985) Mesoplankton near phosphorus from marine waste discharges will have
the Bulgarian Coast. Oceanology of the little impact on the control of eutrophication. The
Academy of the Sciences of the USSR 25:647- observation that phosphorus is important in
652. limiting phytoplankton productivity in these
tzooplankton; biomass; mesoplankton) coastal and estuarine areas suggests that water
The distribution of mesoplankton in Burgas Bay quality planning for the coastal zone is best done
in 53 bottle samples taken in October-November on a regional basis, with consideration given to
1982 is discussed. A drop in the total biomass of local nutrient cycling processes.
zooplankton from north to south can be traced on a
northern meridianal section (Cape Krotiriya to Cape Nakanishi, H., M. Ukita, M. Sekine, M.
Kaliakra), probably resulting from a decrease in the Fukagawa, and S. Murakami (1992)
eutrophication effect of the Danube in this Eutrophication control in the Seto Inland
direction. The plankton off the Bulgarian coast was Sea. In: R. A. Vollenweider, R. Marchetti and
in typical autumn condition. In the southern part of R. Viviani (ed.) Marine Coastal Eutrophication.
Burgas Bay, where there is a discharge current Elsevier, Amsterdam.
carrying eutrophicated sewage from the city of Ifish yield, Japan}
Burgas, various stages in the development of the The Seto Inland Sea, surrounded by the four large
community, from a young community in the inner islands of Honshu, Shikoku and Kyushu, is famous
end of the bay to a mature one at its outlet, were for its scienic beauty and coastal fishery. However,
observed. the beauty of this areas has been threatened by a
high concentration of human population and
Myers, V. B. and R. 1. Iverson (1981) industry which contribute to water pollution and
Phosphorus and nitrogen limited euthrophication problems.
phytoplankton productivity i n In this study, we examine the present water
northeastern Gulf of Mexico coastal pollution problems of this area; we have studied the
estuaries. In: B. J. Neilson and L. E. Cronin history of water pollution in the Seto Inland Sea,
(ed.) Estuaries and Nutrients. Humana Press, the influx of nutrients into all sections of this area
Clifton, New Jersey. and the relationship between euthrophic levels and
(nitrogen; phosphorus; limitation; phytoplankton; fishery productions in each section of the Seto
productivity; nutrients) Inland Sea. We have also analyzed the secondary
An understanding of nutrient limitation of organic pollution which is a result of the primary
estuarine phytoplantkon growth is important in photosynthesis production and the comprehensive
making environmentally sound decisions eutrhopic situation of the sea and each of its
concerning watershed development and the use of sections, in comparison with other coastal seas in
aquatic environments for waste disposal. Japan.
Experiments to determine nutrient limitation of From the results of this evaluation, we propose
phytoplankton productivity were conducted the following four strategies for eutrophication

70
control in the Seto Inland Sea: (1) Elimination of (nitrogen-, phophOTUS; nutrient; Baltic;
nitrogen (N) and phosphorus (P) discharging loads hydrographicl
by means of controlled chemical fertilizer Long-term nutrient trends were studied in the deep
consumption and recycling of human and livestock water (in general 1958-1982) and in the winter
excreta as fertilizer (as a conclusion of the budget of surface layer (in general 1958-1983) of the Baltic
N and P in Japan); (2) Consolidation of treatment proper. Phosphate accumulated in the deep water,
facilities for the point sources of pollutants such as on average, over the whole period under
domestic sewage and industrial waste water; (3) investigation, whereas nitrate has decreased
Diversion of the discharging N and P loads from significantly since 1979. The concentrations of
the Seto Inland Sea into the adjacent open sea aras these nutrients also increased in the winter surface
of Japan w;hich have a larger carrying capacity layer. The highly significant overall trends in this
through the exclusion system such as a sewer layer correlated closely with an increase in salinity.
system (through comparison with each adjacent sea The reasons for the long-term variations in
area of Japan); (4) Dredging of sediments over the nutrients and the consequences of the eutrophication
polluted zones. observed are briefly discussed.

Nalewajko, C. and M. A. O'Mahony (1989) Nehring, D. (1992) Eutrophication in the
Photosynthesis of algal cultures and Baltic Sea. In: R. A. Vollenweider, R.
phytoplankton following an acid pH Marchetti and R. Viviani (ed.) Marine Coastal
shock. Journal of Phycology 25:319-325. Eutrophication. Elsevier, Amsterdam.
JpH; acid; lake; phytoplankton; photosynthesis) (nitrogen, phosphorus, oxygen, Baltic Seal
In acidifying lakes, pH decreases abruptly in Eutrophication is identified as one of the most
response to acid precipitation events. We tested the serious problems in the Baltic Sea. Trend studies
hypothesis that in comparison to a circumneutral covering the period 1958-89 show increasing
lake, phytoplankton photosynthesis in an phosphate and nitrate concentrations in the surface
acidifying lake is less sensitive to a rapid decrease layer in winter and in the deep water below the
in pH (acid pH shock). permanent halocline. Consequences of the
Phytoplankton in Plastic Lake, which is eutrophication are discussed with respect to the
undergoing acidification, was characterized by a biological productivity and the determination of the
predominance of Pyrrophyta, and phytoplankton oxygen conditions in stagnant Baltic deep waters.
photosynthesis decreased to a lesser extent in Conditions for eutrophication in the Baltic Sea are
response to an acid pH shock than the compared with those in the Mediterranean Sea.
photosynthesis of populations from St. Nora Lake,
a circumneutral lake located nearby, in which Nehring, D., S. Schulz, and W. Kaiser (1984)
Pyrrophyta were not abundant. Rates of Long-term phosphate and nitrate trends in
phytoplankton photosynthesis in acid pH shock the Baltic Proper and some biological
experiments were significantly correlated with consequences: A contribution to the
hydrogen ion but not with dissolved inorganic discussion concerning the eutrophication
carbon (DIC) concentrations. of these waters. Rappots et Proc&-verbaux des
Depression of photosynthesis following an acid Rgunions
pH shock occurred in axenic cultures of Chlorella Counseil international pour LExploration de la
pyrenoidosa Chick but was not observed in axenic Mer 183:193-203.
cultures of the acidophilic alga Chlorella (Baltic; nutrients; nitrogen; phosphorus; fish;
saccharophila (Krug.) Nadson or in three species oxygen; primary production; phytoplankton;
isolated from Plastic Lake. However, the three zooplankton)
isolates were not acidophilic during growth. We The phosphate and nitrate levels in the winter
conclude that phytoplankton in acidifying lakes surface layers of the central Arkona, Bornholm, and
consists predominantly of species which are southern Gotland Seas roughly tripled from 1969 to
tolerant to acid pH for short periods (hours) but 1978 (phosphate: 0.2 to 0.6 umol 1-1; nitrate: 1.5
cannot grow at these pHs. to 4.0 umol V 1). In the deep water below the
halocline, long-term accumulation of these
Nehring, D. (1981) Phosphorus in the Baltic nutrients was also found in the Gda'nsk, Gotland,
Sea. Marine Pollution Bulletin 12:194-198. FArb, and Landsort Deeps. Ibis process was most
(Baltic Sea; phosphorus; Gotland Seal significant at a depth of 100 rn in the Gotland
No abstract Deep, where on average phosphate increased from
Nehring, D. 1984) The further development about 1 to 2.5 umol 1-1 between 1958 and 1978,
of the nutrient situation in the Baltic and nitrate from about 4 to 8 umol 1-1 between
proper. Ophelia Suppi.3:167-179. 1969 and 1978.

The biological effects of the nutrient increase on estuary, SW Netherlands, after it has been
the pelagic system of the Baltic are obviously low transformed into a salt-water lake. In summer 1978,
and masked by inadequate observation frequency or after a development of 8 years of opportunistic
by annual fluctuations. On the other hand the colonization by seeds and rhizomes, all potential
increasing yields of the Baltic fisheries do not seem habitats in the lake were occupied and covered 44
to be the result of the growing fishing effort alone km2 of sheltered sediment areas with a water depth
but are the consequence of higher biological of 0-2.5 in. After man-induced changes in
production. environmental conditions the eelgrass population
dropped down in 1980 to about half the 1978
Neilson, B. J. and L. E. Cronin (ed.) (1981) situation. The possible causes of this sudden
Estuaries and Nutrients. Humana Press, decline have been discussed, viz seed production and
Clifton, New Jersey. germination, water temperature, wasting disease,
No abstract transparency of the water and macro-algal
competition. The most plausible explanation for
Nielsen, A. and G. frtebjerg (1984) Plankton mass decline of eelgrass in Lake Grevelingen in
blooms in Danish waters. Ophelia, 1980 is associated with an increase in organic
Supplementum 3:181-188. matter deposition on the bottom, following a
(phytoplankton; primary production; nitrogen; change in nutrient loading of the lake, causing rapid
bloom} deoxygenation and toxification of the sediments,
In the past fifteen years there seems to have been and consequent die back of rhizomes and roots.
an increase in the occurrence of massive plankton Water temperature and salinity may have played a
blooms along the coast of the North Atlantic minor role in the eelgrass wax and wane.
Ocean, including the North Sea, the Kattegat and
the Belt Sea. Some of the plankton blooms have Nilsson, P., B. J6nsson, I. L. Swanberg, and K.
caused fish mortality, either directly by poisoning Sundbdck (1991) Response of marine
of the fish, or indirectly by suffocation due to shallow-water sediment system to an
oxygen deficiency. It is indicated in the paper that increased load of inorganic nutrients.
the increased pollution load on the coastal zone Marine Ecology Progress Series 71:275-290.
may play an important role in the development of (sediment; nutrients; microflora; production;
massive plankton blooms in Danish waters. biomass; composition; productivity; meiofauna;
bacteria; grazing; oxygen)
Niemi, A. (1972) Observations on An outdoor experimental system was used to
phytoplankton in eutrophied and non- investigate the effect of an increased load of
eutrophied archipelago waters of the inorganic nitrogen and phosphorus on the lower
southern coast of Finland. Memoranda trophic levels of the food web of a shallow-water
Societatis pro. Fauna et Flora Fennica 48:63-74. sandy sediment. Estimates of structural changes
lphytoplankton; Finland; biomass; composition) were based on relationships between the biomass of
A comparison is made of the summer course of autotrophs, heterotrophic bacteria and meiofauna
the biomass and the composition of the and their qualitative composition. Effects on the
phytoplankton populations of 3 archipelago zones functions of the sediment community were assessed
in clean and polluted conditions off the southern by measuring primary and bacterial productivity and
coast of Finland. In summer low phytoplankton meiofaunal grazing rates using radiolabelling, as
biomass follow a vigorous spring bloom in the well as measuring changes in oxygen and nutrient
oligotrophic areas. Strongly eutrophic waters are fluxes and carbon pools. The sediment system
characterized by high biomasses (primarily due to responded within 2 to 3 wk to the nutrient
the blue-green alga 0scillatoria agardhii) persisting enrichment. Meiofauna biomass increased, resulting
throughout the summer. In addition to in higher relative importance of oligochaetes and
phytoplankton biomass, the composition of harpacticoid copepods. Primary productivity
phytoplankton and the occurrence of certain increased faster than meiofaunal grazing, resulting
indicator group may be a good criteria of the quality in an increase of microalgal biomass by a factor of
and level of eutrophication. 4. Diatom and filamentous cyanobacteria were
favoured by the increased nutrient levels. The
Nienhuis, P. H. (1983) Temporal and spatial stimulated photosynthetic activity has a negative
patterns of eelgrass (Zostera marina L.) feedback on the producing sediment layer, which
in a former estuary in the Netherlands, was lifted off by oxygen bubbles entrapped in the
dominated by human activities. Marine mucus-rich top most layer. Stimulated growth of
Technology Society Journal 17:69-77. the filamenous green alga Enteromorpha clathrata
(eelgrass; submerged vegetation; Netherlandsl resulted in a biomass of ca 2 g C m-2 after 4 wk,
A review is given of the temporal and spatial which was more than 2 times the biomass of
patterns of the eelgrass population in Grevelingen microautotrophs living in the sediment. Bacterial

productivity responded only weakly to the nutrient
additions and the grazing pressure on bacteria was Nixon, S. W. (1981) Remineralization and
high during the whole 4 wk experiment. Meiofauna nutrient cycling in coastal marine
removed on average about 4 % of the microalgal ecosystems. In: B. J. Neilson and L. E.
biomass and 12 % of the bacterial biomass per day Cronin (ed.) Estuaries and Nutrients. Humana
in treatments with increased nutrient levels. Press, Clifton, New Jersey.
Although autotrophic organisms were more (nutrients; remineralization; cycling)
important carbon sources than bacteria for Our views of remineralization and nutrient cycling
meiofauna, the impact of grazing on in coastal marine ecosystems have changed
microautotrophs was small, whereas grazing may considerably over the last 30 years. The major trend
be important in regulating bacterial growth. has been an increasing appreciation for the
Measurements of ammonium, nitrate, phosphate complexity of processes involved, including some
and silicate flux showed that shallow-water marked changes in our perception of the association
sediment with a high primary productivity in the between bacteria and particulate matter in the sea.
top layer functions as a sink for inorganic Among the more recent development in this area is
nutrients. In terms of biomass, the addition of a growing awareness of the importance of the
nutrients led to a dominance of meiofauna. coupling between benthic and pelagic communities
However in terms of productivity, autotrophs in coastal waters. There appears to be a strong
predominated and thus, during the initial phase of linear correlation between the organic matter
increasing nutrient levels, the entire sediment produced in the overlying water and the amount of
system was dominated by autotrophic processes. organic matter consumed on the bottom in almost
all of the coastal environment for which annual data
Nishijima, T. and Y. Hata (1991) Growth are available. The large amount of organic matter
potentials of red tide phytoplankters in consumed by the benthos (perhaps 25-50 percent of
coastal seawater by AGP assay. Marine that produced) is associated with a large flux of
Pollution Bulletin 23:175-179. inorganic nutrients from the sediments to the
fred tide; algal bloom; growth potential; nutrients) overlying water. The stoichiometry of net benthic
The method for the preparation of algal inocula nutrient regeneration differs from that of pelagic
for AGP (algal growth potential) assay was studied, regeneration, however, and simple Redfield type
and the growth potentials and growth limiting models probably cannot be applied. The amount of
nutrients of red tide phytoplankters in coastal fixed inorganic nitrogen returned to the water across
seawater were determined. The red tide the sediment-water interface appears to be about
phytoplankters Heterosigma akashiwo, and half of that expected on the basis of the flux of
Skeletonerna costatum were used as assay phosphorus. This behavior, along with the fact that
organisms. When using AGP assay the growth an appreciable amount of organic matter in coastal
potentials were found to be dependent on the waters gets remineralized on the bottom,
preparation of the algal inoculm. Accurate growth contributes to the low NIP ratio that is
potentials were obtained only when sufficiently characteristic of these areas and may be responsible
nutrient- starved cells were inoculated into the for the observation that nitrogen is a direct
samples. Insufficient starvation of the cells resulted measurements of the flux of dissolved N2 across
in an overestimation. Based from the results of the the sediment-water interface indicate that
assay methods, the growth potentials of Osaka Bay denitrification is probably responsible for the loss
water were about 104 cells/ml for H. akashiwo and of fixed nitrogen during decomposition in the
about 105 cells/ml for S. costaturn. Moreover, the sediments. If this is a widespread phenomenon,
water was found to have a potential to develop to estuaries, bays, and other coastal waters may be
the density of red tide for both organisms. The major sinks in the marine nitrogen cycle and
growth limiting nutrients of the water for both red important terms in the global nitrogen budget.
tide organisms were determined to be nitrogen and However, the fact that eutrophication appears to be
phosphorus. an increasing problem in many estuaries is dramatic
warning that anthropogenic nutrient inputs can
Nixon, S. W. (1980) Between coastal overwhelm the recycling and remineralization
marshes and coastal waters - A review of processes in coastal waters.
twenty years of speculation and research
on the role of salt marshes in estuarine Nixon, S. W. (1982) Nutrient dynamics,
productivity and water chemistry. In: P. primary production and fisheries yields of
Hamilton and K. B. MacDonald (ed.) Estuarine and lagoons. Oceanologica Acta Supplirnent4:357-
Wetland Processes. Plenum Publishing Co., New 371.
York. flagoon; primary production; fish yield; lake;
Imarsh; estuary; nitrogen; phosphorus} nutrientl
No abstract

73
There is a great range in many of the of nutrient and sediment input and the chemical
characteristics of the lagoons which make up 13% composition of sediments accumulated in the
of the world coastline, including size (a few ha up estuary shows that Chesapeake Bay retains only
to 800,000), shape, annual temperature range, fresh some 3-6% of the nitrogen, 11-17% of the
water input, exchange with the sea, anthropogenic phosphorus and 33-38% of silicon brought into its
inputs, and human uses. While these differences waters during a year in which no major flood
appear to have a marked influence on the species occurred.
composition of lagoon ecosystems around the This behavior suggests that current problems of
world, there are also some remarkable similarities estuarine eutrophication are more a consequence of
in nutrient dynamics, the standing crop of present nutrient inputs than an inevitable or
phytoplankton, annual primary production, and inescapable legacy of past enrichment. It also
fisheries yields of many lagoons. For example, in follows that the management or manipulation of
many of the systems reviewed, the annual nutrient loadings to estuaries may lead to a more
phosphate cycle was similar in timing and rapid response in environmental quality than
magnitude, primary production amounted to some previously predicted.
200-400 g C m-2 yr-1, and fisheries landings were
on the order of 50- 100 Kg ha- I yr- 1. There appears Nixon, S. W. (1988) Physical energy inputs
to be an empirical correlation between primary and the comparative ecology of lake and
production and fisheries yields in a great variety of marine ecosystems. Limnology and
marine systems which may be similar in slope to Oceanography 33:1005-1025.
that found in large lakes. However, the yield of flake; estuary; physical energy; tide; nutrientl
marine systems at any given level of primary Although freshwater and marine systems both
production appears to be 10-20 times greater. The receive light and heat energy from the sun and are
yield of lagoon fisheries per unit area is commonly mixed by the wind, only marine systems receive
as high or higher than that from the most additional mechanical energy from the tide. This
productive coastal and offshore fisheries. input is very small relative to the flux of solar
energy but may exceed that from wind. Some
Nixon, S. W. (1987) Chesapeake Bay obvious. physical consequences of this additional
nutrient budgets - a reassessment. energy input include the development of intertidal
Biogeochemistry 4:77-90. habitats, the presence of stronger currents, and more
lChesapeake Bay; nutrient; budget; nitrogen; vigorous vertical mixing. It is argued that these
phosphorus; silicon; management) (and perhaps other) consequences lead to coastal
Recently published annual mass balances or marine ecosystems which differ in a number of
budgets for nitrogen, phosphorus, and silicon in important ways from temperate lakes.
Chesapeake Bay have pictured the estuary as There is some evidence that coastal marine
retaining a very large fraction, perhaps all, of the systems generally maintain a larger standing crop
nutrients that enter from land drainage, the of benthic animals and that these fauna are more
atmosphere, and anthropogenic discharges. effective in mixing the bottom sediments. As a
However, those budgets have been based on result of better sediment mixing (and perhaps
estimates of the net exchanges of nutrients at the warmer and better oxygenated bottom water),
mouth of the Bay or on the rates of accumulation organic matter deposited on the bottom of coastal
of nutrients and sediments calculated from the marine areas may be more completely metabolized
distribution of various geochernical tracers in the and less C, N, and P retained than in lake
sediments. While conceptually straightforward, the sediments. Materials are more tightly bound to
first approach is subject to large erTors because it particles, like many metals, may behave similarly
requires the determination of a small "signal" in lake and marine sediments. Although many lakes
against a large background of tidal "noise". The are strong sinks for nutrients and metals, marine
second approach has led to overestimates of the bays and estuaries may be much less effective in
nutrient trapping efficiency of the bay because retaining nutrients.
tracer-derived sediment deposition rates have been A major consequence of the input of tidal energy
multiplied by the surface area of the whole bay or appears to be a more intensive yield of fish from
various parts of the bay rather than by the smaller marine systems compared with temperate lakes.
area of active sediment deposition. This approach is The data suggest that this more intense yield is not
also incorrect because the average, long-term rates due to the size or interconnection of marine areas or
of sediment deposition measured by the to higher primary production, Rather, the efficiency
geochernical tracers, including major floods, have of transfer of primary production to fish appears to
been compared to shorter-term records of nutrient be greater. Tropical lakes appear more like marine
input. systems in this regard, and this may be related to
The more appropriate calculation of nutrient lower thermal stability and more effective wind
retention based on contemporaneous measurements

energy transfer because of a small Coriolis effect at the yield of fish form marine ecosystems developed
low latitudes. in Western Europe and

Nixon, S. W. (1989) An extraordinary red Nixon, S. W. and V. M. Berounsky (1984) The
tide and fish kill. In: E. M. Cosper, V. M. role of nitrification in contributing to
Bricelj and E. J. Carpenter (ed.) Novel low oxygen conditions in an urban
Phytoplankton Blooms Causes and Impacts of waterway. Rhode Island Water Resources Center,
Recurrent Brown Tides and Other Unusual Blooms. FY-1983.
Springer-Verlag, Berlin. Initrification; oxygen; Providence River}
(Narragansett Bay; red tide; sewage} No abstract
No abstract
Nixon, S. W., J. R. Kelly, B. N. Furnas, C. A.
Nixon, S. W. (1990) Quanifying the Oviatt, and S. S. Hale (1980) Phosphorus
relationship between nitrogen input and regeneration and the metabolism of
the productivity of marine ecosystems. coastal marine bottom communities. In:
Eighth Marine Technology Conference and K. R. Tenore and B. C. Coull (ed.) Marine Benthic
International Symposium for Ecology, Tokyo, Dynamics. University of South Carolina Press,
August 1990, and Shimane, September 1990, Columbia, South Carolina.
Japan. f phosphate; Narraganett Bay; flux; sediment)
(nutrients; productivity; relationship; estuary; In situ measurements of the exchange of
nitrogen) phosphate between sediment and the overlying
The concept that the input of dissolved inorganic water at three stations in Narragansett Bay, Rhode
nutrients, especially nitrogen, regulated the basic Island, showed that there was almost always a net
level of primary production and, hence, the yield of flux out of the sediments. The magnitude of the
fish from marine ecosystems developed in Western flux ranged from near zero in winter to almost 60
Europe and Scandinavia about a century ago. This umols M-2 hr-l in summer. The flux was strongly
"agricultural model" of marine production was correlated with temperature during the spring
conceived only after quantitative biological warming and did not decrease with increasing
sampling methods and analytical techniques for phosphate concentrations in the overlying water.
measuring nutrients in the sea became available Calculations indicate that the phosphate cycle in
during the late 1800s and early 1900s. coastal waters such as Narragansett Bay is
Verification of the model has been slow because dominated by sediment-water exchanges. Some 120
of difficulties in measuring the rate of nutrient mg-at of inorganic P m-2 yr-1 were released
input to marine areas, problems in measuring annually to the overlying water, or enough
primary production, lack of long-term phosphorus to support about 50% of the annual
comprehensive data, a system-specific view among phytoplankton primary production. The flux of
researchers that discourages broad comparisons, and dissolved organic phosphorus was erratic and lower,
a reluctance to undertake ecosystem-level and appreciable uptake as well as release was often
experiments in marine ecology. Widespread use of observed.
the 14C technique after about 1950, recent efforts Laboratory experiments using replicate cores
to develop nutrient budgets, and the completion of collected from the bay at different times of year
a long-term nutrient addition experiment using showed that oxygen uptake, carbon dioxide release,
marine mesocosms, have now provided data used in and phosphorus exchange by the sediments were
this report to quantify the broad, general also influenced by the availability of fresh organic
relationship between primary production and matter. It was clear from the laboratory and field
fisheries yields from the sea have only been measurements that the regeneration of organic
assembled in the past few years, but the evidence is matter by the benthos results in a return of
now compelling. Despite the great complexity of inorganic nutrients to the water column that is
marine ecosystems, the simple first-order anomalously low in fixed nitrogen relative to
relationship between nutrient inputs and phosphorus. This remarkable feature of benthic
productivity that was first conceived a century ago regeneration, along with the fact that a large
seems generally correct. amount of organic matter is decomposed on the
bottom in shallow areas compared with the open
Nixon, S. W. (1992) Quantifying the sea, appears to be responsible for the
relationship between nitrogen input and characteristically low N/P ratio of coastal marine
the productivity of marine ecosystems. waters and for the importance of nitrogen rather
Advanced marine Technology Conference, No.5, than phosphorus as a major limiting nutrient in
Japan, The concept that the input of dissolved these areas.
inorganic nutrients, especially nitrogen, regulated
the basic level of primary production and, hence,

75
Nixon, S. W., C. A. Oviatt, J. Frithsen, and B. Idiel; dissolved oxygen; metabolism; nitrogen;
Sullivan (1986) Nutrients and the nutrients; phosphorus; salt marsh; silica;
productivity of estuarine and coastal Narragansett Bay)
marine ecosystems. Journal of Limnological Simultaneous measurements of the diel patterns
Society of Southern Africa 12:43-71. of dissolved 02, pH-C02, dissolved organic
(nutrients; primary productivity; estuary; coastal; nitrogen, ammonia, nitrite, nitrate, dissolved
microheterotrophs; recycling; MERL) organic phosphorus, phosphate, and silicate were
Recent research on estuarine and coastal marine made on four occasions in a shallow salt marsh
ecosystems has revealed two particularly interesting embayment (Bissel Cove, Narragansett Bay, Rhode
things about nutrient and productivity. First is the Island, U.S.A.) that was closed to tidal water
observation that these areas are among the most exchanges for 24-h periods. In spite of high rates of
intensively fertilized environments on earth. Second community photosynthesis and respiration, there
is the common finding that much of the appeared to be little diel exchange in dissolved
characteristically high primary productivity of these nutrient concentrations, especially with respect to
shallow waters is supported by nutrients released or inorganic nitrogen. During summer and early fall,
recycled by pelagic and benthic microheterotrophs. inorganic nutrients appeared to cycle within the
Since nutrient inputs to coastal areas have probably sediment-detritus system of the embayment
been increasing and are likely to continue to do so, bottom, rather than being released to the overlying
it is particularly important to understand the water. This suggest that there would be little export
relationship between nutrient loading and nutrient of inorganic nutrients from the marsh during tidal
cycling and the extent to which their interactions exchanges with the estuary. The behavior of this
may set the levels of primary and secondary marsh embayment contrasts with reports from other
production in coastal systems. marsh areas where large net imports or exports of
That some direct relationship exists between the nutrients have been observed. The concentrations of
input of nutrients and the productivity of higher dissolved organic phosphorus and nitrogen were
trophic levels has been a principle of marine higher than inorganic forms, and showed
ecology since the turn of the century. It is substantial variation over each diel sampling
surprisingly difficult, however, to find quantitative period. The relationship between photosynthetic or
evidence showing that estuaries, lagoons, or other respiratory gas exchange and nutrient cycling in
coastal waters respond to eutrophication by waters influenced by benthic community
producing a larger biomass of animals. Part of this metabolism is subtle and complex, and cannot be
difficulty arises because the amount of nitrogen or adequately approximated by simple models such as
phosphorus incorporated in animal tissue is a very the Redfield ratio.
small term in the total nutrient budget of an
estuary, and the accuracy and precision of ecological Nixon, S. W. and M. E. Q. Pilson (1983)
field measurements may not be adequate to the task. Nitrogen in estuarine and coastal marine
In addition, the response of natural systems to ecosystems. In: E. J. Carpenter and D. G.
nutrient enrichment is compounded by changes in Capone (ed.) Nitrogen in the Marine Environment.
climate, hydrography, harvesting effort and Academic Press, New York.
technology, and pollution. Initrogen; estuary; budget; distribution)
Attempts to avoid these problems by carrying out No abstract
controlled nutrient addition experiments in the field
or with mesocosms have been much rarer in marine Nixon, S. W. and M. E. Q. Pilson (1984)
ecology than limnology. The results that are Estuarine total system metabolism and
available for such studies seem to suggest that there organic exchange calculated from nutrient
is a modest enhancement of primary production ratios: An example from Narragansett
with nutrient addition, but that most of this extra Bay. In: V. S. Kennedy (ed.) The Estuary as a
organic matter is rapidly consumed, presumably by Filter. Academic Press, Orlando.
microheterotrophs. In other words, as nutrient [Narragansett Bay; estuary; nutrient; denitrification;
inputs rise, so does the rate of nutrient recycling. net ecosystem metabolism; phosphorus; budget}
Only a small fraction of the added nutrients appears The average ratio of the concentration of dissolved
as an increment in the production of higher trophic inorganic nitrogen (DIN) to phosphorus (DIP)
levels. found in many estuaries differs from that in the
rivers, sewage, rain, and offshore waters that
Nixon, S. W., C. A. Oviatt, J. Garber, and V. Lee contribute nutrients to these systems. These
(1976) Diel metabolism and nutrient differences are largely the result of various
dynamics in a salt marsh embayment. biological processes which take place in the
Ecology 57:740-750. estuary, including denitrification, the net production
of organic matter, and the consumption and
remineralization of organic matter carried into the

76
estuary. Net production in this case represents the and the overlying water column were made in a
total amount exported, harvested, and removed shallow coastal lagoon on the ocean coast of Rhode
through long-term burial that is in excess of the Island, U.S.A. The release of ammonia from mud
amount imported and consumed within the estuary. sediments in the dark (20-440 umol per m2 per h)
It is not the same as the commonly reported net averaged ten times higher than from a sandy tidal
production by the autotrophic components of the flat (0-60 umol per m2 per h), and while mud
system. Together with physical mixing and sediments also released nitrate and phosphate, sandy
chemical exchange processes which take place in sediments took up these nutrients. Fluxes of
the system, these biological transformations nutrients from mud sediments, but not from sandy
provide a dynamic "filter" which often makes the areas, markedly increased with temperature.
quantity and form of nutrients exported from an Ammonia release rates for mud sediments in the
estuary quite different from those it imports. These light (0-350 umol per m2 per h) were lower than
changes can themselves provide insight into the those in the dark and it is estimated that some 25%
internal processes of the estuary. of the ammonia released to the water column on an
For Narragansett Bay (RI, USA) we have annual basis may be intercepted by the benthic
combined measurements of the annual nutrient microfloral community. Estimates of the annual
input with the ratio of the annual time and volume- net exchange of nutrients across the sediment-water
weighted mean concentrations of DIN and DIP interface, weighted by sediment type for the lagoon
found in the Bay, measurements of the annual rate
of denitrification, and data on the chemical as a whole, showed a release of 450 mmol per m2
composition of the plankton to calculate the total of ammonia, 5 mmol per m2 of phosphate, 5
system metabolism of the Bay. The result suggests mmol per m2 of dissolved organic phosphorus, and
that the system is autotrophic, with net production an uptake of 80 mmol per m2 of nitrate. Although
exceeding consumption by about 80 g C m-2 y- I . rates of ammonia and nitrate exchange were
Since only a small portion of this organic matter comparable to those described for the deeper
accumulates in the sediments (- 6 g C m-2 y- 1) or heterotrophic bottom communities of nearby
is removed in the fishery (- I g C m-2 Y-1), some Narragansett Bay, rates of benthic phosphate release
70-75 g C M-2 y- I may be exported from the Bay, were significantly lower. On an annual basis the
an amount equal to 22-24% of the reported Bay benthos released approximately 20 times more
production by the phytoplankton. inorganic phosphate per unit area than did the
lagoon benthos. As a result, the N/P ratio for the
Nixon, S. W., M. E. Q. Pilson, C. A. Oviatt, P. flux from the sediments was 74:1 in the lagoon,
Donaghay, B. Sullivan, S. Seitzinger, D. Rudnick, compared with 16:1 in "average" marine plankton
and J. Frithsen (1984) Eutrophication of a and 8: 1 for the benthic flux from Narragansett Bay.
coastal marine ecosystem - An The lack of remineralized phosphate in the lagoon
experimental study using the MERL is reflected in water column phosphate
microcosms. In: M. J. R. Fasham (ed.) Flows concentrations (always <lum) and water column
of Energy and Materials in Marine Ecosystems. N/P ratios (anual N/P = 27) and suggests that the
Plenum Publishing Co., New York. lagoon may show phosphate limitation rather than
tnutrients; phytoplankton assemblage; the nitrogen limitation commonly associated with
phytoplankton abundance; secondary production; marine systems.
MERLI O'Connor, D. J. (1981) Modeling of
No abstract eutrophication in estuaries. In: B. J.
NOAA (1990) Estuaries of the United Neilson and L. E. Cronin (ed.) Estuaries and
States Nutrients. Humana Press, Clifton, New Jersey.
Vital statistics of a national resource [nutrient; model; phytoplankton; estuary)
base. NOAA, A special NOAA 20th anniversary No abstract
report, jestuary; resource; review) O'Reilly and D. A. Busch (1984)
N .o abstract Phytoplankton primary production on the
northwestern Atlantic shelf. Rappots et
Nowicki, B. L. and S. W. Nixon (1985) Benthic Proc@s-verbaux des Rgunions
nutrient remineralization in a coastal Counseil International our LExploration de la
lagoon ecosystem. Estuaries 8:182-190. Mer 183:255-268. P
jammonia; nitrate; nitrite; phosphate; phosphorus; fAtlantic shelf; phytoplankton; primary
lagoon; Narragansett Bay; flux; N/P ratio) production; sewage)
In situ measurements of the exchange of Phytoplankton production of particulate and
ammonia, nitrate plus nitrite, phosphate, and
dissolved organic phosphorus between sediments dissolved organic carbon was measured using 14C
simulated in situ sunlight incubation method

during 23 survey of the Mid-Atlantic Bight, (silicon; nitrogen; phosphorus; evidence;
Georges Bank, and the Gulf of Maine. Annual phytoplankton; diatoms; flagellates)
phytoplankton production (particulate plus Diatom phytoplankton populations are the usual
dissolved) ranged between 260 and 470 g C m-2 yr- food for zooplankton and filter feeding fishes and
1 in various regions, which places this continental contribute in a direct way to the large fishable
shelf system among the most productive in the populations in coastal zones. Flagellates, on the
world. The highest rates of daily phytoplankton other hand, are frequently poor foods for most
production were consistently found off the coast of grazers and can lead to undesirable eutrophication
New Jersey in the sewage-polluted apex of the New effects. Arguments are presented that silicon is
York Bight, followed by the shallow, well-mixed often the controlling nutrient in altering a diatom
waters on Georges Bank. In general, high daily to a flagellate community. The alteration is
rates of primary production (I g C m-2 yr-1) were governed by the relative magnitudes of the natural
observed during most of the months sampled and fluxes of the nutrients nitrogen, phosphorus and
were not limited to the 'spring bloom'period. silicon to the receiving water body and the recycled
Available data indicate that microheterotrophs fluxes of nitrogen and phosphorus for oceanic,
above the thermocline (not the seabed) supply most estuarine and inland water bodies.
of the mineralized nutrients required by the Okaichi, T. (1987) Red tide problems in the
productive summer phytoplankton communities. Seto Inland Sea, Japan. In: T. Okaichi, D.
Size-fractionation of the 14C-Iabeled particulate M. Anderson and T. Nemoto (ed.) Red Tides:
organic carbon revealed that netplankton (> 20 um) Biology, Environmental Science, and Toxicology.
are major primary producers during the spring and Elsevier, New York.
fall blooms in the Mid-Atlantic Bight, on Georges (red tides; blooms; oxygen; abundance}
Bank, and in the Gulf of Maine; however, the With the development of industries along the
nanoplankton (< 20 um) arc responsible for most coast of Seto Inland Sea, Japan, aquaculture of
of the annual photosynthesis of organic carbon. On yellowtail, oyster and nori (laver) have been
the annual basis, the percentage of particulate suffering from outbreaks of red tides. During these
carbon productivity by nanoplankton increased from two decades 20 billion yellowtail worth about 20
the shallow to the deep water. Euphotic per cent billion yen were killed. Comprehensive surveys on
extracellular release (PER) of dissolved organic the cause of outbreaks of red tides are carried out by
carbon by phytoplankton ranged between 0 and 55 workers at Universities and Prefectures.
%, averaging 15 %, and increased from the shallow Countermeasures to avoid the fishkill are devised
to the deep waters of the continental shelf. No well- and the monitoring systems for red tides, PSP and
defined seasonal cycle in euphotic PER was evident DSP within Prefectures are organized by the Bureau
from our data. of fisheries.

Ochi, T. (1987) The development of anoxic Okaichi, T., D. M. Anderson, and T. Nemoto (ed.)
water and red tide associated with (1988) Red Tides: Biology, Environmental
eutrophication in Hiuchi Nada Inland Science, and Toxicology (Proceedings of
Sea, Japan. In: T. Okaichi, D. M. Anderson the International Symposium on Red
and T. Nemoto (ed.) Red Tides: Biology, Tides, Takamastu Perfecture, Japan, 10-
Environmental Science, and Toxicology. Elsevier, 14 November, 1987). Elsevier, New York.
New York. No abstract
fred tides; oxygen)
The mechanism of anoxic water mass formation Olenin, S. N. (1990) Benthic "desert" and
was investigated in the eastern part of Hiuchi-Nada. transitional ecological zone at the
It was found that the oxygen consumption by the bottom of the eastern part of the Baltic
sediment was unexpectedly small and that the Sea. Oceanology of the Academy of Sciences of
oxygen consumption by organic matter freshly the USSR 29:751-753.
deposited on the bottom from phytoplankton lbenthic; desert; Baltic; transition; macrofaunal
blooms, followed by frequent breakdown of the As a result of studies performed during 1981-1987
thermocline were important -contributing factors in on the slope and floor of the Gotland basin in the
the formation of anoxic water masses in Hiuchi- eastern Baltic Sea, a transitional ecological zone
Nada. was distinguished between the benthic "desert",
which is devoid of macrofauna and situated on the
Officer, C. B. and J. H. Ryther (1980) The floor and slope of the basin below 110- 120 in, and
possible importance of silicon in marine the upper sections of the benthic zone above 80-90
eutrophication. Marine Ecology Progress m, which has a comparatively rich and varied
Series 3:83-91. bottom populations. The zoobenthos of the
transitional zone is extremely poor and is

78
characterized by a predominance of mobile present day levels of sewage effluents. For a 32-fold
invertebrate forms. increase in nutrients, system apparent production
increased by only a factor of 3.5. Seasonal patterns
Olsen, P., M. Cohn, J. B. Mahoney, and E. Feerst of autotrophy during the winter-spring diatom
(1984) Ganyaulax excavata Monitoring in bloom and heterotrophy during summer and early
New Jersey. Coastal Ocean Pollution fall occurred at all treatment levels. With the
Assessment News 3:25-26. exception of the 8X treatment, all treatments above
fnuisance bloom; Gonyaulax; red tide; nutrient; 2X had a greater respiratory demand in the water
New York Bay; New Jerseyj column than the benthos. The highest treatment
No abstract mesocosm (32X) went briefly anoxic during the
second summer of the experiment during a period
Olsson, G. E., B. Sundstr6m, and L. Edler (1987) when little productivity was occurring in the water
In situ studies of the effects of humic column.
acids on dinoflagellates and diatoms. In:
T. Okichi, D. M. Anderson and T. Nemoto (ed.) Oviatt, C. A., P. Lane, F. F. III, and P. Donaghay
Red tides: Biology, Environmental Science, and (1988) Phytoplankton species and
Toxicology. Elsevier, New York. abundance in response to eutrophication
{blooms; humid in coastal marine mesocosms. Journal of
Experiments with 160 1 plastic bags were Plankton Research 11: 1223-1244.
performed in September 1986 in situ on the west Inutrient; species composition; phytoplankton;
coast of Sweden. One series of bags were filled assemblage; abundance; MERLI
with water containing the natural phytoplankton In a mesocosm nutrient enrichment experiment
community dominated by dinoflagellates (>80% of the species (or categories) and abundances of
the total phytoplankton biomass). The other series diatoms, dinoflagellates, flagellates, monads and
was filled with I um. filtered sea water inocculated ciliates were identified and counted over a 16 month
with the diatom Phaeodactylum tricornutum. period. Diatoms and ciliates increased with
Phosphorus or nitrogen was added alone or together increasing nutrient treatment while monads and
with commercially available humic acids to both flagellates, less than 10 um, did not. In some
series. Bags with no additions (C) or with only instances grazing controlled abundances to low
humic acids added (CH) were used as controls. P. levels in nutrient enriched treatments. By contrast,
tricornutum was stimulated by nitrogen addition, in the field diatoms sometimes appeared to decrease
while no positive response was obtained by any while small phytoplankton less than 10 um
nutrient addition to the dinoflagellate bags. appeared to increase under eutrophic conditions.
Dinoflagellate biomass increased several fold when While nuisance species were occasionally present in
humic acids were added together with nitrogen. various nutrient treatments, the intensity and
Also the alkaline biomass phosphatase activity frequency of their presence did not tend to increase
(APA) increased drastically both in the CH and with nutrient treatment. Generally species (or
nitrogen + humic acid bags. Tentative conclusions categories) did not appear to change with nutrient
are: 1) The dinoflagellates were inhibited by heavy treatment.
metals and the humic acids alleviated this
inhibition by working as chelating agents. 2) The Oviatt, C. A., J. G. Quinn, J. T. Maughan, J. T.
diatom was not inhibited by heavy metals because Ellis, B. K. Sullivan, J. N. Gearing, P. J. Gearing,
it could still grow quite well without humic acids. C. D. Hunt, P. A. Sampou, and J. S. Latimer
3) As APA increased in the dinoflagellate control (1987) Fate and effects of sewage sludge
bag with only humic acids we assume that nitrogen in the coastal marine environment: A
in the humic fraction to some extent was available mesocosm experiment. Marine Ecology
for the dinoflagellates. Progress Series 41:187-203.
1production; respiration; fate; nutrients; sludge;
Oviatt, C. A., A. A. Keller, P. A. Sampou, and L. oxygen; grazing; MERL)
L. Beatty (1986) Patterns of productivity A mesocosm experiment to assess the fate and
during eutrophication: A mesocosm effects of sewage sludge in the coastal marine
experiment. Marine Ecology - Progress Series environment was conducted over 4 mo. during the
28:69-80. summer of 1984 with 9 different treatments of
foxygen; net ecosystem production; MERLI sewage sludge and nutrient additions. Evidence from
In a 28 months mesocosm experiment, levels and settling rate studies, accumulations of carbon and
patterns of productivity and respiration were pollutant organics and ratios of A13C in the
observed for a range of nutrient additions selected to sediment indicated up to 83% of the sludge
provide a gradation from conditions in lower particulate inputs settled to the bottom and up to
Narragansett Bay, Rhode Island, USA, to 51% accumulated there. While no toxicity due to
maximum impact for an urban estuary receiving organics or metals was apparent, prolonged hypoxia

and even anoxia occurred in upper treatment levels. and to incorporate such criteria into the design of
At summer temperatures, sludge particulate inputs water quality management strategies applicable to
in excess of 1 g C m-2 d- I caused these hypoxic both coastal marine and freshwater habitats.
conditions. In contrast to low primary production
in low and medium sludge treatments during the Pagou, K. and L. Ignatiades (1988)
last 2 mo. of the experiment, the highest sludge Phytoplankton seasonality patterns in
treatment had high production. We attribute these eutrophic marine coastal waters. Biological
patterns to excessive grazing pressure from Oceanography 5:229-241.
zooplankton and benthic fauna in the lower 2 [phytoplankton; seasonality; Aegean Sea)
treatments and hypoxia-retarded grazing in the upper Phytoplankton seasonality patterns were examined
treatment. In the upper treatments, peaks of in an inshore eutrophic station fertilized
production stimulated peaks of excessive respiration continuously by domestic sewage. Data of certain
suggesting that respiration of sludge particulates physical (temperature), chemical (P-P04, N-N031
was enhanced or co-metabolized with the production and Si-S102 concentrations), and phytoplanktonic
of fresh organic material. (chi a, total cell, diatom, and dinoflagellate
concentrations) parameters were analyzed by time
Paerl, H. W. (1988) Nuisance phytoplankton series analysis (autocorrelation, cross correlation).
blooms in coastal, estuarine, and inland The phytoplanktonic and chemical parameters
waters. Limnology and Oceanography 33:823- exhibited significant seasonal variability-in spite of
847. the constant fertilization of the water-which was
(bloom; phytoplankton; marine; estuary; associated with the seasonal variability of
freshwater; cyanobacteria; dinoflagellates; review) temperature.
Multiple interacting physical, chemical, and
biotic factors, in proper combinations lead to the Parker, C. A. (1983) Localized hypoxia
development and persistence of nuisance algal recurs in the New York Bight. Coastal
blooms. Upon examining combinations of Ocean Pollution Assessment News 3:1-3.
environmental conditions most likely to elicit (oxygen; New York Bight)
nuisance blooms, commonalities and analog No abstract
situations become more apparent among coastal
marine (d i n o f I a g e I I a te - d o m i n a te d), Parker, C. A. and J. E. O'Reilly (1991) Oxygen
estuarine(di nofl agell ate- and cyanobacteria- depletion in Long Island Sound: A
dominated), and freshwater (eyanobacteria- historical perspective. Estuaries 14:248-264.
dominated) ecosystem. A combination of the foxygen; hypoxia; historical; Long Island Sound)
following hydrological, chemical, and biotic factors A retrospective analysis of available data was
will most likely lead to bloom-sensitive waters: a conducted to characterize the spatial distribution and
horizontally distinct water mass; a vertically temporal trends in dissolved oxygen (DO)
stratified water column; warm weather conditions, concentrations in Long Island Sound (LIS) over the
as typified by dry monsoon tropical climates and past four decades. A general cast-west gradient of
summer seasons in temperate zone; high incident decreasing bottom DO was evident in all historical
photosynthetically active radiation (PAR); enhanced data examined. In our review of data from the 1950s
allochthonous organic matter loading (both as DOC , collected by Gordon Riley and colleagues, and
and POC); enhanced allochthonous inorganic from contemporary surveys, we found no evidence
nutrient loading (nitrogen and/or phosphorus); of hypoxia (DO :@ 3.0 mg 1- 1) in the Eastern Basin;
adequate availability of essential metals, supplied however, in the deeper waters of the Central Basin,
by terrigenous inputs or upwelling; underlying there is some evidence for recent (1986) emergence
sediments physically and nutritionally suitable as of moderate hypoxia. The Western Basin
"seed beds" for resting cysts and alkinetes; algal- experienced episodes of hypoxia during the 1970s
micrograzer (protists and rotifers) synergism, which which became more recurrent and possibly more
also enhances nutrient cycling without severe in the late 1980s. The most severe,
consumption of filamentous and colonial nuisance persistent and chronically recurrent hypoxia
taxa; and selective (for non-nuisance taxa) activities occurred throughout the water column of the East
of macrograzers (crustacean zooplankton, larval River and in bottom water of the Western Narrows.
fish). An unprecedented episode of anoxia was observed in
Nuisance bloom taxa share numerous additional both the Western and Eastern Narrows regions of
physiological and ecological characteristics, LIS in 1987. Previously, anoxia occurred rarely,
including limited heterotrophic capabilities, high was short-lived, and was confined to the East River.
degrees of motility, and toxicity. Given such a set Statistical trend analyses revealed a significant
of commonalities, it would appear useful and increase in the summer minimum bottom DO in
timely to identify and address generally applicable the lower and middle reaches of the East River over
criteria for deeming a water body "bloom sensitive" the past 20 years. Beginning in 1981, however, DO

80
declined markedly in the adjacent Narrows bordering Journal of Experimental Marine Biology and
the Nassau County nearshore. The improvements Ecology 20:1-41.
in East River water quality over the previous 15-20 (sediments; benthic fauna; species; anoxic;
years appear to have been gained at the expense of effluent)
poorer water quality in the western sound. Changes are described in the benthic fauna of
Mechanisms potentially responsible for the recent Lochs Linnhe and Eil in response to organic
decline the bottom DO in western LIS are enrichment of the loch sediments, brought about by
suggested. the input of effluent material from a pulp and paper
mill. Fluctuations in the amount of effluent
Pastorok, R. A. and G. R. Bilyard (1985) discharges are related to subsequent successional
Effects of sewage pollution on coral-reef changes in the fauna. As organic enrichment
communities. Marine Ecology Progress Series proceeds, there is progressive elimination of
21:175-189. species, accompanied by an increase in the numbers
Isewage; coral reef, lagoon; species composition; of the surviving species, Excessive organic
abundance; benthic algae; primary production; deposition leading to anoxic conditions at the
nutrient; oxygen; Kaneohe Bay; turbidity; sediment/water interface results in the elimination
sedimentation} of all but a few annelid species. The observed
Sewage pollution is an increasing problem in successional changes are discussed in the light of
tropical marine environments. In this review we recent hypotheses concerning ecological changes in
synthesize present knowledge of the effects of relation to environmental stress.
sewage pollution on coral-reef communities, and
suggest directions for future research. A wide range Pearson, T. H. (1990) Marine pollution
of sewage impacts on coral-reef communities has effects of pulp and paper industry wastes.
been reported. Little or no impact has been Helgoldnder Meerestuntersuchungen 33:340-365.
observed on some reefs in well-flushed waters that (pulp; paper; oxygen demand; benthic; abundance;
receive small quantities of effluent, whereas large sediment; composition}
discharges of effluent into poorly-flushed lagoons The scale of waste discharges to the marine
and bays have caused major changes in species environment from the pulp and paper industry in
composition and abundance. The 3 components of various parts of the world is outlined and a brief
sewage effluent most detrimental to coral description of the major characteristics of such
communities are nutrients, sediments, and toxic wastes is given. The information available on the
substances. Nutrient enrichment by sewage effluent direct toxicity of these wastes to marine fauna and
may enhance bentbic algal biomass and primary flora is assessed including both lethal and sublethal
production in the water column. Increased primary effects. The environmental impact of waste
production in the water column favors benthic discharge and subsequent ecosystem modifications
filter-feeding invertebrates which, with the benthic are considered in detail; the complex adjustments to
algae, may out-compete corals and other reef- the nutrient and carbon budgets of the water column
building organisms. Anthropogenic inputs of and sediment involved in direct alterations to their
dissolved nutrients and organic particulate matter oxygen balance and other indirect eutrophication
may also depress oxygen levels. While heavy effects are described. It is concluded that whereas the
sediment loads on corals may be lethal, lesser direct toxicity of these wastes has minimal effect in
quantities may inhibit growth, cause changes in the the marine environment the complex problems
growth forms of colonies, decrease coral cover, alter created by increasing the oxygen demand on the
species composition of reef-building organisms, receiving waters can have considerable impact in
and inhibit coral recruitment. Toxic substances may inshore areas. Methods of minimizing the effect of
induce metabolic changes in corals, decrease rates of such impacts and of forecasting their extent are
growth and reproduction, or reduce viability of outlined.
corals. Although further research is needed on all
three major components of sewage effluent, the Pearson, T. H. (1987) Benthic ecology in an
most critical need is for comprehensive, long-term accumulating sludge-disposal site. In: J.
studies of sewage impacts. The combined effects of M. Capuzzo and D. R. Kester (ed.) Oceanic
particulate and toxic substances are especially Processes in Marine Pollution. Vol.]. Biological
poorly documented at this time. Processes and Wastes in the Ocean. Robert E.
Krieger Publishing Company, Malaba, Florida.
Pearson, T. H. (1975) The benthic ecology lbenthic; sludge; Eh; sediments; pH; abundance;
of Loch Linnhe and Loch Eil, a sea-loch biomass; species; annelids; nematods; deposit-
system on the west coast of Scotland. feederl
IV. Changes in the benthic fauna The distribution of benthic fauna in the area of the
attributable to organic enrichment. sludge-disposal site in the Firth of Clyde, Scotland,
is related to the gradient of increasing organic

enrichment created by dumping. Faunal succession The influence of effluents from cellulose
in relation to enrichment is related to changing industries on the benthic faunas of comparable
productivity along the gradient. Community fjordic systems on the west coasts of Sweden and
changes ar6 ascribed to progressive maximization of Scotland is assessed. Generalizations are made
carbon utilization as input levels increase; this about sequential changes in the interrelationship of
involves responses at many levels of organization. benthos and sedimentary structure. Estimates are
It is concluded that the benthic ecosystem has fully made of the rates of such changes along a gradient
adapted to the continuously high organic carbon of environmetal stress brought about by the organic
input. enrichment of marine sediments and these can be
used to assess the probable effect of large inputs of
Pearson, T. H., A. B. Josefson, and R. Rosenberg organic material to marine sediment. The
(1985) Petersen's benthic stations advantages and disadvantages of using indicators
revisited. 1. Is the Kattegat becoming species, comparing faunal groups, and using
eutrophic? Journal of Experimental Marine diversity indices in the description of ecological
Biology and Ecology 92:157-206. changes are briefly considered.
(benthic; macrofauna; abundance; biomass;
increase; decrease; composition; Balticl Pearson, T. H. and R. Rosenberg (1978)
The macrobenthic fauna present at the stations Macrobenthic succession in relation to
sampled by C.G.J. Petersen in 1911-1912 in the organic enrichment and pollution of the
Kattegat were re-assessed in 1984 using equipment marine environment. Oceanography and
and techniques similar to those used in the original Marine Biology: Annual Review 16:229-311.
surveys. The total biomass of organisms present in (macrobenthos; benthic; organic enrichment;
1984 was found to be significantly reduced in the sediments; abundance; biomass; spatial; temporal;
north and west of the area and in areas shallower diversity; similarity; indicator; opportunistic}
than 25 m. These changes were largely attributable No anstract
to decreases in the populations of large echinoid,
Echinocardium. When echinoids were excluded, Pearson, T. H. and S. 0. Stanley (1979)
significant increases in the biomass of other Comparative measurement of the redox
organisms present in 1984 were found to the east of potential of marine sediments as a rapid
the area and in areas deeper than 25 m. In general means of assessing the effects of organic
the majority of species were found to be smaller in pollution. Marine Biology 53:371-379.
individual size in 1984 when compared with the [sediments; redox potential; macrofauna; benthos;
earlier surveys. Dominance comparisons based on abundance; species; biomass)
biomass changes showed that deposit-feeders had A rapid method for measuring the redox potential
decreased and suspension-feeders and carnivores had (Eh) values of marine sediments is described. This
increased in dominance. Most notably method has been used to relate changes in the areas
Echinocardium had decreased and the ophiuroid of highly reduced sediment in a sea loch system to
Amphiura filiformis (Mijller) had increased in the continuously fluctuation inputs of organic
dominance at over 70% of the stations sampled. In waste from a pulp and paper mill. Subsequent
general ophiuroids and annelids had increased and faunal changes in these areas can be related to
echinoids and molluscs had decreased in dominance. changes in the measured sedimentary redox level.
Comparisons of similarity values indicated that The method is suggested as a rapid means of
community composition has changed markedly assessing the potential impact of an additional
over the area, with only =30% similarity of species organic input to a marine sediment.
between 1911-1912 and 1984 at most stations.
The possible causes underlying these general Pease, B. C. (1977) The effect of organic
changes are considered, amongst which changes in enrichment from a salmon mariculture
fish predation pressures, the direct effects of facility on the water quality and benthic
trawling and long-term temperature fluctuations are community of Henderson Inlet,
suggested. Whilst such factors may be partially Washington. Dissertation for Ph.D., University
responsible, the most probable causes is considered of Washington.
to be a general eutrophication of the area. Imariculture; benthos; infauna; epifauna;
sediments)
Pearson, T. H. and R. Rosenberg (1976) A The effect of fish excretion and respiration at a
comparative study of the effects on the salmon mariculture facility on the local water
marine environment of wastes from quality was evaluated. In the immediate vincinitiy
cellulose industries in Scotland and of the facility, dissolved oxygen concentrations
Sweden. Ambio 5:77-79. were consistently depressed while ammonia
(benthos; abundance; diversity; indicator} concentrations were increased, but neither effect was
significant based on accepted criteria. The

82
mariculture facility had no apparent effect on the Perry, D. M., J. B. Hughes, and A. T. Herbert
local phytoplankton population. Major fluctuations (1991) Sublethal abnormalities in
in the local water quality were a result of natural embryos of winter flounder,
hydrographic conditions and phytoplankton Pseudopleuronectes americanu,s, from
activity. Long Island Sound. Estuaries 14:306-317.
The effect of fish feces and excess fish food from [abnormalities; winter flounder; disease; Long
the mariculture facility on the benthic environment Island Sound; Boston Harbor)
was also evaluated. There was a definite Sublethal abnormalities were examined in
accumulation of organic matter under and extending developing embryos of the winter flounder,
at least 15 rn from the salmon pens. This organic Pseudopleuronectes americanus, as part of a study
accumulation caused significant changes in the of its early reproductive success in Long Island
structure of the benthic epifaunal and infaunal Sound and two sites in Boston Harbor. These sites
communities. Epibenthic mats of the sulfide- represented varying levels of anthropogenic
oxidizing bacterium, Beggiatoa, were observed contamination that were possibly affecting
under salmon pens along with an increased reproduction, as ascertained by chromosomal
abundance of several large predatory species ( abnormalities and lowered developmental rates in
Cancer gracilis, Lepidopsetta bilineata, and the embryos. Effort was focused on examination of
Myoxocephalus polyacanthocephalus). The benthic blastula and tail-bud embryos from field-captured
infauna community under the salmon pens declined females spawned at the laboratory. Abnormalities
in species richness and became numerically observed included evidence of cytotoxicity and
dominated by the polychaete worm, Capitella chromosome damage. Embryos of fish from New
capitata, a well-known ubiquitous indicator of Haven were usually the most aberrant, while
organic pollution. Therefore, the mariculture embryos from other sites, notably Hempstead,
facility would have little or no effect on the marine Shoreham, and both Boston Harb or stations,
environment if the settleable solids were collected showed subtle indications of abnormality.
and removed, or depressed in such a way that there
was no accumulation in the underlying sediments. Persson, L.-E. (1987) Baltic eutrophication:
A contribution to the discussion. Ophelia
Pekkari, S. (1973) Effects of sewage water 27:31-42.
on benthic vegetation. 0 1 K 0 S [Baltic; macrozoobenthos; abundance; diversity}
Supplymentum 15:185-188. Quantitative sampling of macrozoobenthos has
Iseaweeds; sewage; benthic; abundance; abnormal) been performed in the southern Baltic in the 1920s
The influence of sewage water on b6nthic and the 1950s. Some of these stations (43) were
vegetation was studied in the Stockholm revisited in 1982/83. The total abundance and
Archipelago and in the Gulf of Bothnia. The most biomass was significantly increased in 1982/83
conspicuous effects of sewage effluents in the Gulf compared to the 1950s. It is suggested that the
of Bothnia are the excalves of marine and freshwater main part of the biomass increase might be due to
vegetation distinctly differing from the vegetation reduced stocks of predatory flatfishes in the 1920s
in the vicinity. Disappearance and abnormal shape and 1930s.
of plants are conspicuous effects of sewage water in The most conspicuous difference in the
the inner part of the Stockholm Archipelago. comparisons is the widespread distribution of the
polychaete Capitella capitata in 1982/83. This is
Perciasepe, R. (1992) Progress Report of the attributed to increased trawling activities in recent
Baywide Nutrient Reduction decades and not to organic enrichment.
Reevaluation. Chesapeake Bay Program,
fabundance;oxygenj Phillips, D. J. H. and S. Tanabe (1989) Aquatic
In 1987, the parties to the original Chesapeake pollution in the Far East. Marine Pollution
Bay Agreement signed a new Chesapeake Bay Bulletin 20:297-303.
agreement. The 1987 Chesapeake Bay Agreement JFar East; algal blooms; oxygen; red tidesl
set a specific goal-to achieve at least a 40 percent An overview of the problems experienced in the
reduction of nitrogen and phosphorus entering the Far East relating to aquatic pollution is presented,
mainstern Chesapeake Bay by the year 2000. with emphasis on the quality of coastal waters. The
Although this progress report's fingdings are pollution problems faced by developing and
preliminary, trends and generalizations of nutrient developed nations in the region tend to vary in type
loads, water quality, and hibatat improvements are according to socioeconomic factors. The appropriate
becoming evident. Most of the background studies treatment and disposal of sewage remains a
for the Nutrient Reduction Reevaluation have been particular problem in the developing nations.
drafted. To date, seven model runs have been Industrial and agricultural effluents also exert
completed for use in this report. measurable effects on marine resources in many of
these regions. The environmental impacts of major

83
engineering and development projects are also a by the equation ks = 0.72 + 0.26c . The water
cause for concern in certain areas. In the developed column oxygen demand of the harbour was
nations, the impacts of conservative contaminants determined experimentally and revealed a significant
(especially the highly persistent and toxic dependence on oxygen concentration. It was
organochlorines) are considered to be the most estimated that over 80% of the oxygen supplied to
significant problem at present. The region as a the harbour was used within the water column. The
whole exhibited a great dependence on seafood as a sediment oxygen consumed about 18% of the
primary or preferred source of protein, and the oxygen entering the harbour and was relatively
protection of coastal waters is therefore of vital most important in the early stages of stratification
concern. In addition, tourism provides much-needed when the hypolimnetic dissolved oxygen
foreign exchange in many countries of the region, concentrations were high. The main sources of
and the tourist facilities are mostly situated on the oxygen were atmospheric reaeration (80%), lake-
coast. Additional regulatory initiatives are required harbour exchange (10%) and photosynthesis (10%).
in many areas, coupled to strong legislative action
and robust monitoring of coastal pollution, if the Portnoy, J. W. (1991) Summer oxygen
vital marine resources are not to be irreversibly depletion in a diked New England
damaged. estuary. Estuaries 14:122-129.
(oxygen; estuary; organic; fish kill; marsh}
Phoel, W. C., K. L. Webb, and C. F. D'Elia The diked and freshened Herring River estuary
(1981) Inorganic nitrogen regeneration (Wellfleet, Massachusetts) experiences regular
and total oxygen consumption by the summer hypoxia and one- to three-week period of
sediments at the mouth of the York main stream anoxia, often accompanied by fish
River, Virginia. In: B. J. Neilson and L. E. kills. Stream hypoxia results from the temperature-
Cronin (ed.) Estuaries and Nutrients. Humana dependent increase in oxygen demand of organic
Press, Clifton, New Jersey. matter released by diked salt marsh deposits; periods
(nitrogen; regeneration; oxygen} of total anoxia are induced by heavy rains which
In situ measurements of inorganic nitrogen fluxes increase the runoff of wetland organic matter.
and riverbed oxygen consumption were made on Historic reductions in tidal flushing have extended
sediments in 3, 9, and 16 rn of water at the mouth the low salinity region of the estuary normally
of the York River during stratified and destratified characterized by high organic loads and minimal
water conditions. Ammonium was regenerated, the flushing. Recurrent main stream anoxia has
rate of which increased with depth and oxygen depressed both migratory and resident aquatic fauna.
concentration in the overlying water. Nitrate and
nitrite fluxes from the sediment were minimal or Porumb, F. (1992) On the development of
non-existent during stratification at the 16-m Noctiluca scintillans under
station but increased and the nutrients were taken eutrophication of Romanian Black Sea
up by the sediments under destratified conditions. waters. In: R. A. Vollenweider, R. Marchetti
At the 3-m station, which is above the halocline and R. Viviani (ed.) Marine Coastal
when developed, nitrate and nitrite appeared to be Eutrophication. Elsevier, Amsterdam.
the major forms of nitrogen being released by the (species, abundance, noctiluca scintillans, Black
sediments. Oxygen consumption by the riverbed at Sea}
the 16 and 9-m stations was higher during the This paper analyses the development of Noctiluca
increased oxygen tensions associated with vertical scintillans during the 1980s and 1990s as a
destratification. The 3-m stations maintained the consequence of the intense eutrophication of the
higher rates of oxygen consumption throughout the Romanian waters of the Black Sea.
sampling period. The in situ incubation of bottom
water alone at all three stations indicated negligible Price, K. S., D. A. Flemer, J. L. Taft, G. B.
rates of oxygen uptake. Mackiernan, W. Nehlsen, R. B. Biggs, N. H.
Burger, and D. A. Blaylock (1985) Nutrient
Polak, J. and G. D. Haffner (1979) Oxygen enrichment of Chesapeake Bay and its
depletion of Hamilton Harbour. Water impact on the habitat of striped bass: A
Research 12:205-215. speculative hypothesis. Transactions of the
foxygen; eutrophic; demand; sediment} American Fisheries Society 114:97-106.
Processes involved in the oxygen cycles of the (Chesapeake Bay; fish; striped bass; oxygen;
highly eutrophic Hamilton Harbour were studied. habitat; nitrogen; phosphorus; aquatic vegetation;
Sediment oxygen demand ks(02 m-2 day-1) was chlorophyll)
measured by an in situ method and was determined Stocks of striped bass Morone saxatilis have
to be dependent on oxygen concentration c (ml-1) declined in the Chesapeake Bay system over the last
in the water phase. This dependence was expressed decade. We present evidence for the working
hypothesis that the decline has resulted, in part,

84
from loss of deep-water habitat for adults, caused by Nahant Bay Beaches. MIT Sea Grant
limiting concentrations of dissolved oxygen that are Program, MITSG 82-15.
related, in turn, to nutrient enrichment and greater f seaweeds}
planktonic production. A related hypothesis is that Since the early 1900's the sandy beaches of
changes in the near-shore habitat for juvenile Nahant Bay, Massachusetts have been chronically
striped bass, involving severe declined in fouled by hundreds of tonnes of an abnormal form
submerged aquatic vegetation due to nutrient-driven of the filamentous brown alga Pilayella littoralis.
planktonic shading, also have contributed to the In warm weather, foul-smelling gaseous
decline of striped bass. Nutrients (nitrogen and decomposition products sporadically cause great
phosphorus) and in chlorophyll a, an indicator of discomfort in the local residential and recreational
phytoplankton biomass, have increased i many populations. At the request of state and federal
areas of the bay and tributaries over the past 20 to officials, a multidisciplinary research team was
30 years. These trends are qualitatively correlated assembled to find out why this alga grows so
with greater deoxygenation of the deep channel in abundantly in the surf zone and shallow sublittoral
the mid and upper bay. During the late 1970s, of the fouled beaches. The research team was
summer oxygen concentrations as low as 2 ml/liter specifically asked to evaluate the influence of
approached to within 7-8 m of the surface, allowing effluents from a municipal sewage treatment plant
water stressful to striped bass to intrude onto shoal outfall located on the bay's ocean boundary. The
areas of the bay. The volume of Chesapeake Bay concentrations of algal nutrients were measured in
bottom waters containing 0.5 mg 02/liter or less the outfall boil and at fourteen other locations
was about 15 times greater in July 1980 than in throughout the bay at seventeen monthly intervals.
July 1950. The combination of the expanding In addition, dyes, drift-cards, and computer
hypoxic pool and summer temperature above simulations were used to estimate the trajectory of
preferred levels for adult striped bass may contribute effluents released from the treatment outfall. All
to and "oxygen-temperature squeeze" that forces the available information consistently indicated that
adults onto shoal areas of the bay or out of the the effluent plume is completly dispersed long
upper bay. Many of these shoal areas now lack before reaching the algal drift community, and at
suitable cover for juvenile striped bass and their most, only 1-2% of the macronutrients available to
prey. Strong intraspecific competition among the alga come from the outfall. Tidal exchange
striped bass may be occurring there. with Massachusetts Bay was found to contribute
roughly 74 times as much nitrogen, the
Provini, A., G. Crosa, and R. Marchetti (1992) macronutrient apparently limiting the alga's
Nutrient export from the Po and Adige growth, as the outfall does on each tide. As a
river basins over the last 20 years. In: consequence, the municipal sewage effluents
R. A. Vollenweider, R. Marchetti and R. Viviani released into Nahant Bay were exonerated as a cause
(ed.) Marine Coastal Eutrophication. Elsevier, of the algal fouling problem there.
Amsterdam.
(phosphorus, nitrogen, Adriatic Sea) Quinn, H., J. P. Tolson, C. J. Klein, S. P.
Available data on nutrient concentration and flow Orlando, and C . Alexander (1989)
of the Po and Adige river waters were analyzed in Susceptibility and Status of Gulf of
order to evaluate the nitrogen and phosphorus load Mexico Estuaries to Nutrient Discharges.
carried into the Adriatic Sea, as well as the changes NOAA National Ocean Service, (nutrients;
that have occurred since 1968. Load data results nitrogen; phosphorus; estuary; susceptibility; N/P
were highly correlated with the flow, following ratio; classification; review}
relations of the linear type. The yearly mean loads No abstract
amount to 13,000 t of total P and 110,000 t of
total mineral N for the Po, and 1200 t and 13,000 t Quinn, H., J. P. Tolson, C. J. Klein, S. P.
respectively for the Adige. These values are more Orlando, and C. Alexander (1989)
than two times higher than those evaluated in Susceptibility of East Coast Estuaries to
1968. Nitrogen increased quite regularly, while the Nutrient Discharges: Albemarie/Pamlico
phosphorus load had already reached its maximum Sound to Biscayne Bay. NOAA National
value at the end of the 1970s. Nutrient increase Ocean Service, [nutrients; nitrogen; phosphorus;
during 1968-88 can be explained by a higher estuary; susceptibility; N/P ratio; classification;
production of detergents and fertilizers, and the review}
constructions of numerous sewer networks in the No abstract
Po and Adige basins. Quinn, H., J. P. Tolson, C. J. Klein, S. P.
Quinlan, A. V. (1982) An Ecodynamic Orlando, and C. Alexander (1989)
Analysis of Algal Blooms Fouling Susceptibility of East Coast Estuaries to
Nutrient Discharges: Passamaquoddy Bay

85
to Chesapeake Bay. NOAA National Ocean Radziejewska, T. and I. Drzycimski (1988)
Service, Inutrients; nitrogen; phosphorus; estuary; Meiobenthic communities of the Szczecin
susceptibility; N/P ratio; classification; review} Lagoon. Kieler Meeresforschungen, Sonderheft
No abstract 6:162-172.
[composition; distribution; meiobenthos; lagoon;
Quinn, H., J. P. Tolson, C. J. Klein, S. P. Baltic; nematode; ostracod; copepods)
Orlando, and C . Alexander (1991) Based on data collected at 7 stations 4 times a year
Susceptibility of West Coast Estuaries to in 1985 and 1986, composition and distribution of
Nutrient Discharges: San Diego Bay to meiobenthic communities in the Polish part of the
Puget Sound. NOAA National Ocean Service, Szczecin Lagoon, a eutrophic and polluted water
Inutrients; nitrogen; phosphorus; estuary; body connected with the Baltic Sea, is presented.
susceptibility; N/P ratio; classification; review} The data show a tendency to reduced total
No abstract meiobenthic densities and diversity from the lower
to upper reaches of the lagoon. The meiobenthic
Rabalais, N. N. and D. F. Boesch (1986-1987) communities studied were dominated by nematodes,
Extensive depletion of oxygen in bottom ostracods ranking second in numerical importance.
waters of the Louisiana shelf during Harpacticoid copepods were most abundant at the
1985. Coastal Ocean Pollution Assessment News outer stations which are influenced by Baltic
3:45-47. inflows. Most of the 10 harpacticoid species
(oxygen; Louisiana; Gulf of Mexico) recorded in the lagoon were found at the outer
No abstract stations (lower reached) as well. Similarity analysis
allowed to separate three zones within the lagoon:
Rabalais, N. N., R. E. Turner, J. W.J. Wiseman, (1) the outer zone, its stations showing most
and D. F. Boesch (1991) A brief summary of abundant and diverse meiobenthic communities; (2)
hypoxia on the northern Gulf of Mexico the innermost zone (upper reaches) with the least
continental shelf: 1985-1988. In: R. V. abundant and qualitatively impoverished
Tyson and T. H. Pearson (ed.) Modern and Ancient communities; and (3) the intermediate zone.
Continental SheVAnoxia. The Geological Society,
London, England. Raffaelli, D., J. Limia, S. Hull, and S. Pont
[oxygen; hypoxia; Gulf of Mexico; Louisiana; (1991) Interactions between the amphipod
phytoplankton) Corophium volutator and macroalgal
Oxygen-deficient conditions occur from April to mats on estuarine mudflats. Journal of the
October on the inner to middle continental shelf of Marine Biological Association of the United
the northern Gulf of Mexico and may cover up to Kingdom 71:899-908.
9500 km2 during mid-summer off the Louisiana (macroalgae; seaweeds; mudflat; amphipod;
coast. Hypoxic bottom waters are found in 5-60 in Corophium; Capitella)
water depth, 5-60 km offshore and extend up to 20 One of the most obvious effects of eutrophication
in above the bottom. Salient forcing functions in sheltered coastal areas and estuaries is enhanced
contributing to hypoxic water formation, growth of opportunistic macroalgae, which may
maintenance and break-up are presented. form extensive mats over intertidal mudflats during
Stratification is directly correlated with hypoxia in the spring and summer. In the Ythan estuary,
time and space, suggesting that reaeration of densities of the amphipod Corophium volutator
bottom waters is controlled by physical processes (Pallas) in the sediment underlying weed mats were
that are influenced by regional wind fields, river significantly lower than those in weed-free
discharge and continental shelf scale currents. sediments, and are dominated by species
Phytoplankton biomass reaches the bottom waters characteristic of organically enriched, low oxygen
in the hypoxic zones in large amounts (>5 ug/1), environments such as Capitella capitata. Long-term
fueling water column and benthic respiration rates, data sets on Corophuim abundance in the Ythan
but to a currently unknown degree. Increased suggest that this species has declined dramatically
nutrient loadings in the two major rivers and throughout those parts of the estuary affected by
changes in the proportion of those nutrient weed mats.
essential to phytoplankton growth have probably Two fields experiments have been carried out to
changed both the phytoplankton community assess the effects of weed mats on assemblages. In
species composition and community production. It the first experiment, sediments subjected to a range
is not yet clear whether the extent and severity of of weed densities displayed faunal responses
hypoxia on the Louisiana continental shelf have consistent with those observed by survey of weed-
changed as a result of these riverine water quality covered and weed-free areas. Under a high biomass
changes. of weed, the amphipod Corophuim volutator
disappeared almost completely from the mudflat.
The second experiment indicated that the physical

86
presence of weed filaments may be as important as concentration was due to a higher production rate of
the effects of weed on sediment chemistry in GXT 61 showing that the toxin profile is not a
affecting the density of Corophium, probably conservative property in this species.
through interference with the amphipod's normal
feeding behavior. It is concluded that weed mats Renk, H., J. Nakonieczny, and S. Ochocki (1988)
have a significant impact on Corophium, which is Primary production in the southern Baltic
an important component of the diet of fish and in 1985 and 1986 compared with long-
shore-birds in the estuary. term mean seasonal variation. Kieler
Meeresforschungen, Sonderheft 6:203-209.
Raffaelli, D. G. and C. F. Mason (1981) [primary production; Baltic; phytoplankton;
Pollution monitoring with meiofauna, Chlorophyll a)
using the ratio of nematodes to copepods. On the basis of long-term investigations, the
Marine Pollution Bulletin 12:158-163. course of seasonal variations of mean daily primary
Imeiofauna; nematode; copepod; ratio; sandy production and mean chlorophyll-a concentration in
beaches) the Southern Baltic waters are presented. The mean
The ratio of nematodes to copepods was calculated annual primary production of the Gda'nsk Deep,
for meiofauna samples from 17 beaches and Borholm Deep and the Gotland Deep amounted to
additional values for intertidal and sublittoral sites 12.4 gC m-2, 88.8 gC M-2 and 107.2 gC m-2,
were obtained from the literature. Intertidal ratios respectively. The primary production of the
increased with decreasing particle size, but ratios Southern Baltic in 1986 was higher than in 1985,
from polluted sites were always extremely high. and higher than the long-term mean value. The
Sublittoral ratios increased with depth. It is mean annual production for the Southern Baltic in
suggested that this ratio has potential for 1986 amounted to 130 gC m-2. On the basis of
monitoring organic pollution of sandy beaches. long-term observations of primary production and
chlorophyll-a of the Southern Baltic, certain upward
Raman, A.- V. and K. P. Prakash (1989) trends in phytoplankton production were observed.
Phytoplankton in relation to pollution in
Visakhapatnam harbour, east coast of Revelante,N. and M. Gilmartin (1976) The
India. Indian Journal of Marine Sciences 18:33- effect of Po River discharge on
36. phytoplankton dynamics in the Northern
lphytoplankton; nutrients; Visakhapatnam harbour; Adriatic Sea. Marine Biology 34:259-27 1.
sewage; abundance; diversity) (phytoplankton; Adriatic Sea; community;
Sporadic outbursts of phytoplankton, notably nutrients; density; primary production;
diatoms and phytoflagellates, are observed from nanoplankton; microplankton; blooml
Visakhapatnam harbour waters subjected to An oceanographic transect, extending from
pollution. The harbour waters are characterized by Yugoslavia across the Northern Adriatic Sea to the
high proportions of inorganic nutrients attributable Po delta in Italy, was occupied during 1972 and
to sewage and industrial waste discharges. High 1973 to establish the effect of Po river discharge on
phytoplankton standing crop and chlorophyll the phytoplankton communities of the region.
concentrations accompanied by low species Density distribution showed distinct seasonal
diversity at the harbour locations indicate severe features: a winter-spring period of low stability
eutrophication of waters in this area as against near throughout the water column, and a summer period
normal conditions in the open sea. of stratification. The total water-column plant
nutrients (nitrate, nitrite, phosphate, silicate)
Reguera, B. and Y. Oshima (1990) Response showed a 1.3 to 4.5-fold decrease eastward, with
of Gymnodinium catenatum to increasing semi-eutrophic conditions restricted to an area off
levels of nitrate: Growth patterns and the Po delta. Nanoplankton usually dominated the
toxicity. In: E. Gran6li, B. Sundstr6m, L ' phytoplankton community, in terms of cell
Edler and D. M. Anderson (ed.) Toxic Marine density, surface chlorophyll a concentrations, and
Phytoplankton. Elsevier, New York. surface primary production rates. However, all
(Gymnodinium; nitrate; phytoplankton; growth; significant maxima in these characteristics resulted
red tides} from increases in the microplankton component.
Gymnodinium catenatum, responsible for PSP Changes in the frequency of major microplankton
outbreaks in NW Spain, was cultured in K media groups characterized three periods of the annual
with increasing levels of nitrate. Growth rates were cycle: September-Decernber, neritic, temperate
calculated from measurements of in vivo diatom flora with some littoral elements (e.g.
fluorescence and cell counts. Toxicity was Nitschia seriata); Jani@ary-May, neritic, temperate
determined by HPLC. Toxin concentrations per cell diatom flora of different composition (e.g. Lauderia
reached a peak with nitrate concentrations of 110- borealis, Skeletonema costatum); May-August,
220 uM. Much of the increase in toxin dinoflagellates (e.g. Prorocentrum micans) at

87
western stations and dinoflagellates plus neritic, Revelante, N. and M. Gilmartin (1980)
warm-water diatoms at eastern stations. The Microplankton diversity indices as
seasonal cycle was characterized by spring and fall indicators of eutrophication in the
maxima tending to coincide with maximum Po Northern Adriatic Sea. Hydrobiologia
river discharge and/or periods of low water-column 70:277-286.
stability and vertical mixing. The higher nutrient Jphytoplankton; diversity; Adriatic Sea;
input at western stations was correlated with the co- Mediterranean; indicator; chi. al
dominance of only a few species of microplankton Phytoplankton community diversity indices are
during bloom periods, suggesting that these species used to characterize effects of eutrophication in the
(S. costatum, N. seriata, and 5 others) can serve as Northern Adriatic Sea. A derived Shannon diversity
indicators of eutrophic conditions in this region. frequency spectrum provided a single biological
Assimilation ratios of both the micro- and quantification which allowed an interpretation of
nanoplankton suggested borderline nutrient temporal and regional differences and which can
conditions. Phosphate was implicated as the also be used to evaluate future changes in species
limiting nutrient. diversity. The data base comprised a 4+ year time
series involving 300 taxa.
Revelante, N. and M. Gilmartin (1977) The
effects of Northern Italian rivers and Reys, E. and M. Merino (1991) Diel dissolved
Eastern Mediterranean ingressions on the oxygen dynamics and eutrophication in a
phytoplankton of the Adriatic Sea. shallow, well-mixed tropical lagoon.
Hydrobiologia 56:229-240. Estuaries 14:372-381.
lphytoplankton; Mediterranean; Adriatic Sea; [oxygen; dynamics; eutrophication; lagoon;
standing crop; diversity; chl. a; density; Mexico; primary productivity}
microplankton; nanoplankton; Nitzschia seriata; Bojorquez Lagoon (BL), located on the Mexican
indicator; dinoflagellate; coccolithophoresl Caribbean, has received sewage and dredging
An analysis of the distribution of the impacts as a result of tourism development. The
phytoplankton standing crop during a period of lagoon supports a high diversity of primary
high stability characterized by. the relative influence producers compared to sheltered adjacent lagoons
of northern Italian rivers and Mediterranean waters dominated by Thalassia testudinum communities.
on the Adriatic Sea. The region influenced by The Diurnal Curve Method (Odum and Hoskin,
northern Italian rivers was marked by low 1958) was used to measure community metabolism
community diversities, relatively high chlorophyll and asses eutrophication in BL by comparing it to
a standing crops and cell densities, high the nonimpacted lagoons and to other systems
microplankton to nanoplankton community studied with this method. Dissolved oxygen
volume ratios, and the dominance of Nitzschia community input to the water column in BL ranged
seriata. To the contrary, the region influenced by between 8.3 9 02 m-2 d- I and 41.5 9 02 m-2 d- I
Mediterranean waters exhibited high community during 1985 and 1986, and averaged 17.1, whereas
diversities with prominent oceanic and warm water dissolved oxygen community consumption ranged
elements, low chlorophyll a standing crops and from 6.4 9 02 m-2 d- I to 37.6 g 02 m-2 d- I and
microplankton cell densities, low microplankton to
nanoplankton ratios, and a relatively conspicuous averaged 15.2. These values are higher than those
coccolithophore flora. found for the adjacent lagoons and similar coastal
Nanoplankton populations remained relatively lagoons, and are similar to results from other
constant throughout the Adriatic. However lagoons with sewage or seafood waste discharge.
eutrophication resulted in a marked increase in the Net*flux of oxygen from the community to the
microplankton component of the community, with water column averaged 1-9 9 02 m-2 d- I and ranged
the diatom Nitzschia seriata serving as a prime from -9.8 g 02 m-2 d- I to 8.1 g 0,2 m-2 d-l.
indicator of the process, even in regions several These values are low compared to the adjacent
hundred kilometers downstream from nutrient lagoons, and close to zero, as in dystrophic
sources. environments. Primary productivity, as estimated
Even under relatively low rates of river discharge, by oxygen input, increased in BL during the period
the influence of northern Italian rivers could be of study, indicating that eutrophication is
traced along the entire western side of the sea to the proceeding, but the lagoon has not reached yet a
Otranto Strait, with low north to south and high level of "critical eutrophication" as defined by Mee
west to east gradients resulting from the effect of a (1988).
cyclonic surface circulation pattern on inflowing
Mediterranean waters in the south and river Riegman, R., A. Rowe, A. A. M. Noordeloos, and
discharge in the north. G. C. Cad6e (1993) Evidence for
eutrophication induced Phaeocystis sp.
blooms in the Marsdiep area

88
(Netherlands). In: T. J. Smayda and Y. Ritz, D. A., M. E. Lewis, and M. Shen (1989)
Shimizu (ed.) Toxic Phytoplankton Blooms in the Response to organic enrichment of
Sea. Elsevier, Amsterdam. infaunal macrobenthic communities under
JPhaeocystis; ammonium/nitrate ratio) salmonid seacages. Marine Biology 103:211 -
Recently, a global epidemic of novel and nuisance 214.
phytoplankton blooms has been suggested. Imacrobenthos; benthic; infauna; organic
Anthropogenic linkage is still questionable because enrichment; abundance; biomass; diversity)
the involved mechanisms are unclear. In all coastal The response of infaunal macrobenthic
areas of the European continent, anthropogenic communities beneath salmonid seacages to solid
enrichment of N and P (but not Si) has favoured organic wastes (food pellets and fish exdreta) was
non-diatom blooms during the nutrient controlled tested by means of the ABC-method (abundance,
period (late spring-autumn). The colonial biomass comparison methods: Warwick 1986) on
flagellate, Phaeocystis sp., regarded harmfull samples collected from the SAFCOL seafarm at
because of its massive foam production during the Badger Cove, southeast Tasmania, from February
wane of the bloom, has increased in Dutch coastal to July 1988. The intermittent nature of the
waters. This increase coincides with a decrease of addition of organic waste allowed us to monitor
the average N/P ratio from 38 down to 13 dutring decline and recovery of the macrofauna, i.e.,
the nutrient controlled period (summer) as a harvesting of a cage permitted a period of recovery
consequence of altered nutrient discharges from whilst restocking precipitated a decline. The ABC-
Lake Ussel and the river Rhine. Competition method proved to be a sensitive indicator of
experiments in continuous cultures showed that the community health. Under a normal feeding regime
Phaeocystis could become dominant at N/P ratios the macrofaunal community structure indicated a
below 16. At higher N/P ratios Phaeocystis did moderately disturbed condition. Only 7 wk after the
not become dominant. In laboratory cultures, cage was harvested, species richness had increased
colony formation was absent under P-limitation. markedly and the community adopted an
Under N-limitation, Phaeocystis formed only undisturbed condition. Further improvement was
colonies when nitrate was the nitrogen source. apparent 14 wk post-harvest. Similarly, a decline to
Dutring growth on ammonium, no colonies were a moderately disturbed condition was apparent 7 wk
formed under limiting conditions. The importance after restocking and species richness had declined.
of the ammonium/nitrate ratio for Phaeocystis No such changes occurred under a cage which
colony formation under nutrient limiting conditions contained fish continuously over the same period.
was illustrated in 1991 by a summer bloom which
occurred in the Marsdiep area after the Rosenberg, R. (1985) Eutrophication-the
ammonium/nitrate ratio had dropped remarkably. future marine coastal nuisance? Marine
Pollution Bulletin 16:227-231.
Riisg&rd, H. U. and E. Poulsen (1981) Growth (oxygen; benthic fauna; fish; Sweden; bloom;
of Mytilus eudulis in net bags transferred review)
to different localities in a eutrophicated Increased inputs of nutrients to marine coastal
Danish Fjord. Marine Pollution Bulletin areas over the last decades have created a basis for
12:272-276. eutrophication of the waters surrounding Sweden.
1growth; bivalves; Mytilus; fjord; H2S; oxygen) In combination with relatively low water exchange
The growth rate has been measured in Mytilus in these vertically stratified and almost non-tidal
edulis transferred in net bags to seven localities in waters, local and regional effects of increased
the brackish Danish fjord, Limfjorden, in which macro-algal biomass, and decreased oxygen
certain areas are heavily eutrophicated. The increase concentrations in bottom water leading to
in shell length, shell weight and flesh body weight mortalities of benthic animals and decreased fish
was measured after growth periods of 14-18 days. catches have at times been observed. The effects
The increase in, e.g., flesh dry weight ranged from were first noted in the Baltic, but are now obvious
twice the start weight to a fout-fold increase of the also in Swedish and Danish coastal areas in the
start weight. The net growth efficiencies were Kattegat and the Belt Sea. Similar symptoms have
estimated to be between 54 and 73%. Algal recently also been recorded off the Danish North
concentration was in no case the limiting growth Sea coast. Other shallow coastal and shelf areas,
factor, but low growth rates were observed in areas where stratification occurs, can be regarded as
with seasonal oxygen depletion and release of toxic potentially eutrophic risk areas.
H2S from the sediments. It is suggested that
measurements of actual growth in Mytilus edulis Rosenberg, R. (1992) Eutrophication-related
can be a useful technique in the study of biological marine ecosystem studies in western
effects in marine recipients. Sweden. . In: G. Colombo, I. Ferrari, V. U.
Ceccherelli and R. Rossi (ed.) Marine

89
Eutrophication and Population Dynamics. Olsen & eastern Skagerrak enriched? Journal of
Olsen, Fredensborg. Experimental Marine Biology and Ecology
Jecosystem; sediment transport; Kattegat} 105:219-251.
A six year multidisciplinary programme in the lbenthic; fauna; abundance; biomass; composition;
Kattegat on the Swedish west coast is summarized. organic enrichmentl
There are indications that primary production and In summer 1985, 23 of the stations established in
sedimentation of organic material have increased 1914 by C.G.J. Petersen in the northern Skagerrak
lately due to eutrophication. Although primary and the Oslofjord, were revisited to compare the
production is not exceptional (annual mean 144 g present state of the macrobenthic assemblages with
C per square meter) the special geomorphological the. previous condition. The methods used were
and hydrographical featurew of the southern similar to those of the original survey.
Kattegat (shallow, semi-enclosed, no tide, strong The total benthic biomass was significantly
halocline) promote development of periodic higher in 1985 by, on average, a factor of 1.8 over
bottom-oxygen deficiency. The ecological the whole area. Most of this change was due to a
importance of vertical sedimentation and of lateral highly significant increase of worms, notably
transport of ephemeral blankets into areas of polychaetes, and a significant increase of ophiuroid
deposition is discussecl. Several examples of and echinoid echinoderms. Similarity values in
rapidly occurring negative ecological effects caused terms of biomass suggested that community
by eutrophication are given. composition had changed considerably over the 70-
yr period, in particular in the inner and central
Rosenberg, R., R. Elmgren, S. Fleischer, P. Oslofjor (11% similarity) but also in the outer
Jonsson, G. Persson, and H. Dahlin (1990) Oslofjord and the Skagerrak (33% similarity). There
Marine eutrophication case studies in was no difference in the comparative size of
Sweden. Ambio 19:102-108. individual organisms over the period.
(review; Sweden; nutrient; effects} The observed increase in biomass is greater than
This article summarizes the eutrophication of two the increase along the Swedish Skagerrak coast
sea areas: 1) the Baltic Sea and in particular a observed in the late 1970s, and well beyond the
coastal area which is affected primarily by effluents observed limits of annual variability in such areas.
from a tertiary sewage plant; and 2) the Kattegat Different causes for the observed changes are
and in particular the Laholm Bay, which is affected discussed and it is suggested that a general organic
primarily by nutrient inputs from agriculture and enrichment has taken place in the Oslofjord.
forest land. During this decade, inputs of N and P
to these large sea areas have increased by factors of Rosenberg, R., B. Heilman, and B. Johansson
approximately 4 to 6 and >8, respectively. Internal (1991) Hypoxic tolerance of marine
nutrient transport and atmospheric deposition is benthic fauna. Marine Ecology Progress Series
discussed. It is concluded that N availability 79:127-131.
generally limits primary production, except in [hypoxia; oxygen; benthic; fauna; tolerance}
Bothnian Bay, where P has this role. P is, In stratified coastal marine waters hypoxia is a
however, the limiting factor at times in some growing problem affecting bottom-dwelling
coastal areas, and in the Baltic for N-fixing blue- animals. Earlier studies suggest oxygen
green-algae, which bloom in warm summers when concentrations of about 2 ml 1-1 (Rosenberg, 1980)
low N/P ratios prevail. Noted eutrophication effects as the lower tolerance limit for many benthic
are localized elevated primary production, decreased species in coastal areas. We exposed several
depth distribution and species richness of conspicuous infaunal species on the NE Atlantic
macroalgae. Eutrophication is most likely a continental shelf, contained within their sediment
stimulus to high benthic infaunal biomass and fish habitat, to gradually reduced oxygen concentrations.
biomass in well ventilated sea areas. Other areas Tolerance to hypoxia for the 8 species examined
suffer today from almost permanant (the Baltic was in the range of 0.5 to 1.0 ml P 1 (8 to 15%
proper) or seasonal (the Kattegat) hypoxia with saturation), which they could tolerate for several
devastating effects on benthos and demersal fish. days to weeks. The ophiuroid Amphiura filiformis
The N cycle and future possible consequences in left its protected position in the sediment at an
relation to nutrient input are discussed. It is oxygen concentration of 0.85 ml 1-1 (13% sat.),
concluded that a reduction by at least half of whereas A. chiajei emerged from the sediment at
external N and P to the Baltic sea, and N inputs to
the Kattegat is needed to mitigate the negative 0..54 ml 1-1 (8% sat.).
effects of eutrophication. Rosenberg, R. and L.-O. Loo (1988) Marine
Rosenberg, R., J. S. Gray, A. B. Josefson, and T. eutrophication induced oxygen deficiency:
H. Pearson (1987) Peterson's benthic Effects on soft bottom fauna, western
stations revisited. 11. Is the Oslofjord and Sweden. Ophelia 29:213-225.

90
(oxygen; benthic; macrofauna; increase; mortality}
Marine eutrophication is studied in a Ryther, J. H. and W. M. Dunstan (1971)
multidisciplinary project in the Kattegat, western Nitrogen, phosphorus, and eutrophication
Sweden. Input of nutrients has increased several- in the coastal marine environment.
fold during this century; nitrogen = 6 times, Science 171:1008-1013.
phosphorus = 10 times. This has caused elevated [nutrients; nitrogen; phosphorus; phytoplanktonj
concentrations of nutrients in the water and The distribution of inorganic nitrogen and
increased primary production. Some adverse effects phosphorus and bioassay experiments both show
at the bottom have been noted annually from about that nitrogen is the critical limiting factor to algal
August through October in the 1980's, e.g. fish growth and eutrophication in coastal marine waters.
catches have dropped and mortalities have been About twice the amount of phosphate as can be
reported for fish and benthic animals including used by the algae is normally present. This surplus
Nephrops norvegicus. results from the low nitrogen to phosphorus ratio
In this paper we evaluate the effects on benthic in terrigenous contributions, including human
macrofauna which are suggested to be caused waste, and from the fact that phosphorus
mainly by oxygen deficiency occurring especially regenerates more quickly than ammonia from
around the depth of a strong halocline ( = 15 in), decomposing organic matter. Removal of
but also along a bottom transect down to 57 in. At phosphate from detergents is therefore not likely to
some sites the benthic communities were- slow the eutrophication of coastal marine waters,
impoverished, and at others only some species and its replacement with nitrogen-containing
seemed to be especially sensitive, e.g. the bivalve nitrilotriacetic acid may worsen the situations.
Abra alba and the brittle star Amphiura filiformis.
In shallow ( < 10m ) exposed waters an annually Saad, M. A. H. and E. 1. M. Hemeda (1992)
occurring mortality of molluscs was observed over Effect of pollution on the Western
large areas from 1980 onwards. Affected species Harbour of Alexandria. 1. Environmental
were mainly the bivalves Cardium edule and Mya characteristics. In: R. A. Vollenweider, R.
arenaria. The causes behind this mortality may also Marchetti and R. Viviani (ed.) Marine Coastal
be oxygen deficiency occurring during calm weather Eutrophication. Elsevier, Amsterdam.
periods. JpH)
The Western Harbour of Alexandria has an area of
Rydberg, L., L. Edler, S. Floderus, and W. Grandli about 1962 acres and a water depth ranging from 6-
(1990) Interaction between supply of 14 in. This harbour is protected by two water
nutrients, primary production, breaks and is divided into two areas; the inner and
sedimentation and oxygen consumption outer ports. The water quality of the harbour has
in SE Kattegat. Ambio 19:134-141. deterioated due to the effects of various pollutants
Inutrients; nitrogen; phosphorus; oxygen; Baltic) discharged from ships at anchor and sewage waste
Intensive measurements of nutrient fluxes, carbon outfalls. Variations of some of the environmental
and nitrogen assimilation, sedimentation and conditions in the harbour were studied.
oxygen consumption within the SE Kattegat are Temperature measurements generally decreased with
used in an effort to follow the coupling between depth. The highest average water temperatures were
increasing nutrient supply and decreasing deep-water recorded in July and August, whereas the lowest
oxygen concentration. The investigated area is a were in January and February. The Secchi disc
part of the strongly stratified Baltic estuary, with a readins showed remarkable variations: the
hampered deep-water exchange and a large supply of minimum and maximum regional average Secchi
nitrogen from nearby anthropogenic sources. values were accompanied by the highest and lowest
Measurements were undertaken during 1981-1988. regional averages of suspended matter (SM). The
The results are discussed, mainly in terms of annual irregularity in the vertical distribution of SM
or seasonal mean values. The average primary possibly coincided with turbulence of the wsater
production, 11.4 mol C m-2 (corresponding to 45 column from navigation and the effect of pollution.
mmol C m-2 d-1 during the "productive season" Salinity values generally increased with depth. The
from March to October), was surprisingly weli discharge of sewage wastes from numerous ships
correlated with the uptake of inorganic nitrogen and land-based sources decreased the surface salinity.
which averaged 1.68 mol N m-2 (11.1 mol C m-2: The highest regional average salinity values were
using the Redfield ratio). The oxygen consumption found at locations far from the direct effect of
sewage pollution. The pH values generally
(deep water and benthic) which averaged 20 mol 02 decreased with depth: low pH values were found at
m_2 d-1, was well correlated with the measured locations directly affected by sewage wastes.
nitrate uptake and with the external supply of
nitrate to the surface water. The consumption was Sales, D., A. G6mez, and D. Cantero (1983)
twice as high as that in the open Kattegat. Incidence of urban sewage disposal in the

91
salt-ponds areas of the south of the Bay control and were correlated with temperature and
of Cadiz. Marine Pollution Bulletin 14:447- carbon concentrations (r2=0.85). Anaerobic
452. metabolism was the dominant metabolic pathway
(Cadiz Bay; oxygen; salt pond) in control and treated sediments, with 50 to 70% of
The highest pollution indices in the marsh and annual carbon remineralization due to sulfate
tide-lands situated to the south of the Bay of Cadiz reduction.
have been evaluated. At the present, the salt
industry of this area is being transformed into Sampou, P. A. (1989) Effects of
mariculture installations. This study consisted of eutrophication on the biogeochemical
the systematic evaluation of different parameters at cycling of carbon, oxygen, sulfur and
26 sampling stations during the period 1976-1981. energy in coastal marine ecosystems.
The results suggest that extensive zones are too Dissertation for Ph.D., University of Rhode Island.
seriously affected to make use of them as marine Jbiogeochernical cycling; energy flow; MERL;
piscifactories. sewage; nutrients; production; ecosystem; oxygen)
The effects of eutrophication on the
Samanidou, V., K. Fytianos, and G. Vasilikiotis biogeochernical cycling of the major elemental
(1989) Distribution of nutrients in the cycles, carbon, oxygen, and sulfur and on energy
Thermaikos Gulf, Greece. Toxicological and flow, were investigated in experimental marine
Environmental Chemistry 20-21:29-37. ecosystems (mesocosms). Eutrophic environmental
(nutrients; Greece; methods; nitrogen; phosphorus; conditions were created within mesocosms by daily
distribution; sewage) addition of varying amounts and sources of
The distribution of nutrients (phosphate, nitrate, nutrients and sewage sludge. Inorganic nutrient
nitrite and ammonium) in water samples from loading resulted in an increase in the magnitude and
eleven sampling stations in the Thermaikos Gulf duration of winter-spring phytoplankton
was studied during a period of one year (1985- populations and production with a correlated
1986). Significant seasonal fluctuations were increase in sedimentation and accumulation of
observed mainly for nitrate and ammonium, with a carbon in surface sediments. Net ecosystem
maximum during winter and a minimum in the production during the winter and spring was
summer. The obtained data showed that the followed by net system heterotrophy during the
nutrients are present at relatively high summer and fall with a concomitant decrease in
concentrations near the agricultural areas, in the sedimentary carbon concentrations. Carbon
harbour of Thessalonki and in the area where sedimentation and accumulation (up to 19 mol C
municipal sewages are discharged. M-2) in surface sediments stimulated sediment
oxygen consumption and sulfate reduction
Sampou, P. and C. A. Oviatt (1991) Seasonal metabolism. Sulfate reduction increased by an order
patterns of sedimentary carbon and of magnitude in carbon enriched sediments and was
anaerobic respiration along a simulated a dominant pathway for benthic carbon
eutrophication gradient. Marine Ecology remineralization in all the mesocosms. However,
Progress Series 72:271-282. the fraction of benthic metabolism using sulfate for
forganic carbon; oxygen uptake; sulfate reduction) a terminal electron acceptor did not increase along
Concentrations of organic carbon and rates of the nutrient addition gradient when annual
dissimilative sulfate reduction in surface sediments integrated estimates of metabolism were compared.
of marine mesocosms were examined along an Whole system carbon and oxygen budget showed
experimental eutrophication gradient. the sediments to be an important and influential
Phytoplankton biomass increased due to addition of component in eutrophic marine ecosystems. A
inorganic nutrients (N, P, Si). This increase was significant fraction of alloclithonous sewage carbon
especially pronounced during the winter-spring was remineralized via sulfate reduction in the
diatom blooms, which increased in magnitude and sediments. This led to the dissociation of carbon
duration along the nutrient gradient. Net system cycling from oxygen and energy flow in the system
production in winter and spring resulted in carbon due to an increased role of sulfur biogeochemistry.
deposition and accumulation in surface sediments Accumulation of solid sulfides in the sediments
(maximum net accumulation 17 mol C m-2). from sulfate reduction during the experiment
Benthic remineralization of carbon exceeded explained the discrepancy between carbon dioxide
depositional supply during summer and fall. and oxygen consumption at the sediment water
Sediment carbon concentration approached interface (respiratory quotients > 2) and represented
background levels in December and February, a large storage of energy originally contained in
suggesting very little annual accumulation of organic matter.
sediment'carbon. Sediment oxygen consumption Oxygen dynamics in eutrophic marine systems
and sulfate reduction rates in organic enriched were regulated by phytoplankton cycles and sulfur
sediments were an order of magnitude higher than cycling in the sediments. Initiation of hypoxia

92
(oxygen less than 2 mg 1-1) was caused by rapidly indicator of trace-metal pollution as it occurs over a
declining phytoplankton populations and a large widespread area and chemical analysis for trace
imbalance between day oxygen production and metals is a relatively simple procedure. Due to the
night oxygen consumption in the water column. high growth potential of eelgrass leaves (about 5-6
However, high sulfate reduction rates and the cm leaf turion- I day- 1), growth rates of leaves can
accumulation of hydrogen sulfide in surface be used as a very sensitive method of assessing the
sediments was found to be influential in the demise effects of industrial waste products both in the field
of the macrofauna and the precipitation and and in laboratory tests.
maintenance of oxygen concentrations less than I
mg 1- 1. Sandulli, R. and M. d. N. Giudici (1989) Effects
of organic enrichment on meiofauna: A
Sand-Jensen, K. and J. Borum (1983) laboratory study. Marine Pollution Bulletin
Regulation of growth of eelgrass (Zostera 20:223-227.
marina L.) in Danish coastal waters. Imeiofauna; sewage; nematode; harpacticoid;
Marine Technology Society Journal 17:15-21. copepod; ratio}
Isubmerged vegetation; eelgrass; growth; shading) A 84 day laboratory experiment indicates the
Eelgrass is the dominant macrophyte in coastal decline of intertidal meiofauna abundance may be
Danish waters. The productivity of eelgrass is at directly related to treatment level and time of
the same level as fertile terrestrial crops and it can exposure to sewage sludge. The observed changes
be substantially higher than the phytoplankton revealed a marked decrease of nematodes and total
production in open waters when compared per unit disappearance of mesobenthic harpacticoids while
surface area. Eelgrass may contribute significantly the non-interstitial copepod species were littl 'e
to the total production of all autotrophic affected. The possibility of a new interpretation of
components in shallow coastal areas. the Nematode/Copepod ratio as an index of organic
The above-ground biomass of eelgrass is more pollution in sediments is discussed.
variable than the below-ground biomass due to a
higher turnover rate and exposure to physical Schiewer, U., R. B6rner, and N. Wasmund (1988)
factors of the former. In shallow water, biomass of Deterministic and stochastic influence of
eelgrass increases rapidly from spring to early nutrients on phytoplankton function and
summer and stabilizes at a high level during late structure in coastal waters. Kieler
summer. Growth is continuous throughout the year Meeresforschungen, Sonderheft 6:173-183.
although it occurs at a higher rate during spring and [phytoplankton; nutrients; composition;
summer. Individual plants produce about 16 leaves abundance; chlorophyll al
and rhizome segments per year. During spring and Knowledge of how phytoplankton responds to
summer each plant produces a new leaf every 9-15 nutrient inputs is essential for water management
days and their life span is about 2 months. The and for minimizing eutrophication. Only processes
rapid turnover is important in keeping down the that are deterministic, i.e. that can respond as
epiphytic load by producing new tissue. It also algorithms, are controllable. The study area is the
produces a continuous supply of senescing leaves chain of inshore waters (so-called Bodden) south of
which presumably leads to stabilized energy the Darss-Zingst peninsula- shallow eutrophic
transformations within the detritus food web. Since waters of estuarine character in the Southern Baltic.
the leaf biomass increases from spring to summer, Monitoring programmes and laboratory
the ratio of daily productivity to biomass (P/B) experiments have revealed an annual periodicity of
falls because of increased self-shading. Water the phytoplankton and of the physico-chemical
temperature and availability of nutrients do not factors influencing it. On the basis of these results,
affect seasonal variation in leaf productivity. experiments were carried out in enclosures to study
Nutrient availability may, however, affect the the effects of nutrient loading on phytoplankton.
biomass as a yield limiting factor. The purpose was to test the feasibility of
Eutrophication of coastal waters changes in the influencing phytoplankton development under field
balance among different autotrophic components. conditions during the transition period from late
Growth of phytoplankton, epiphytes and free- spring to mid-summer.
floating algae, which take up inorganic nutrient This contribution presents results from the 1985
from the free water phase, is greatly stimulated. shallow water enclosure experiments (FLAK 85)
This will result in decreased growth rates, depth which demonstrate that
penetration and, eventually, survival of eelgrass - the scale of phytoplankton reactions and the
though shading. species involved are stochastic in character and are
Biomass and growth measurements of epiphytic governed by stochastic interactions between
algae provide a convenient method for assessing the meteorological events and water exchange processes
effect of inorganic nutrient loading in different parts in the chain of Bodden;
of a receiving area. Eelgrass is suitable as an

93
all processes affecting phytoplankton growth are processes. Kieler Meeresforschungen, Sonderheft
deterministic in character, conforming to simple 6:221-240.
batch theories: simultaneous addition of nitrogen Initrogen; phosphorus; seaweeds; productivity;
and phosphorus favours green algae, and in Fucus; Phycodrys; Baltic)
exceptional cases one algal species became Considerable amounts of nutrients enter, as a
dominant result of human activity, the Western Baltic Sea in
- nutrient loadings do not affect the time of the form of discharge of seepage from land or as
transition to the mid-summer phytoplankton atmospheric input. Benthic macrophytes which
population, the most important regulating factor preferably inhabit the nutrient recipient shallow
obviously being temperature. near-shore areas, and where they often constitute the
major primary producers may thus play an
Schindler, D. W. (1981) Studies of important role in eutrophication processes by
eutrophication in lakes and their uptake and accumulation on nutrient elements,
relevance to the estuarine environment. enhanced primary production and following
In: B. J. Neilson and L. E. Cronin (ed.) Estuaries degradation of the produced organic material.
and Nutrients. Humana Press, Clifton, New Jersey. Nutrient uptake, binding capacity and illumination
flake; estuary; comparison; Vollenweider; nutrient; as well as nutrient dependent growth of the two
composition) community forming seaweeds Phycodrys rubens
No abstract and Fucus vesiculosus have been investigated and
related to the seasonal patterns of the nutrient
Schlungbaum, G. and G. Nausch (1988) conditions in their respective habitats. Nutrient
Nutrient turnover at the sediment/water concentrations, both phosphorus and nitrogen, in
interface in shallow eutrophic coastal the algal beds are markedly higher than in the
waters. Kieler Meeresforschungen, Sonderheft surface water of the open Kiel Bight. In general, the
6:184-190. seaweeds seem not to be nutrient limited under
(sediment; interface; nutrient; phosphorus; natural conditions except for nitrogen in Phycodrys
phosphate; phytoplankton; primary productionj during summer. Nutrient tissue contents are
The nutrient situation in eutrophic shallow saturated only for nitrogen during winter. The
coastal water systems is characterized by particular significance of these findings is discussed in view
features. In addition to water exchange with the of the observed increasing nutrient levels and
open sea, interactions at the sediment/water changes of the vegetation in the Kiel Bight.
interface play a significant role. With the help of
investigation of phosphate sorption/desorption Schultz, S., G. AErtebjerg, G. Behrends, G. Breuel,
balances and on the denitrification capacity of P. Ciszewski, U. Horstmann, K. Kononen, E.
shallow coastal waters, the following general Kostrichkina, J. Leppdnen, F. Mohlenberg, 0.
conclusions can be made: The phosphate Sandstr6m, M. Viitasalo, and T. Will6n (1992)
concentrations in the water are primarily determined The present state of the Baltic Sea
by physico-chemical reactions with the sediment, pelagic ecosystem - an assessment. In:
and less by the rhythm of the phytoplankton G. Colombo, I. Ferrari, V. U. Ceccherelli and R.
primary production. Through intensive interactions Rossi (ed.) Marine Eutrophication and Population
between sediment and water, a phosphate Dynamics. Olsen & Olsen, Fredensborg.
11 equilibrium" concentration which fluctuates very (Baltic Sea; assessment; pelagic biology;
little is maintained over the whole year. High phytoplankton assemblage)
primary production rates are possible at these low In recent years, the decrease of salinity as well as
equilibrium concentrations of phosphate sorption of the temperature and oxygen in the deep water has
capacities. With the development of nearly continued in the Baltic Sea. In addition, the
anaerobic conditions, nitrate reduction can occur at content of phosphate and nitrate lids declined in the
a rate, when the appropriate amount of nitrate is winter surface water at least during the last two
made available to the reaction. Under optimal years. However, both the primary production
conditions for nitrate reduction, nitrate capacity and the assimilation numbers have been
concentration is also not a parameter characterizing increasing for most sub-areas of the Baltic Sea,
the nature of the water body. indicating a higher phytoplankton activity. The
stock of zooplankton increased in tha Arkona,
Schramm, W., D. Abele, and G. Breuer (1988) Bornholm and Eastern Gotland Seas. Changes in
Nitrogen and phosphorus nutrition and the species composition or in the structure of the
productivity of two community forming phytoplankton and zooplankton community have
seaweeds (Fucus vesiculosus, Phycodrys not been observed.
rubens) from the Western Baltic (Kiel The present status of the Baltic Sea pelagic
Bight) in the light of eutrophication system reflects clear signs of eutrophication. The
reasons for this are seen in the anthropogenic

impact in conjunction with the features of the Several researches for eutrophication in the
Balitc Sea and a stagnation period lasting thirteen northern Hiroshima Bay were carried out to
years. investigate nutrient budgets in aquatic circulation
processes, i.e. primary production, settling flux of
Schulz, S. and G. Breuel (1981) Response of particulate organic matter (POM) and the benthic
plastic-bag enclosed summer remineralization.
phytoplankton to a nutrient input under It became apparent from these investigations that
in situ conditions in the Arkona Sea primary production contributed more to the organic
(Baltic Sea). Kieler Meeresforschungen, pollution in the northern bay than land based
Sonderheft 5:9 1 - 100. organic loadings by a factor of 10, that approx. 70-
(phytoplankton; nutrients; Baltic; primary 80% of the POM originating in phytoplankton
production; Arkona; biomass; composition) were easily remineralized into inorganic nutrients
In July 1878 several experiments with enclosed during the settling process in water, and that
summer phytoplankton communities were carried approx. 60% of nitrogen and 70% of phosphorus in
out in the Arkona Sea using 25 1 cylindrical bags POM settled on the sediments might be returned to
made of 0.2 mm polyethylene film. These the water column by the release from benthic
experiments concerned the response of sediment.
phytoplankton to nutrient input. Phytoplankton
biomass, primary production and chlorophyll Seitzinger, S. P. and S. W. Nixon (1985)
content increased rapidly in enriched water, in Eutrophication and the rate of
contrast to the controls where changes were denitrification and N20 production in
negligible. Nutrients were depleted to the same coastal marine sediments. Limnology and
levels as in the controls after a few days. The g- Oceanography 30:1332-1339.
algae were the first to respond to nutrient input, fdenitrification; N20 production; nutrient; MERLI
followed later by diatoms and dinoflagellates. In the Large (13 m3, 5m deep) microcosms with coupled
course of the experiments, however, the energy was pelagic and benthic components were used to
stored to an increasing extent in the cyanophytes at measure the effect of nutrient loading and
the expense of other groups where energy content eutrophication in coastal marine ecosystems on the
remained constant or decreased slightly. rates of benthic denitrification (ND and N20
Schulz, S., G. Breuel, and A. Irmisch (1988) production. After 3 months of daily nutrient
Simulated eutrophication in enclosure additions, average denitrification rates ranged from
experiments in the Arkona Sea. Kieler about 300 umol N m-2 h- I in the sediments of the
Meeresforschungeb, Sonderheft 6:73-84. control microcosm to 880 in the most enriched
(Arkona Sea; Baltic; plankton; nutrients; primary microcosm, which received 65 times the nutrient
production; biomass; productivity) input of the control. Increases in the production of
In spring, summer and autumn enclosure N20 were more dramatic and increased by a factor
experiments were performed in the central part of of about 100, from 0.56 N m-2 h- I in the control
the Arkona Sea. The. natural water with the to 51 in the most enriched microcosm. Although
plankton community was enriched by nutrients to there was a clear increase in the denitrification rate
about winter levels, and to the double of these in the more eutrophic systems, the amount of fixed
concentrations. nitrogen removed was a constant or progressively
In spring and summer, the phytoplankton smaller fraction of the nitrogen input. Even in the
responded with rapid uptake of nutrients and an most enriched microcosm, at least 16% of the N
increase in primary production and biomass. In input was removed by denitrification.
autumn, the uptake of nutrients was also fast,
whereas productivity did not increase, and biomass Seliger, H. H. (1989) Mechanisms for red
only in diatoms. Not only did the production tides of Pyrodinium bahamense var.
increase with higher nutrient supply, but also the compressum in Papua New Guinea, Sabah
productive season was prolonged. This fact is and Brunei Darussalam. In: G. M.
important for the function of the pelagic system in Hallegraeff and J. L. Maclean (ed.) Biology,
the Baltic, because the biomass and nutrients Epidemology and Management of Pyrodinium red
remaining in the spring bloom determine to a great Tides. Fisheries Department, Ministry of
extent the productivity of the whole year. Development, Manila.
[red tides; blooms; abundance)
Seiki, T., E. Date, and H. Izawa' (1991) Mechanisms are proposed to explain the
Eutrophication in Hiroshima Bay. Marine development of red tides of the toxic dinoflagellate
Pollution Bulletin 23:95-99. Pyrodinium bahamense var. compressum along the
(nutrients; primary production; budgetl northern and southern coasts of Papua New Guinea
and along the northwest coast of Borneo, from

Sabah to Burnei Darussalam. The "spread" of red Skjoldal, H. R. and I. Dundas (ed.) (1991) The
tides to areas not previously affected and the Chrysochromulina polylepis bloom in
increased frequency of annual occurrences of the Skagerrak and the Kattegat in May-
paralytic shellfish poisoning in already colonized June 1988: Environmental conditions,
areas are suggested to be the result of coastal possible causes, and effects. ICES,
hypertrophication. Increased nutrients along Copenhagen.
coastlines permit sufficient reproduction of (bloom; Chrysochromulina; causesl
dinoflagellates in entrained surfaces patches to No abstract
compensate dilution losses, enhancing emigration
over longer distances. Smayda, T. J. (1989) Primary production
and the global epidemic of phytoplankton
Sellner, K. G. and M. M. Olson (1985) blooms in the sea: A linkage? In: E. M.
Copepod grazing in red tide of Cosper, N. M. Bricelj and E. J. Carpenter (ed.)
Chesapeake Bay. In: D. M. Anderson, A. W. Novel Phytoplankton Blooms; Causes and Impacts
White and D. G. Baden (ed.) Toxic Dinoflagellates. of Recurrent Brown Tides and Other Unusual
Elsevier, New York. Blooms. Springer-Verlag, Berlin.
(secondary production; zooplankton; bloom; (primary production; phytoplankton bloom)
Chesapeake Bay; red tide; grazing; dinoflagellates} No abstract
Copepod grazing of dinoflagellate blooms in
Chesapeake Bay and its tributaries was investigated Smayda, T. J. (1992) A phantom of the
using 14C-labelling procedures. Grazing rates for ocean. Nature 358:374-375.
Eurytemora affinis, the late winter - early spring Idinoflagellate; red tides; foodweb)
dominant herbivore, on Heterocapsa triquetra were No abstract
low with a mean rate of 0.011 +- 0.002 ml
copepod- 10. However, at a cell density of 2.43 X Smayda, T. J. and Y. Shimizu (ed.) (1993)
105 ml-1, the ingestion rate corresponds to 1.88 Toxic Phytoplankton Blooms in the Sea
(Proceedings of the Fifth International
ugC copepod-lh-1, or approximately one-third of Conference on Toxic Marine
the copepod's daily ration. Fecal pellet production Phytoplankton, Newport, Rhode Island,
every 6-9 min. Low grazing rates for the spring-fall U.S.A., 28 October -1 November 1991).
dominant herbivore, Acartia tonsa, were also Elsevier, Amsterdam.
recorded where the copepod was feeding on blooms No abstract
of Gymnodinium sp. and Katodinium rotundatum.
Mean rates ranged from 0.005 - 0.048 ml copepod- Smayda, T. J. and A. W. White (1990) Has
10, corresponding to estimated ingestion rates of there been a global expansion of algal
0.038 - 0.102 ugC copepod-lh-1, well below the bloom? If so, is there a connection with
rate recorded for Eurytemora. These data indicated human activities? In: E. Gran6li, B.
that A. tonsa may not remove significant quantities Sundstr6m, L. Edler and D. M. Anderson (ed.)
of dinoflagellate carbon in Chesapeake Bay and its Toxic Marine Phytoplankton. Elsevier, New York.
sub-estuaries. [phytoplankton; algae; . bloom; nutrient
enrichment}
Silva, E. S. (1985) Ecological factors No abstract
related to Prorocentrum minimum blooms
in Obidos Lagoon (Portugal). In: D. M. Smith, D. W. (1984) Responses Of
Anderson, A. W. White and D. G. Baden (ed.) aufwuchs, phytoplankton and
Toxic Dinoflagellates. Elsevier, New York. macrophytes to municipal and industrial
fred tide; bloom; environmental condition} wastes in San Francisco Bay and bay
Prorocentrum minimum Schiller red waters have analog systems. Dissertation for Ph.D.,
often been observed in Obidos Lagoon and have University of California, Berkeley.
caused toxicity of bivalve there. Particular attention tauhwuchs; phytoplankton; seaweeds; San
is given to two of those blooms, separated by Francisco Bay; method; mesocosm)
about 10 years, in May-June 1973 and in January- The use of aquatic organisms which colonize
February 1983. A comparative study of artificial substrates (aufwuchs) is currently being
environmental conditions during the two red waters developed as a method of assessing localized effects
of A minimum is presented. of municipal and industrial discharges into San
Francisco Bay, California by state regulatory
Skjoldal, H. (1992) agencies. The method is intended to provide an
Personal Communication. inexpensive and robust bioassay which will serve
as an index of wastewater effects on the
phytoplankton community, since the effects on

phytoplankton cannot be directly observed. In this nitrogen levels within the water column.
study functional or physiological responses Particulate materials including plankton biomass,
(photosynthesis, respiration, chlorophyll a, and as well as dissolved inorganic phosphorus, were
biomass) were measured in the laboratory, in elevated substantially. The benthos showed
mesocosms and in the field to characterize the increased biomass and metabolic rates, especially of
response of the aufwuchs community to wastewater heterotrophs responding to fallout of organic
and some of its individual stimulatory and toxic particulate materials. Nutrient recycling within the
components. The mesocosm consisted of 12 Bay was the major immediate source for the
outdoor 3000L tanks with a constant supply of nutrients for the observed rapid metabolic activity.
baywater. The response of aufwuchs under Sewage accounted for approximately 80 percent of
experimental conditions confirmed and elucidated the inorganic nitrogen and 90 percent of the
the causes of the stimulatory response of aufwuchs inorganic phosphorus delivery to Kaneohe Bay.
grown on substrates moored in dilution-fields in Diversion lowered the land-derived inorganic
San Francisco Bay. nitrogen and phosphorus input to the Bay by 70-80
The effects of colloidal and microbial components percent. Virtually all components of the system
of wastewater on earlier colonizing and later rapid have responded to this diminished nutrient subsidy,
growth phase of the auchwuchs community were but the water column nutrient washout and
determined to be unimportant with respect to the biological recovery are predictably occurring more
measurable differences caused by exposure to rapidly than the benthos responses.
wastewater. Wastewater was found to stimulate
growth of aufwuchs. Inorganic nitrogen was Snelgrove, P. V. R. and J. B. Lewis (1989)
determined experimentally and deductively to be the Response of a coral -associated crustacean
causative stimulatory component of wastewater. community to eutrophication. Marine
Aufwuchs were also shown to be sensitive to toxic Biology 101:249-257.
compounds found in the wastewater. [crustacea; coral; Madracis; Barbados; composition;
The stimulatory and toxic response of aufwuchs density)
amplified the response of phytoplankton. Various Seventy-five samples of the crustacean fauna
aspects of the Bay's physical and chemical associated with the coral Madracis mirabilis
environment such as light turbulence and ambient (Duchassaing and Michelotii) were obtained at five
nutrient concentration may result in this sites along the west coast of Barbados in 1985.
amplification by moderating factors. Aufwuchs, Samples were analyzed to determine whether any
then, may serve as an early warning of potential community response to eutrophication was evident.
effects of wastewater on phytoplankton. Ordination (multidimensional scaling) and
Strategies of substrate deployment and harvest classification (group average clustering) based on
were also assessed. Increased biomass, facilitated by Bray-Curtis dissimilarity measures separated the
relatively long incubation intervals (7-14 days), fauna of highly eutrophied sites from those that
was shown experimentally to maximize the were less eutrophied. Dominance plots based on
precision of the technique. The precision of the relative abundance and log-normal plotting methods
method is adequate from a regulatory point of view were ineffective in distinguishing the fauna of
to delineate the region where biota are influenced by eutrophied sites. The crustacean fauna responded to
wastewater. eutrophication mainly by reduction of density of
animals. No differences in species richness, number
Smith, S. V. (1981) Responses of Kaneohe of site endemic species or rare species were
Bay, Hawaii, to relaxation of sewage observed between highly eutrophied and less
stress. In: B. J. Neilson and L. E. Eugene (ed.) eutrophied sites. Nine "indicator" species of
Estuaries and Nutrients. Humana Press, Clifton, eutrophy were identified on the basis of strong
New Jersey. density differences between sites.
(nitrogen; sewage; nutrient; cycling; Hawaiij
Kaneohe Bay is a subtropical coral reef/estuary Soulsby, P. G., D. Lowthion, and M. Houston
complex which was subjected to increasing sewage (1982) Effects of macroalgal mats on the
loading; that sewage was diverted in 1977 and ecology of intertidal mudflats. Marine
1978. We have treated the loading and diversion as Pollution Bulletin 13:162-166.
a controlled , total-ecosystem experiment to Imacroalgal mats; mudflat; benthos; abundance;
evaluate the chemical and biological responses to biomass)
external subsidy of nutrients. We here consider the The standing biomass and distribution of the
Bay's response to sewage loading and to its invertebrate fauna of mudflats covered by
diversion largely in the context of a nitrogen macroalgal mats has been determined and compared
budget. with that of mudflats unaffected by algae. The
Even the most heavily impacted portion of the faunal biomass of algal-covered mud was
Bay showed only modest increases in dissolved consistently higher than uncovered mud, mainly

97
due to the presence of large populations of the snail Stachowitsch, M. (1991) Anoxia in the
Hydrobia ulvae. The structure of the faunal Northern Adriatic Sea: rapid death, slow
composition of algal-covered mud was altered and recovery. In: R. V. Tyson and T. H. Pearson
the numbers of at least one species of polychaete (ed.) Modern and Ancient Continental Shelf
worm markedly reduced. The effects of these Anoxia. The Geological Society, London, England.
changes on the bird population exploiting the JAdriatic; oxygen; benthos; anoxia; mortality}
mudflats is discussed. The Northern Adriatic Sea is characterized by the
full range of features associated with sensitive
Soulsby, P. G., D. Lowthion, and H. A. C. marine ecosystems. In the Gulf of Trieste,
Montgomery (1985) The role of sewage mortalities due to anoxia have been reported in
effluent in the accumulation of 1974, 1983 and 1988. The anoxic event in
macroalgal mats on intertidal mudflats in September 1983 led to mass mortality in a highly
two basins in southern England. developed and wide-ranging community in the Gulf.
Netherlands Journal of Sea Research 19:257-263. This destroyed over 50% of the epifaunal biomass
Imacroalgae; nutrients) in only two days, and over 90% within four days.
Two intertidal basins in southern England, The increased frequency, severity, and range of
Portsmouth Harbour and Langstone Harbour, both recent oxygen deficiencies suggest that, in addition
support large crops of the macroalgae Ulva and to meteorological and hydrographic factors,
Enteromorpha each summer. Both Harbours have anthropogenic influences are a contributing or
received discharges of sewage but the discharge in enhancing factor in the onset of anoxic conditions
Portsmouth has been removed. Data from surveys and associated mass mortalities. These
of algal distribution, standing biomass, tissue anthropogenic influences may be subsumed under
nutrient concentration and water column nutrient the heading eutrophication. If eutrophication only
concentration have been evaluated in order to' slightly increases the rate of oxygen decrease below
deternine the relationship between effluent discharge the thermocline, it may advance the point at which
and macroalgal growth. No direct relationship critical oxygen levels are attained by several days.
between water column nutrient concentration and Since only a few days of anoxic conditions are
macroalgae biomass or area cover was found in the required to destroy this benthic community (with an
field. Concentrations of Nitrogen and equally short time required for reoxygenation) the
Nitrogen: Phosphorus ratio in the tissues suggest degree of eutrophication may determine whether an
that Nitrogen is stored in excess during the growing ecological catastrophe occurs or not.
season. It is concluded that the evidence strongly The recolonization process between 1984 and
indicates that further increases in sewage derived 1987 has been slow: the total biomass and the
nutrient will not result in higher standing crops of relative contribution of the designating species have
algae than currently observed. remained distinctly below previously recorded
values. Additional anthropogenic influences
Spencer, C. P. (1985) The use of plant (intensive dredging) may further impede
micro-nutrient and chlorophyll records as recolonization. A further, small-scale mortality was
indices of eutrophication in inshore registered in 1988.
waters. Netherlands Journal of Sea Research
19:269 -275. Stambler, N., N. Popper, Z. Dubinsky, and J.
findices; chlorophyll; nutrients) Stimson (1991) Effects of nutrient
The records for 14 years of plant micro-nutrient enrichment and water motion on the coral
concentrations and the quantities of chlorophyll in Pocillopora damicornis. Pacific Science
Liverpool Bay are reviewed. The complicated and 45:299-307.
varied hydrographical regimes which characterize [nutrient; coral; phosphorus; nitrogen; density;
the area preclude the use of the former variables as growth)
precise indicators of the extent of enrichment of the Exposure of the hermatypic coral Pocillopora
water. There is nonetheless evidence that the area damicornis (Linnaeus) to elevate levels of dissolved
can sometimes support high standing crops of inorganic phosphorus did not affect the colony or
phytoplankton. Advective dispersal processes and the zooxanthellae. Exposure to elevated levels of
variations in their intensity are probably dissolved inorganic nitrogen and inorganic nitrogen
particularly important in controling the magnitude + phosphorus led to an increase in algal density,
of these crops of phytoplankton. It is suggested and as a result, to an increase in the chlorophyll
that adequate long term records of the frequency and concentration. These latter two experimental
durations of such blooms are likely to be a more enrichments slowed skeletal growth rate of the
useful indication of the extent or eutrophication of corals, probably because of a decrease in the
the area and of any long term trends which might photosynthetic rate of the algae and perhaps a
occur than are the hydrochernical data. decrease in the translocation of photosynthetic
products from the algae to the coral. The algae

98
probably used the photosynthetic energy for their annual production varies from lows of 150 g C m-2
own increased growth. Experimental manipulation in Fraser River plumes to highs over 500 g C m-2
of water motion used in these experiments did not in sheltered boundary waters of inlets. Recent
affect the coral or the symbiotic algae. increases in ammonia and nitrate from land drainage
and domestic sewage, mainly through the Fraser
Stockner, J. G. and D. D. Cliff (1979) River, are related to increases in phytoplankton
Phytoplankton ecology of Vancouver standing stocks in the Strait.
Harbor. Journal of the Fisheries Research Board
of Canada 36: 1 - 10. Stockner, J. G. and K. S. Shortreed (1988)
lphytoplankton; Vancouver Harbor; primary Response of Anabaena and Synechococcus
production; sewagel to manipulation of nitrogen: Phosphorus
Phytoplankton production and distribution were ratios in a lake fertilization experiment.
examined over a 2-yr period in the Burrard Inlet Lininology and Oceanography 33:1348-1361.
system, which includes a true fjord (Indian Arm), a [lake; nutrient addition; picoplankton; chlorophyll;
shallow blind inlet (Port Moody arm), and a N/P ratio; bacteria)
turbulent narrows region that is contiguous to the Additions of inorganic nitrogen and phosphorus to
Port of Vancouver. Greatest annual production selected areas of oligotrophic Kennedy Lake began
occurred in Port Moody Arm with a mean of 532 g in 1978 and resulted in development of late-summer
C m-2 yr-1 while the lowest values were in Indian blooms of nitrogen-fixing cyanobacteria Anabaena
Arm and the Narrows region, averaging about 260 circinalis in 1981 and 1982. In subsequent years the
g C M-2 yr 1. Nitrate and zooplankton grazing were blooms were successfully eliminated by increasing
the main factors limiting phytoplankton production the molar N:P ratio of added nutrients to 35: 1. The
in Indian Arm, while flushing and poor light nutrient additions also resulted in dramatic increases
conditions influenced phytoplankton growth in the in both autotrophic (predominantly Synechococcus
Narrows and outer Burrard Inlet. Most of the sp.) and heterotrophic (bacteria) picoplankton
discharges of domestic and industrial wastes have numbers, with respective maxima of 4.0 x 105 ml-
been diverted to the Fraser River, and Vancouver I and >4.0 x 106 ml- 1. Chlorophyll concentrations
Harbor can be considered relatively clean and were generally <2 ug liter- I during untreated
pollution-free because of strong tidal mixing and
seaward flushing. The only sign of eutrophication conditions and increased to as much as 50 ug liter- I
in the inlet is in Port Moody Arm where sufficient during blooms. In years following the Anabaena
nutrients from sewage discharges and a relatively blooms, hypolimnetic nitrate concentrations were
stable mixed-layer depth create near optimal significantly higher in the treated basin of the lake
conditions for phytoplankton growth. Daily than they were in an untreated basin or than they
production here is among the highest recorded in were before the blooms. Development of nitrogen-
the literature for Pacific coastal marine waters. fixing cyanobacteria blooms is dependent on both a
low N:P supply ratio and a sufficient phosphorus
Stockner, J. G., D. D. Cliff, and K. R. S. supply.
Shortreed (1979) Phytoplankton ecology of
the Strait of Georgia, British Columbia. Sukhanova, 1. N., M. V. Flint, G. Hilbaum, V.
Journal of the Fisheries Research Board of Canada Karamfilov, A. 1. Kopylov, E. Matveea, T. N.
36:657-666. Rat'kova, and A. F. Sazhin (1988) Exuviaella
(phytoplankton; primary production; chlorophyll a; cordata red tide in Bulgarian coastal
Strait of Georgia) waters (May to June 1986). Marine Biology
Observations of phytoplankton production, 99:1-8.
abundance, and distribution were made at 16 [Exuviaella; red tide; bloom; Burgas Bay;
stations in the Strait of Georgia from 1975 to phytoplankton; plankton; species compositionj
1977. The discharge of turbid Fraser River water The structure and some functional characteristics
exerts a strong influence on phytoplankton of the plankton community at the times of a
production and distribution in surface waters by Exuviaella cordata red tide were investigated in
rapid light attenuation and horizontal advection. At Burgas Bay, Bulgaria, in May and June 1986.
plume boundaries and back eddied where light Characteristics of main plankton components
conditions improve, very high production occurs (phyto-, bacterio-, nanoheterotrophic plankton,
(>4-5 g C m-2 d- 1), because of rapidly replenished ciliates and mesoplankton) in the bloom area are
nutrients supplied by the Fraser River. Advection, presented. Development of the Ecordata bloom was
turbulence, zooplankton grazing, and summer determined by abiotic conditions among which
nitrate depletion collectively impart a eutrophication and salinity decrease caused the patch
heterogeneous distribution pattern to phytoplankton character of its rapid development. Attainment of
in the surface waters of the Strait of Georgia. Mean maximum red tide (ca. I x 109 cells P 1; 1 x 103 g
111-3) from background (500 to 800 x 103 cells 1-1;

99
600 g m-3) took 3 to 7d. Growth rate (u) during lbenthos; food web; pelagic ecosystem)
that period was 1.2 to 2.2 doublings per day. A Results from studies in experimental enclosures
sharp bloom decline (3 to 4d) was caused by containing both water column and benthic
parasitic flagellates destroying the alga's communities show that the benthos has an
chloroplast. Diel biomass losses due to grazing important effect on the structure and productivity of
remained below 5%. Metabolites and degradation pelagic food webs whether that change is nutrient
products of Ecordata revealed no pronounced toxic enriched or nutrient limited. Research over a 10-yr
effects on the other components of the planktonic period in 13m-3 mesocosms showed that changes
community. The rapid bloom degradation due to in the pelagic food web were correlated with
effects of parasitic flagellates indicates the high different sediment communities and with the
potential ecosystem "self-regulation". presence or absence of a benthos. The abundance of
copepods was inversely correlated with numbers of
Sullivan, B. K. and P. V. Banzon (1990) Food macrofauna. At both low and high nutrient levels,
limitation and benthic regulation of systems without benthos had greatly enhanced
population of the copepod Acartia numbers of carnivorous holozooplankton including
hudsonica Pinhey in nutrient-limited and ctenophore, medusans, chaetognaths, and fish. Our
nutrient-enriched systems. Limnology and observations indicate that the presence of the
Oceanography 35:1618-1631. benthos shortens the pelagic food web and inhibits
(secondary production; copepods) the response of pelagic fauna to nutrient enrichment
Evidence that population size of copepods in in well-mixed coastal waters. The strength of
benthic-pelagic COUDling, which is controlled by
coastal marine ecosystems can be both resource and
predator limited was obtained from experimental the amount of turbulence in the water column, may
enclosures with and without the sediment be more important to food web structure than the
community in place. The novel finding was that rate of nutrient supply and could determine which
meroplankton and attached benthos were apparently subsystem responds to eutrophication.
effective competitors and predators of planktonic
copepods in both nutrient-limited and nutrient- Sullivan, B. K. and P. J. Ritacco (1985)
enriched systems. During the spring period of Ammonia toxicity to larval copepods in
increase Acartia hudsonica, as well as other eutropbic marine ecosystems: A
abundant species of copepods, reached considerably comparison of results from bioassays and
higher numbers in four mesocosms without enclosed experimental ecosystems. Aquatic
sediments than in four other mesocosms with the Toxicology 7:205-217.
normal sediment community in place regardless of jammonia toxicity; copepods; eutrophic marine
rate of nutrient input to the mesocosms. ecosystems; mortality;MERL)
Measurements of population parameters of A. In an experiment designed to simulate
hudsonica indicated equivalent rates of reproduction, eutrophication of a shallow coastal ecosystem,
growth, and recruitment in nutrient enriched nutrients were added to experimental ecosystems
treatments, with and without sediments. (MERL mesocosms) in six different treatment
Suppression of population growth in the presence levels. We observed large reductions in the numbers
of sediments was therefore not due to food of normally dominant copepods of the species
limitation but, by inference, to higher rates of Acartic tonsa and A. hudsonica associated with a
mortality associated with high abundances of high concentrations of unionized ammonia (NH3)
potential predators of benthic origin. In unenriched in the two most nutrient enriched treatments.
control mesocosms, resource competition with the Comparison of 48h LC50 values of 10-15 uMl-
benthos may also have contributed to low INH3 obtained from laboratory bioassays with
population densities in tanks with sediments
(reproductive rates were more food limited in concentrations of NH3 associated with increased
enclosures with sediment). These results mortality in the MERL tanks indicated that
demonstrate important interactions between benthic bioassay data correctly predicted trends of high and
and pelagic fauna that have often been overlooked low mortality as well as fluctuations in the
but nevertheless may be important in regulating numbers of copepods in MERL tanks. Actual
dynamics of pelagic copepod populations in mortality rates of the mesocosm copepods was
shallow, well-mixed waters. sometimes higher than predicted, however.

Sullivan, B. K., P. H. Doering, C. A. Oviatt, A. Sullivan, B. K. and P. J. Ritacco (1988) Effects
A. Keller, and J. B. Frithsen (1991) Interactions of nutrients and copper on copepod
with the benthos after pelagic food web population dynamics: A mesocosm
structure in coastal waters. Canadian study. In: M. S. Evans (ed.) Toxic
Journal of Fisheries and Aquatic Science 48:2276- Contaminants and Ecosystem Health; A Great
2284. Lakes Focus. John Wiley & Sons, Inc., New York.

100
1copepod; biomass; copper; survival; egg species in natural water. Marine Biology
production; MERLI 70:267-273.
No abstract fred tide; nutrient; phytoplankton; growth;
abundance; macronutrient; micronutrient;
Swartz, R. C., F. A. Cole, D. W. Schults, and W. chlorophyll Q
A. DeBen (1986) Ecological changes in the Three semi-continuous culture experiments were
Southern California Bight near a large conducted by using natural algal populations
sewage outfall: benthic conditions in [dominants: Skeletonema costatum, Thalassiosira
1980 and 1983. Marine Ecology Progress sp., Gymnodinium sp., Heterosigma sp. (formerly
Series 31:1-13. called Olisthodiscus luteus) and Eutreptilla sp.]
isewage; species richness; biomass; density; collected from Tanigawa Harbor, Japan, where red-
macrobenthos; Californial tides occurred occasionally. The effects of macro-
The structure of the macrobenthic community, and micro-nutrients on the growth of different
sediment toxicity, and sediment contamination species of algae were specially evaluated by a
changed greatly between 1980 and 1983 along a dialysis bag culture technique. Two types of
pollution gradient from the Los Angeles County responses for nutrients were clearly observed:
Sanitation Districts (LACSD) sewage outfalls on macro- and micro-nutrient dependency. The former
the Palos Verdes Shelf, California, USA to a group includes S. costatum, Thalassiosira sp. and
reference site in the northern Santa Monica Bay. Eutreptilla sp. which mainly stimulated their
Stimulation of the species richness, biomass, and growth by macro-nutrients. Heterosigma sp. was
density of the benthos along the 60 in contour 5 to the latter group which was stimulated by micro-
I I km from the outfalls was significantly reduced nutrients. Gymnodinium sp. showed an
in 1983 when compared to 1980. The clam intermediate type. A red-tide of Heterosigma sp.
Parvilucina tenuisculpta and the polychaete Tharyx formed during the experiment in the harbor and is
sp. A were the numerically dominant species in the discussed with the results of the present culture
area of benthic stimulation in both years. Stations experiments and some field observations.
I to 3 kni from the outfalls in 1980 were occupied
by a depauperate fauna strongly dominated by the Takano, H. (1987) Red-tides at the mouth
opportunistic polychaete Capitella spp. In 1983 of Sumida River, Tokyo, during the last
species richness and biomass significantly increased eleven years, 1976-1986. In: T. Okaichi,
close to the outfalls, Capitella spp. was much less D. M. Anderson and T. Nemoto (ed.) Red Tides:
abundant, and the assemblage was dominated by Biology, Environmental Science, and Toxicology.
ostracods of the genus Euphilomenedes that are Elsevier, New York.
characteristic of background conditions or areas [phytoplankton; abundancel
only slightly affected by wastewater discharges. At The quantities of chlorophyll a, including
the reference site in Santa Monica Bay there were pheopigmcnts, in the brackish water of Sumida
no significant changes in species richness, River were measured at a fixed station near the
biomass, or density between 1980 and 1983, and laboratory at least once in ten days from January
the brittlestar Amphiodia urtica remained the 1976 to December 1986. Values higher than 30
dominant species. Sediment contamination by most ug/I were obtained mainly from the middle of April
measured chemicals and parameters of organic to the 'end of September, and in February and
enrichment decreased on the Palos Verdes Shelf, but October in some years. The highest value was 356
not at the reference station, between 1980 and ug/l at the end of July 1977. The main growth
1983. Sediments collected within 3 km of the areas of plankton were in Tokyo Harbour.
outfalls in 1980 were acutely toxic to the Populations were swept to the sampling station by
phoxocephalid amphipod Rhepooxynnius abronius. tidal currents.
There was no significant toxicity associated with
any sediment from the shelf in 1983. These Tamminen, T. (1982) Effects of ammonium
ecological changes correspond with the reduction in effluents o n planktonic primary
the mass emission of BOD and chemical production and decomposition in a
contaminants from the LACSD outfalls between coastal brackish water environment. 1.
1980 and 1983. Severe storms in the winter of Nutrient balance of the water body and
1982-1983 may also have contributed to effluent tests. Netherlands Journal of Sea
improvements in sediment quality and the partial Research 15:455-464.
recovery of the macrobenthos. fammonia; primary production; decomposition;
nutrients; archipelago; Baltic; effluent}
Takahashi, M. and N. Fukazawa (1982) A No abstract
mechanism of "Red-Tide" formation. 11.
Effect of selective nutrient stimulation Tamminen, T. (1982) Effects of ammonium
on the growth of different phytoplankton effluents on planktonic - primary

101
production and decomposition in a of enclosed (220 1) planktonic communities were
coastal brackish water environment. 11. examined during a 12-day 32 factorial experiment
Interrelations between abiotic and biotic on the southern coast of Finland, in early August,
components of the planktonic ecosystem. at the commencement of the annual blue-green
Netherlands Journal of Sea Research 15:349-361. algal bloom in the area. Particulate and excreted
fammonia; phosphate; effluent; phytoplankton; primary productivity, heterotrophic activity (3H-
primary production; decomposition; nutrients; glucose turnover rate), 33 14C
archipelago; Baltic; abioticl P04 assimilation, -
No abstract methylamine (ammonium analogue) assimilation
and chlorophyll a were measured on four dates
Tamminen, T. (1984) Ammonium effluent (activity parameters in >3 um and <3 um size
tests with phytoplankton and fractions). Nitrogen fixation (acetylene reduction
bacterioplankton communities and with a method) was measured at the end of the experiment.
test alga. Ecological Bulletins 36:120-124. Phytoplankton responded rapidly to the nutrient
(ammonia; phytoplankton; bacterioplankton; addition, with up to a 3-fold increase of chlorophyll
primary production; effluent) a on the first date, when the single effect of
Stimulative and toxic effects of industrial ammonium was the most significant. Ammonium
effluents containing high concentrations of was the main limiting nutrient for phytoplankton
ammonium were assayed in natural brackish water production during the experiment, but also
communities of phytoplankton and significant phosphate effects and combined effects
bacterioplankton and with laboratory cultures of a of ammonium and phosphate were observed.
test alga (Chlorella sp.). Acute effluent effects on Nitrogen-fixing blue-green algae were only
planktonic communities were measured with short- phosphate-limited. The interaction of nitrogen and
term primary productivity and heterotrophic activity phosphorus cycles in the regulation of primary
assays. Effects on Chlorella were determined as productivity demonstrates that the concept of the
biomass increase during 14 d. limiting nutrient cannot be rigorously applied at
Activity of bacterioplankton was inhibited by the the community level. Hetertrophic activity was
lowest effluent additions (corresponding stimulated by both nutrients, and the results
approximately to 100 ug NH4-N 1-1). emphasized the functional role of <3 um
picoplankton in both nutrient and carbon cycles.
Phytoplankton was stimulated over a wide range of Mean percentages of the total particulate
effluent additions (5 to 1000 ug NH4-N 1-1), and assimilation were 44% (C02), 69% (P04) and 68%
toxicity was observed with effluent additions over (methylamine) in the < 3um fraction during the
1000 ug NH4-N 1-1. Chlorella showed the largest experiment (n=45-54). The result suggest that
stimulation (up to 1000% of the control), and bacterioplankton would directly affect primary
inhibition occurred only with the highest effluent production through competition for inorganic
additions (over 6000 ug NH4-N 1- 1). The different nutrients.
responses of these organisms to effluent additions Tamminen, T. (1990) Eutrophication-and the
are discussed with regard to applying effluent tests Baltic Sea: Studies on phytoplankton,
to environmental monitoring and research. A bacterioplankton, and pelagic nutrient
coupling of test results and field data is considered cycles. Dissertation for Ph.D., University of
to be a special advantage if the pollutant under Helsinki.
study is significant in the cycles of matter in the [Baltic; phytoplankton; bacterioplankton;
environment. nutrients; cycles; bloom; nitrogen; phosphorus;
Tamminen, T., S. Kaitala, K. Kivi, and J. Finland)
Kuparinen (1985) Response of a planktonic Planktonic processes were studied at the SW coast
brackish water community to single and of Finland, the Baltic Sea, in order to examine
combined additions of ammonium and causal connections relevant to eutrophication in the
phosphate in a factorial mesocosm ecosystem. The annual pattern of nutrient
experiment. In: J. S. Gray and M. E. limitation was investigated by means of
Christiansen (ed.) Marine Biology of Polar Regions experiments on several system levels, and by
and Effects of Stress on Marine Organisms. John analyzing in situ nutrient ratios. Nitrogen was
Weily & Son Ltd., New York. shown to be the basic limiting nutrient throughout
jammonia; phosphate; mesocosm; Finland; the growth seasons, and colimitation by
phytoplankton; primary productivity; model) phosphorus occurred in early summer.
Effects of single and combined addition of The shift from new production of the spring
bloom to regenerated production of the summer
ammonium (38 and 75 mg N m-3) and phosphate season was studied in terms of nitrogenous
(10 and 20 mg p -3) on the structure and function nutrition. Clear preference succession from nitrate

102

to ammonium was demonstrated, but nutrient
availability overruled inherent preferences in Thompson, G. B. and J. Ho (1981) Some
nutrient assimilation. During regenerated effects of sewage discharge upon
production, utilization of organic nutrients was phytoplankton in Hong Kong. Marine
(organic N) uptake, while bacterial regeneration of Pollution Bulletin 12:168-173.
organic P provided an important P sources also for [sewage; phytoplankton; abundance; assemblage;
phytoplankton. diversity; Hong Kong)
Research strategies for the dynamically fluctuating The coastal waters of Hong Kong constitute a
pelagic environment were outlined with special transition from estuarine conditions in the west to
emphasis on the interrelation between fluctuations more oceanic conditions in the east, with a major
in the physical environment and within the discharge of untreated sewage located at the mid-
planktonic food web at different time scales. point. Chlorophyll a was determined and net
Continuous interaction between experimental food phytoplankton was sampled at 45 stations
web fluctuations, and system modelling appears throughout this transition. Over a period of 20
inevitable in order to understand nutrient cycle months, Chlorophyll a values rarely exceeded 2 ug
phenomena in the special conditions of the Baltic 1-1 in unpolluted coastal waters. Estuarine waters
Sea. generally contained 2-6 ug 1-1 and, in waters
Nitrogen discharge was shown to promote influenced by sewage discharge, values sometimes
eutrophication in the Baltic Sea. Effective reduction exceeded 20 ug V 1. There was no evidence of a
of nitrogen loading is therefore of vital importance. reduction in taxonomic diversity in polluted areas
Taslakian, M. J. and J. T. Hardy (1976) Sewage except in summer, when the net phytoplankton was
nutrient enrichment and phytoplankton dominated by Chaetoceros spp. In the autumn and
ecology along the central coast of early winter, Skeletonenia costatum was abundant
Lebanon. Marine Biology 38:315-325. in the central polluted areas.
Isewage; phytoplankton; Lebanon; nutrients; Tolmazin, D. (1995) Changing coastal
abundance; diversity; dominance; blue-green algae; oceanography of the Baltic Sea. 1.
dinoflagellates; diatom; indicator) Northwestern Shelf. Progress in
The abundance and taxonomic diversity of Oceanography 15:217-276.
phytoplankton has been studied in relation to [Black Sea; environment; nutrient; nitrogen;
sewage pollution (proximity to outfalls) south of oxygen; hypoxia; anoxia; mortality; benthos)
Beirut, Lebanon. Surface-water samples were This article describes the hydrography of the
collected from a series of beach stations extending Northwestern Shelf (NWS), of the Black Sea
from the American University of Beirut to 20 km emphasizing the changes induced by water
south from June, 1973 to July, 1974. Samples management in the Dnieper and Dniester river
were preserved, concentrated by settling, and the basins. The existing literature and previously
concentration of each taxon of phytoplankton unpublished data have been reviewed and
enumerated in an inverted microscope. Water synthesized to describe water property fields and
samples from the vicinity of two major sewer transport mechanisms of NWS and the Dnieper and
outfalls (Carton and Khalde sewers) showed very Dniester estuaries before the early 1960s, or the so-
high concentrations of NH4+, N02-, N03- and called precontrol period, when the effect of artificial
3
P04- , a greater total concentration of river flow control upon the coastal waters was
phytoplankton, and a lower taxonomic diversity insignificant.
than samples remote from outfalls. A considerable After the hydroenergy complexes and water
variation in the occurrence of species and withdrawal and disposal systems on the river
dominance occurred along the pollution gradient. became fully operational in the early 1970s (the so-
Blue-green algae and dinoflagellates were dominant called postcontrol period), the annual river discharge
in polluted waters, while diatoms dominated in from the Dnieper and Dniester rivers had noticeably
cleaner water away from major sewage outflow. decreased and seasonal river flow patterns had been
From the dominance and relative distribution of the artificially modified. Instead of a powerful and short
taxa along the pollution gradient, certain taxa early spring flood, typical for the natural conditions
(Oscillatoria spp., Spirulina spp., Phormidium in the Dnieper river, the hydrographs in the post
spp., Synechococcus custos and S. elongatus, control period exhibit two smaller peaks of river
Gymnodinium spp., and Prorocentrum spp.) discharge of much longer period. One of them
emerge as indicator species of pollution. These (winter-early spring) is caused by intense
changes correspond to a typical degradation of a hydroenergy generation and weir discharge through
complex community to a less mature state by the the cascade of storage reservoirs. Another is
inflow of nutrient-rich sewage (eutrophication) associated with the spring flood, modified by
along a coastal region about 10 kin long. intense water consumption and storage in this
period. High average river discharge in late May-

103
early June strengthened the summer pycnocline artificial substrate along a eutrophication
which inhibits vertical mixing in the estuaries and gradient, Barbados, West Indies. Marine
coastal waters. Owing to a slow summer Ecology Progress Series 77:261-269.
circulation, the rate of natural purification of the 1corals; Barbados; settlement; reef)
entire coastal system has been reduced. This Artificial substrate settlement plates (terracotta
coupled with the increased nutrient, organic and tiles) were set out on 3 fringing reefs for a period of
pollutant transports, decreased the dissolved-oxygen 12 mo to study settlement patterns of juvenile
concentration and led to anoxic events and mass scleractinian corals along a eutrophication gradient
mortalities of marine organisms in the previously on the west coast of Barbados, W.I. A total of 716
productive regions. These effects have primarily coral planulae settled on 120 experimental plates
plagued the benthic communities along the entire after 12 mo of exposure. Statistically higher
western coast of the NWS since 1973. (p<0.05) average number of juvenile corals per
Winter convective overturn in the Black Sea plate (X = 9.2+- 3.3; N = 40) was recorded on a
reaches its maximum depth at the southern less eutrophic reef compared to 2 more eutrophic
boundary of the NWS. Thus, the NWS waters reefs (X = 6.9 +- 3. 1; N = 40 and X = 1.9 +- 1.3;
descend beneath the seasonal and main thermoclines N = 40). Differences in juvenile coral settlement
in the open sea and are spread by the prevailing between reef zones, within each reef, were
currents across the entire sea in the cold dependent on the reef s position along the
intermediate layer (CIL). By the dynamic eutrophication gradient. Statistically higher number
mechanism the projected man-made modifications of coral planulae (X = 7.2 +- 4.5; N = 60)
in the riverine-estuarine systems of the NWS will successfully settled on vertical plates compared to
affect and change the large-scale thermocline horizontal plates (X = 4.8 +- 3.3; N = 60). Coral
structure and marine life of the Black Sea. planulae did not settle on upper surfaces of
horizontal plates. In terms of relative abundance,
Tomascik, T. (1990) Growth rates of two the most common coral species in the juvenile
morphotypes of Montastrea annularis population on the experimental plates were Poritea
along a eutrophication gradient, asteroides Lamarck which accounted for 42% of the
Barbados, W.I. Marine Pollution Bulletin settled planulae, followed by Agaricia spp. (23%);
21:376-381. and P. asteroides (Pallas) (19%). Juveniles of
1coral; Barbados; growth rate; reef; growth form; Montastrea annularis (Ellis & Solander),
morphotype) Siderastrea spp. and Diploria spp., while present at
Growth rates (skeletal linear extension) of 2 northern reefs, were absent from the most
columnar and lobatc morphotypes of the reef- eutrophic reef.
building coral Montastrea annularis were compared
within and among seven fringing reefs along a Tomascik, T. and F. Sander (1985) Effects of
previously described eutrophication gradient on the eutrophication on reef-building corals. 1.
west coast of Barbados using X-ray radiography. Growth rate of the reef-building coral
Both morphotypes of M. annularis exhibit similar Montastrea annularis. Marine Biology
patterns of increasing average growth rates with 87:143-155.
improving environmental conditions. However, the (coral; reef; growth; zooxanthellae; SPM}
average growth rates of the columnar morphotype Fourteen environmental variables were monitored
are statistically higher (P<0.001) on all reefs in the at seven locations along the west coast of Barbados
study, when compared to the lobate morphotype. It on a weekly basis over a one-year period, 1981 to
is suggested that while environmental conditions 1982. The physicochemical and biological data
exert a measurable effect on the growth rates of indicate that an environmental gradient exists as a
both morphotypes, the differences in the average result of increased eutrophication of coastal waters.
growth rates between the two growth forms, under Growth rates (linear extension) of Montastrea
similar environmental conditions, may be related annularis (Ellis and Solander), measured along the
to genetic conditions. environmental gradient, exhibit high correlation
Index master chronologies computed for a with a number of water quality variables.
columnar morphotype of M. annularis indicate a Concentration of suspended particulate matter is the
general pattern of decreasing growth rates within best univariate estimator of M. annularis skeletal
the past 30 yr at each of the seven fringing reefs. It extension rates (r2 = 0.79, P<0.0001). The results
is postulated that this general pattern of decreasing suggest that suspended particulate matter may be an
growth rates may be directly related to the energy source for reef corals, increasing growth up
deterioration of water quality along the west coast to a certain maximum concentration. After this,
of the island. reduction of growth occurs due to smothering,
reduced light levels and reduced zooxanthellae
Tomascik, T. (1991) Settlement patterns of photosynthesis.
Caribbean scleractinian corals on

104

Tomascik, T. and F. Sander (1987) Effects of Barbados, West Indies. The data suggest that P.
eutrophication on reef-building corals. 11. porites is a gonochoric species with a brooding
Structure of scleractinian coral mode of reproduction, but a low incidence (2.7%)
communities on fringing reefs, Barbados, of hermaphroditism was detected in a population
West Indies. Marine Biology 94:53-75. sampled from a reef subjected to urban and
[corals; community; reef; species composition; industrial pollution. Gonadal development occurs
zonation; diversity; sedimentation} within the mesenteries between the retractor
Seven fringing reef complexes were chosen along muscles and the mesenterial filaments.
the leeward coast (west) of Barbados to study the Gametogenesis occurs during nine to ten months of
effects of eutrophication processes upon the the year, with the peak reproductive activity
scleractinian coral assemblages. The structure of occurring predominantly in the fall and winter
scleractinian coral communities was studied along a (November to January). Gametogenesis was
eutrophication gradient with a quantitative therefore loosely synchronized between colonies-,
sampling method (line transect) in terms of species however, gonad in all stages of development were
composition, zonation and diversity patterns. On present within colonies throughout the reproductive
the basis of these data the fringing reefs were season. The reproductive season of two P. porites
divided into three ecological zones: back reef, reef populations sampled from two polluted reefs began
flat, and spur and groove. Statistically discernible one to two months earlier than that of a P. porites
and biologically significant differences in population sampled from a less polluted reef. The
scleractinian coral community structure, benthic simultaneous presence of ova and larvae within a
algal cover and Diadema antillarum Philippi colony between November and April suggests that
densities were recorded among the seven fringing larvae may be released repeatedly during an extended
reefs. High correlations between environmental breeding season. No correlation was found between
variables and biotic patterns indicate that the effects the average number of gonads and polyp size.
of eutrophication processes (nutrient enrichment, However, the gonad index (average number of
sedimentation, turbidity, toxicity and bacterial gonads based on the sum of male and female
activity) were directly and/or indirectly affecting the gonads) showed an inverse relationship with a
community structure of scleractinian coral number of environmental variables. It is suggested
assemblages. In general, species diversity was most that zooxanthellae photosynthesis through reduced
sensitive in delineating among-reef, and among- light levels may significantly lower the energy
zone, differences, which were attributed to available from photosynthates to the maturing ova
intensification of eutrophication processes. Porites and/or embryos, thus depressing larval development
asteroides Lamarck, P. porites (Pallas), Siderastrea and maturation. Coral colonies samples from two
radians (Pallas), and Agaricia agaricites (Linnaeus) polluted reefs contained lower numbers of larvae
were the most abundant coral species in the than colonies sampled from a less polluted reef.
polluted southern reefs. The absence and/or low The 2:1 sex ratio observed in a P. porites
abundance of coral species previously characterized population sampled from a polluted reef may result
as well adapted to high turbidity and sedimentation from rapid asexual reproduction (fragmentation),
[i.e. Montastrea cavernosa Linnaeus, Meandrina indicating that the mode of reproduction may be
meandirtes (Linnaeus)] indicate that eutrophication influenced by environmental conditions.
processes may adversely affect these species. It is
suggested that sediment rejection abilities, Topping, G. (1976) Sewage and the sea.
combined with feeding and reproductive strategies, In: R. Johnston (ed.) Marine Pollution. Academic
are the primary biological processes of scleractinian Press, London, England.
corals through which eutrophication processes [sewage; nutrients; benthos; review; oxygen;
directly and/or indirectly affect the structure of coral phytoplankton}
communities. No abstract

Tomascik, T. and F. Sander (1987) Effects of Tracey, G. A. (1988) Effects of
eutrophication on reef-building corals. eutrophication on growth and
111. Reproduction of the reef-building bioenergetics of the blue mussel,
coral Porites porites. Marine Biology 94:77- Mytilus edulis. Dissertation for Ph.D.,
94. University of Rhode Island.
freef; corals; reproduction; larval development; Imussel; growth; energetics; physiology; toxicity;
maturation; zooxanthellae; gonad index) Narragansett Bay; MERL; sewage; phytoplankton;
The sexual reproduction of Porites porites foodl
(Pallas), a shallow water hermatype coral, was Eutrophication may be an important environmental
studied over a one-year period (June, 1982 to June problems in marine water receiving excess
1983) on three fringing reef complexes lying along inorganic or organic nutrients from urban or
an eutrophication gradient on the west coast of agricultural activity. Nutrient-induced changes may

105
include both altered nutritional and toxicological various levels of organic matter to examine the
environments. Shifts in food quality may occur by relationship between level of sediment organic
the mixture of sewage sludge with natural matter and growth of individuals. Growth was
particulates of shifts in phytoplankton species independent of absolute levels of sediment organic
composition to food organisms of lower nutritional matter, but was significantly correlated with
value. In addition, the toxicity of environmental increases in the levels of sediment organic matter
conditionsmay be adjusted by environmental on addition of algal powder to the sediment. These
conditions created by eutrophication. These aspects results indicate that Capitalla species predominating
of eutrophication were examined in this research. in organically enriched areas have a physiological
Growth and bioenergetics of the blue mussel, requirement for organic materials discharged from
Mytilus edulis, was used to indicate ecosystem sources of organic enrichment, e.g. abundant
alteration caused by eutrophication. Bioenergetics growth of algae, fish farming, industrial effluent,
measurements included the physiological responses sewage, if they are to grow normally. Although
of clearance rate, assimilation efficiency, respiration association of some Capitella species with
rate and ammonia excretion. Apparatus used in the sediment organic enrichment has previously been
development of the research included the use of recognized as a reflection of their opportunistic
experimental marine ecosystems (mesocosms). characteristics, a physiological requirement for
Mesocosms were amended daily with varying organic materials discharged from sources of
amounts and sources of nutrients, and mussels were organic enrichment may be the single factor most
exposed to the mesocosm waters. Additional responsible for the concentration of Capitalla
research involved examination of effects on mussels species in organically enriched areas. However, it is
caused by a dense algal bloom in Narragansett Bay. unlikely that they can directly ingest and assimilate
A portion of one study involved documentation of such organic materials. Since the additional organic
mortality in mussels along a transect in materials are easily decomposed in the sediment,
Narragansett Bay. Capitella species may require either (1) some
Mussels are sensitive to copper and toxicity is specific microorganisms, the levels of which
strongly dependent upon environmental conditions increase with levels of the organic materials, or (2)
characteristic of a particular eutrophic state. Factors certain substance(s) produced during decomposition
which influence the eutrophication-toxicity of the organic materials.
relationship include the rate of biological removal
of Cu from the water column and the partitioning Tsutsumi, H., T. Kikuchi, M. Tanaka, K. Imasaka,
of Cu between dissolved and particulate phases. and M. Miyazaki (1991) Benthic faunal
Sewage sludge does not serve as a potential food succession in a cove organically polluted
resource for mussels, and acts to dilute out by fish farming. Marine Pollution Bulletin
available food. Food availability appears largely 23:233-238.
dependent upon plant biomass as indicated by (benthic; fauna; succession; abundance;
chlorophyll a concentration. Mussels may be community)
extremely sensitive to shifts in phytoplankton In the past two decades, fish farming using net
species composition, especially during the cages has developed in the coastal water throughout
reproductive phase of the species. A picoplanktonic Japan. Such fish farming has allowed the
algal bloom caused reduced feeding in mussels production of large amounts of valuable fish and
when present in sufficient densities. their supply to the markets in major cities on a
The results of the research indicate that regular basis. However, fish farming is often
eutrophication may profoundly effect the growth followed by serious organic pollution of the water
and physiological responses of mussels. and bottom sediments in the vicinity of the cages
Mechanisms of effects include alteration of food since approximately 90% of the food for the fishes
quality and modification of pollutant availability. results in organic discharge to the environment
Of all physiological measurements, clearance rates around the fish farm. Organic pollution of soft
were generally most sensitive. bottom sediment is apt to be accompanied by the
development of reducing conditions in the sediment
Tsutsumi, H., S. Fukunaga, N. Fujita, and M. and deoxidization of the bottom water, as a results
Sumida (1990) Relationship between of the decomposition of abundant organic matter.
growth of Capitella sp. and organic The benthic communities in the organically
enrichment of the sediment. Marine polluted areas are subjected to catastrophic
Ecology Progress Series 63:157-162. environmental disturbances. We have assessed the
(Capitella; sediments; organic matter; growth; environmental conditions and abundance of benthic
algae; opportunistic} communities in a specific cove since 1966. The
Laboratory colonies of Capitella sp., which is results of the present study clearly show the
found densely distributed in organically enriched or dramatic changes in fauna and the reduction in the
polluted areas, were cultured in sediments with abundance of members of the benthic communities

106

in the cove that have accompanied the progress of growth and effects on production of
the organic pollution associated with fish farming. plants and associated communities.
Marine Ecology Progress Series 23:179-191.
Tubbs, C. R. and J. M. Tubbs (1983) (submerged plants; community; growth; estuary;
Macroalgal mats in Langstone Harbour, nutrients; estuarine ponds; epiphytes; biomass;
Hampshire, England. Marine Pollution light; mechanism)
Bulletin 14:148-149. Eight experimental ponds containing submerged
(green algae; seaweeds; sewage; birdl vascular plants (predominantly Potamogeton
The spread of green algae over mudflats in per
fibliatus and Ruppia maritima) were subjected in
Langstone Harbour has been caused by increased duplicate to 4 levels (including controls) of
discharges of both treated and untreated sewage fertilization from June to August 1981. Seston and
effluent. The decline in some species of estuarine phytoplankton chlorophyll a increased with
birds may be a secondary effect of the blanketing of fertilization, and pronounced algal blooms were
the muds by algae, but Soulsby et al., 1982 evident under high dosage. Of the total seston,
concluded that the evidence for a causal link was phytoplankton exerted the greatest influence on
inconclusive. This note examines the points they attenuation of photosynthetically active radiation
put forward in support of this. (PAR), such that there was insufficient light for
submerged vasculat plant growth at the sediment
Turner, R. E. and N. N. Rabalais (1991) surface during blooms. An extensive epiphytic
Changes in Mississippi River water community developed on plants in all nutrient-
quality in this century. BioScience 41:140- treated ponds at densities similar to those observed
147. in nature on senescent plants. At high nutrient
Inutrients, nutrient ratios) treatments the accumulation of epiphytic material
No abstract resulted in >80% attenuation of the incident
radiation at the leaf surface. Biomass-of submerged
Turner, R. E. and N. N. Rabalais (1994) macrophytes decreased significantly under high and
Coastal eutrophication near ' the medium nutrient treatments compared to control
Mississippi River delta. Nature 368:619- and low treatments within 60d following initial
621. fertilization. Apparent production of vascular plants
(nutrientsl (based on oxygen production and 14C-bicarbonate
Changes in delivery of river-borne nutrients such uptake) was reduced at the higher nutrient
as dissolved phosphate, nitrate and silicate, owing treatments for both R. maritinta and P. perfoliatus.
to land-use changes and anthropogenic emissions, Most of this reduction in macrophyte
are known to result in eutrophication-enhanced photosynthesis could be explained by attenuation of
phytoplankton blooms-and more severe hypoxic PAR associated with epiphytic material. However,
events in many enclosed bays and seas. Although without PAR attenuance in the overlying water,
similar ecological effects might be expected on observed levels of epiphytic growth would be
continental shelves, the occurrence of such insufficient to reduce light below compensation
eutrophication has been unresolved. Here we levels needed to sustain vascular plant growth. At
present evidence of eutrophication of the the high fertilization rates, integrated primary
continental shelf near the outflow of the production of pond communities was significantly
Mississippi River, obtained by quantifying reduced with the loss of vascular plants, even
biologically bound silica (BSi) in diatom remnants though phytoplankton and epiphytic growth were
within dated sediment cores. BSi accumulation rates enhanced.
are greatest in water depths of 20 to 50 in within
100 kin of the river mouth, and have increased by Tyson, R. V. and T. H. Pearson (ed.) (1991)
as much as 100% this century. The increases were Modern and Ancient Continental Shelf
substantial by 1980, by which time riverine Anoxia. Special Publication No. 58 of
nitrogen loading had doubled relative to the The Geological Society. The Geolocical
beginning of the century, even though silica Society, London.
loading had declined by 50% over the same period, No abstract
Thus changes in river-borne nutrient loadings can
modify coastal coastal food webs and affect the Ursin, E. and K. P. Andersen (1978) A model
amount and distribution of oxygen in bottom water of the biological effects of
on the scale of continental shelves. eutrophication in the North Sea. Rappots
et Proc&-verbaux des Wunions
Twilley, R.R., W. M. Kemp, K. W. Staver, J. C. Counseil International pour LExploration de la
Stevenson, and W. R. Boynton (1985) Nutrient Mer 172:366-377.
enrichment of estuarine submerged (model; North Sea; nutrients; yield; fish)
vascular plant communities. 1. Algal

107
In a model of the pathways of phosphorus Large amounts of organic waste-water are
through the North Sea ecosystem, first, a steady discharged into the Ems-Dollard estuary, mainly in
supply of nutrients from rivers etc. was simulated, autumn and early winter. In November 1975 and
and next, an increasing supply. With apparently June 1975 a number of samples were taken from
realistic data for water exchange and nutrient supply the sediment of the tidal flats and analyzed for
the model predicted a slight increase in the yield of bacterial numbers, organic carbon and mud content.
fish, far from sufficient to describe the observed Cores were taken to quantify macro- and meiofauna.
yield increase over the period 1960-1970. Oxygen saturation was measured in the water
column over the sediment.
Valente, R. M., D. C. Rhoads, J. D. Germano, and In November the oxygen saturation was less than
V. J. Cabelli (1992) Mapping of benthic 20% in a considerable part of the Dollard. It is not
enrichment patterns in Narragansett Bay, clear to what extent the inward increase in organic
Rhode Island. Estuaries 15:4-17. matter content of the sediment was influenced by
(sediments; redox potential; remote sensing; the waste-water discharge.
benthos; methods; stage; infauna; assemblage; The correlation between the viable counts of
bacteria; habitat; review; Narragansett Bayl aerobic heterotrophic bacteria and the organic matter
A synoptic reconnaissance survey was performed content, as observed in the other parts of the
over a five-day period in August 1988 to assess estuary, was absent in the most polluted part of the
benthic habitat quality throughout Narragansett Dollard. However, the differences between the
Bay, Rhode Island, using REMOTSO analysis numbers counted in November and June were not
sediment-profile photography and analysis in significant in that area.
combination with measurements of the levels of Numbers of sulphate-reducing bacteria decreased
Clostridium perfringens spores (a fecal indicator) in sharply between November and June, indicating a
sediments. Three main areas of degraded benthic dependency on the waste-water discharges. Numbers
habitat quality related to either excessive organic of sulphide-oxidizing bacteria decreased in summer
enrichment or physical disturbance were identified concomitantly with the sulphate-reducing bacteria.
based solely on the REMOTSO analysis: the These data suggest that, in contrast to the other
Providence River Reach, Greenwich Bay and its parts of the estuary, in the Dollard obligate
associated coves and harbors, and an area located autotrophs play a predominant part in the
populations of sulphide-oxidizing bacteria during
along the southwest side of Prudence Island. the time of high waste discharges.
Sediments at many stations in these areas exhibited Population diversities of meio- and macrofauna of
shallow apparent redox-potential discontinuity the tidal flats were strongly decreased in the vicinity
(RPD) depths, high apparent oxygen demand, and of the outfall. Total biomass of macrofauna
low-order benthic successional stages. Elevated
Clostridium perfringens spore counts in surface decreased to zero, whereas meiofauna biomass
sediments were attributed to inputs from wastewater significantly increased. Some possible explanations
treatment facilities. The highest spore counts for the effects on the benthic fauna are given.
occurred at the head of the bay, where wastewater Veer, H. W. v. d. (1989) Eutrophication and
treatment discharges and associated combined sewer mussel culture in the western Dutch
overflows are numerous. Using data from the
Wadden Sea: Impact on . the benthic
REMOTSO analysis and the sediment inventory of ecosystem; A hypothesis. Helgolander
C. perfringens spores a distinction was made Meeresuntersuchungen 43:517-527.
between organic enrichment of the bottom from Imussel; Dutch Wadden Sea; benthos; nutrient;
sewage, versus nonsewage enrichment or physical macrofauna; biomass)
disturbance. The combination of techniques Since 1950, two large-scale changed have taken
employed in this investigation could be used to place in the western Dutch Wadden Sea, namely the
design more efficient monitoring to assess eutrophication of the area and introduction of an
eutrophication effects in estuaries and determine the extensive mussel culture. Although eutrophication
effectiveness of regulatory or management in the fresh waters started already around 1950,
initiatives to reduce organic overenrichment of nutrient concentrations in the western Wadden Sea
benthic habitats. remained fairly constant until about 1970, due to
Van Es, F. B., V. A. M.A., L. A. Bouwman, and the retention of nutrients in Lake Ussel, the main
H. G. J. Schr6der (1980) Influence of organic source. From 1970-1980 concentrations increased
strongly, and during the last years the situation has
pollution on bacterial, macrobenthic and stabilized. Mussel culture was introduced in 1950
meiobenthic populations in intertidal and expanded during the next decade to an area of 70
flats of the Dollard. Netherlands Journal of km2, all situated in the sublittoral area. From 1960
Sea Research 14:288-304. the area of mussel culture remained about constant
(oxygen; bacteria; benthos; biomass; diversity; with fluctuating yields of between 35 and 120
Ems estuary; tidal flat; macrofauna; meiofaunal

108

millions of kg fresh weight. Due to a lack of data sediments in the Northern Adriatic Sea, in the area
for the period until 1970 the impact of of Rovinj (Yugoslavia), was investigated,
eutrophication and mussel culture on the comparing meiofaunal density at the stations which
ecosystem, cannot be assessed. From 1970 onwards are under direct influence of sewage and the stations
an increased biomass and production of the which are far from the source of pollution. It is
macrofauna in the intertidal zone has been observed, concluded that raw domestic sewage does not have a
which is attributed to eutrophication. The negative influence on the density and distribution of
hypothesis is postulated that the introduction of meiofauna, but Parker's index of pollution indicates
mussel culture between 1950 and 1960 has resulted that meiofauna at these stations were under stress.
in an increased food competition in the area, leading
to a decreased stock of the macrofauna in the Vogt, H. and W. Schramm (1991)
intertidal. Eutrophication from about 1970 onwards Conspicuous decline of Fucus in Kiel
has improved the food conditions and as a result Bay (Western Baltic): What are the
both in macrofauna in the intertidal and the mussel causes? Marine Ecology Progress Series 69:189-
in the sublittoral area would have increased in 194.
biomass, allowing higher maximum yields of the (seaweeds; Baltic; Fucus; model; biomass)
mussel culture. The importance of monitoring In the framework of investigations on possible
programs is stressed to follow these trends in the effects of eutrophication on the macrophytobenthos
near future and to check the above hypothesis in in Kiel Bay (Western Baltic), a large-scale survey of
areas where it is decided to introduce or intensify the distribution and occurrence of the genus Fucus
mussel culture. was carried out in 1987/88. For large-scale
quantitative mapping, underwater television was
Veer, H. W. v. d., W. v. Raaphorst, and M. J. N. employed or direct observations from the water
Bergman (1989) Eutrophication of the surface were made. Species composition and
Dutch Wadden Sea: External nutrient quantitative biomass data were obtained by dredging
loadings of the Marsdiep and Vliestroom and SCUBA-diving. Comparison with the results
basin. Helgolander Meeresuntersuchungen of earlier investigations revealed a drastic decline in
43:501-515. Fucus biomass from between 40 000 and 45 000 t
jDutch Wadden Sea; nutrients; nitrogen; wet wt down to only 2400 t wet wt in 1987/88,
phosphorus) which means a decrease by 94 to 95 %. Whereas
The increasing P and N content in the two main Fucus vesiculosus as well as F. serriatus were still
tidal basins in the western Dutch Wadden Sea, the frequent at depths below 2 rn down to 13 rn in the
Marsdiep and the Vliestroom basin, has been seventies, during our investigation Fucus spp. was
reconstructed from the 50s onwards. The area is not found in water depths greater than 2 m.
enriched with nutrients by two sources both Possible causes for the observed changes are
originating from the river Rhine, one being the discussed, and it is concluded that decreased light
discharge from Lake IJssel and the other the levels or increased epiphyte growth as a result of
exchange with the coastal zone of the North Sea. eutrophication, and the reduction of substrate for
Due to a buffering by Lake Ussel for about 15-20 algal growth due to stone fishing and sand
years, the eutrophication of the western Wadden Sea deposition, are the main causes for the decline of
showed a time lag compared with the continuously Fucus spp.
increasing nutrient concentrations in the river
Rhine and the coastal zone of the North Sea. At Vollenweider, R. A., A. Rinaldi, and G. Montanari
present, the primary production in part of the area (1992) Eutrophication, structure and
still seems to be nutrient limited in summer, while dynamics of a marine coastal system:
loadings have already been decreasing in recent results of ten-year monitoring along the
years. So far, no severe, negative effects on the Emilia-Romagna coast (Northwest
ecosystem have been reported. Some remarks are Adriatic Sea). In: R. A. Vollenweider, R.
made on the eutrophication in other parts of the Marchetti and R. Viviani (ed.) Marine Coastal
Dutch Wadden Sea in relation to the hydrographic Eutrophication. Elsevier, Amsterdam.
characteristics of these areas. (oxygen, phosphorus, Adriatic Sea)
The process of eutrophication, which has been in
Vidakovi'c, J. (1983) The influence of raw progress for some two decades in the coastal area of
domestic sewage on density and the Northwest Adriatic Sea, shows mainly cyclic-
distribution of meiofauna. Marine Pollution seasonal features. In winter and spring, large
Bulletin 14:84-88. blooms of diatoms usually occur determing the so-
(sewage; meiofauna; abundance; density; Adriatic called "dirty waters", while in summer blooms of
Sea) dinoflagellates cause "red tides". Low salinity and
The influence of raw domestic sewage on density high nutrient loads from the River Po and most
and distribution of meiofauna in the sea bottom minor rivers favour the production of these blooms.

109
Recurrent episodes of anoxia in bottom waters Vuorinene, I. and E. Ranta (1987) Dynamics of
cause fish kills and other nuisances that impair marine meso-zooplankton at Seili,
fisheries. Algal blooms over the years, and two Northern Baltic Sea, in 1967-1975.
episodes of mucilage formation during the summers Ophelia 28:31-48.
of 1988 and 1989 have become a serious menace to Izooplankton; Baltic; Kelatellal
the regional tourist industry. Long-term (1967-1975) dynamics of meso-
zooplankton population densities at Seili (Northern
Vukadin, H. (1991) Impacts of nutrient Baltic Sea) are described. The data were obtained
enrichment and their relation to the algal from quantitative net hauls taken at 10 day internals
bloom in the Adriatic Sea. Marine Pollution during the open water season and once a month in
Bulletin 23:145-148. winter. Water salinity and temperature were recorded
(nutrients; algal bloom; sewage; stratification) concurrently. Altogether 34 taxa (species, species
Extraordinary manifestations of eutrophication in groups, instars) were frequent enough for time
the Adriatic Sea during the last few years have been series analysis.
due to the combined effects of different physico- The most abundant rotifers were Synchaeta
chemical and meteorological factors. Permanent species (S. baltica and S. monopus for the most
inputs of nutrients, particularly in the northern part). Among cladocerans Podon polyphemoides
Adriatic via river runoffs and municipal sewage and Eubosmia longispina maritima were
during calm summers, cause marked stratification numerically dominant, while Acartia spp. and
of the water column and reduction of the horizontal Eurytemora affinis hirundoides were the most
advection. These two effects provide the ideal abundant copepods. The nauplii of Balanus
conditions for single species bloom. An attempt improvisus were the most frequent and abundant
has been made to calculate the nutrient balance meroplankton.
which allows a better interpretation of algal blooms In winter meso-zooplankton biomass is dominated
in the Adriatic. by copepods (nauplii), in spring rotifers increase,
followed by cladocerans and then by copepod
Vukadin, 1. and V. Hu1ji'c (1981) Nutrient copepodids and adults in autumn. From 20 to 100%
salts in the surface microlayer and of the total meso-zooplankton biomass is
subsurface layer of Kastela Bay waters, attributable to copepods. There is considerable year-
Thalassia Jugoslavica 17:89093. to-year variation in meso-zooplankton
Inutrients; nitrogen; phosphorus; microlayer; composition. For example, biomass in the
subsurface; Kastela Bayl summers of 1968 and 1974 was dominated (40-
This paper gives values of the inorganic form of 60%) by rotifers, while in 1967, 1970 and 1973
the main nutrient salts in the surface and subsurface rotifers were less important (10-15%) in the
layers of Kastela Bay waters (the Middle Adriatic). biomass.
Analyses of all nutrients dissolved show an The salinity at Seili rose substantially during
enrichment in nutrients in the surface layer if 1970-1971, followed by another inflow of saline
compared with the subsurface water. Only silicon water in 1973. These salt water intrusions kept the
in the surface layer showed no significant salinity at an elevated level to the end of the
enhancement with respect to silicon in the sampling period. The meso-zooplankton taxa
subsruface water. responded to the �alinity change: I I out of 34 taxa
increased in numbers (seasonal effect removed),
Vuorinen, I. and E. Ranta (1988) Can signs of while most (23) of the taxa decreased in numbers.
eutrophication b e found i n the However, in most cases these changes were not
mesozooplankton of Seili Archipelago distinguishable from random fluctuations. Presently
Sea? Kieler Meeresforschungen, Sonderheft it is difficult to assess the significance of other
6:126-140. factors (human-caused eutrophication, or biotic
Imesozooplankton; Archipelago Sea; hydrography; interactions) to the documented abundance changes.
composition)
In this study, we evaluate whether it is possible Wxrn, M. and S. Pekkari (1973) Outflow
to distinguish the effects of eutrophication from studies. Nutrients and their influence on
other, i.e. hydrographical, factors affecting the algae in the Stockholm Archipelago
zooplankton. We illustrate our arguments with during 1970. No. 1. Oikos Supplymentum
examples from an old set of zooplankton data 15:155-163.
which includes 9-year records of mesozooplankton Inutrient; nitrogen; phosphorus; Baltic)
and hydrography collected from Seili, off the south When comparing 1970 with other years in order
coast of Finland. The present study shows that to study the effects of different treatments of sewage
hydrographic changes override or mask, at least the water in the Stockholm Archipelago some strong
time period studied, possible eutrophication effects. influences depending upon climatic causes have to
be taken into consideration such as,

110
1) the accumulation of nutrients inherited from the in the sewage treatment resulting in a high N : P
years before, ratio (= 65 : 1) in the discharge. In the near future
2) the discharge below the lasting ice cover, also the nitrogen discharge will be reduced, and it is
3) the vigorous spring flood, and feared that a lowered N : P ratio in the receiving
4) the extremely small discharge from Lake waters may favour an undesired increase of nitrogen
Mdlaren during the summer. fixing blue-green algae. This study is focused on
The years 1968-1970 had 3) and 4) in common. the development of a common nitrogen fixing
In the Baltic proper an upwelling ?f phosphate- species in the area, Aphanizomenon flos-aquae, at
rich water appeared in 1970. In the Aland Sea the varied nutrient loadings in 1983-85. The biomass
N:P ratio varied between 11: 1 and 19:1 when total as well as the number of heterocysts was low near
N and total P are compared. When the mineralized the treatment plant and increased with increasing
fractions were compared (below the photosynthetic distance from the sewage discharge. The low
layer) the N:P ratio varied between 2. 1:1 and 2.6: 1. biomass and number of heterocysts in
This is far from an N:P ratio by assimilation of 7:1 Himmerfj5rdan Bay is interpreted as an effect of
(all ratios on a ug basis). competition with other algae and of low level of
phosphorus relative to nitrogen, i.e. unfavourable
Walker, D. I. and R. F. G. Ormond (1982) Coral conditions for nitrogen fixation. The abundance of
death from sewage and phosphate Aphanizomenon was highest in 1994 when the
pollution at Aqaba, Red Sea. Marine phosphorus load was higher than the other years of
Pollution Bulletin 13:21-25. investigation. As a tentative conclusion it is
tsewage; coral; Red Sea; algal growth; sediment) suggested that nitrogen fixing algae may increase as
Localized pollution of coral reef areas is occurring a result of reduction in the nitrogen discharge to
at Aqaba, Red Sea, as a result of sewage discharge, Himmerfjdrdan Bay in the future. At the reference
and as a result of spillage of phosphate dust during station outside the Bay, nutrient conditions
loading of phosphate mineral onto ships. The rate favoured nitrogen fixing algae, as interpreted from
of death of colonies of the coral Stylophora inorganic N:P ratio less than 10:1 during the
pistillata was found to be 4-5 times as great in the summers of the investigation period. Here the
polluted area as in a control area. Coral damage in variation in biomass between the years was mainly
the control area is generally caused by grazing or by due to the variation in water temperature;
extreme low tide, but the cause of coral death in the Aphanizomenon was most abundant during the
polluted area was not readily apparent. The growth warm summer of 1984 and less abundant during the
of algae, both on damaged corals , and on glass cold summer of 1995. A significant correlation was
slides placed out in the reef, was greatly stimulated obtained between the heterocyst frequency and
in the polluted area, but it appeared that such algal concentration of inorganic phosphorus in the
growth was not the direct cause of coral death. trophogenic layer at the reference station.
Corals in the polluted area may be under stress
because of reduced light intensity, inhibition of Walsh, J. J., G. T. Rowe, R. L. Iverson, and C. P.
calcification by excess phosphate, and increased McRoy (1981) Biological export of shelf
sediment load. It was found that in the polluted area carbon is a sink of the global C02 cycle.
there was a greater weight of sediment settling on Nature 291:196-201.
the glass slides for a given weight of algae. But in [carbon; cycle; nutrients; shelf; C021
addition, since algal growth was faster in the Measurements of carbon metabolism, production
polluted area, the sediment load was increased by and exchange along food webs suggest that large
the sediment trapping capacity of the enhanced algal fractions of the organic matter produced on
growth. Thus, it is suggested that increased algal continental shelves must be exported to continental
growth stimulated by increased nutrient slopes. The annual loss of organic matter from
concentrations may be important in greatly continental shelf ecosystems is far greater than in
increasing the sediment load experienced by corals. the open ocean. If part of the loss of nearshore
primary production has increased in those coastal
Wallstr6m, K. (1988) The occurrence of zones where anthropogenic inorganic nutrient
Aphanizomenon f 1 o s - a q u a e supplies have been consistently increasing since the
(Cyanpphyceae) in a nutrient gradient in industrial revolution, then burial and diagenesis of
the Baltic. Kieler Meeresforschungen, this material in slope depocentres could represent
Sonderheft 6:210-220. the 'missing BMTs of the carbon' in global C02
Iblooms; blue-green algae; N/P ratio; nitrogen; budgets.
phosphorus)
The investigation area, Himmerfjdrdan Bay Wang, Z. (1987) Evaluation of water
situated at the Swedish coast of the northern Baltic quality in the Zhujiang Estuary, China.
Proper, receives waste water from a sewage In: T. Okaichi, D. M. Anderson and T. Nemoto
treatment plant. Phosphorus is efficiently reduced

ill
(ed.) Red Tides: Biology, Environmental Sciences, Water movement generated by onshore swells
and Toxicology. Elsevier, New York. appeared to be a major influence on total
Jphytoplankton; abundance; red tide) sedimentation both in time and space. The amount
This paper presents preliminary results of a and proximity of bottom sediments also played a
monthly comprehensive investigation of major role in determining spatial differences.
eutrophication and red tides in the Zhujiang (Pearl Total sedimentation in the area of the outfall was
River) estuarine area, beginning in February, 1987. disrupted. Pronounced gradient in total
The purpose of this paper is to discuss the trophic sedimentation were demonstrated both laterally
level of water quality in the Zhujiang estuary (parallel to shore) and in a seaward direction.
during Spring and Summer. A single parameter as However, these differences appeared to derive from
well as a multiparameter Trophic State Index was the outfall structure and the local topography rather
adopted to assess the trophic status in the estuarine than the direct sedimentation and accumulation of
area. Using nutrients (DIN and DIP), COD loading, sewage particles.
and changes in phytoplankton (cell counts and The lack of organisms at the near sewer station
chlorophyll a) as the major parameters, a synoptic was attributed to (1) the scouring of organisms
assessment was made. from the substrate when total sedimentation was at
a high level, (2) burial of available substrate during
Weaver, A. M. (1978) Aspects of the effects periods of high sediment accumulation.
of particulate matter on the ecology of Like total sedimentation, macrodetritis
kelp forest (Macrocystis pyrifera) (L.) (fragmented plants and animals) content of the
C.A. Agardh) near a small domestic trapped sediments displayed a seasonal behaviour,
sewer outfall. Dissertation for Ph.D., Stanford generally increasing and decreasing with total
University. sedimentation.
(particulate; sewage; Ulva ; sedimentation; kelpl The fluctuations in macrodetritis levels appeared
An investigation of the relation of particulate to be a major influence upon the levels of organic
matter to the ecology of a kelp forest was carbon in the particulates within the study area.
undertaken near a small sewer outfall located on a The outfall station had consistently higher
flat shale reef in I I in of water which supported a macrodetritis levels than adjacent stations and the
canopy of the kelp Macrocystis pyrifera. The fall macrodetritis at all station within the sewage
sewage effluent had an unusually high particulate filed contained large quantities of detached Ulva not
content which varied several fold through time. The found elsewhere.
area immediately in front of the outfall terminus, as The occurrence of the Ulva bloom (which took
compared to adjacent stations, was conspicuously place annually for three consecutive years) appeared
more turbid, had higher sedimentation and many to be based on: 1) the high water clarity during
patches of exposed shale devoid of organisms, periods of low sedimentation which allowed
including Macrocystis. sufficient light for Ulva growth at 11 in depth; 2)
The purpose of the study was fourfold: 1) to the presence of sewage effluent which is known to
observe how the quality and quantity of particulate stimulate the growth of Ulva.
matter fluctuated through time and space; 2) to Distinct concentration gradients of copper were
determine some of the major reasons for these often, but not always, demonstrable in the
fluctuations; 3) to determine the influence of the sediments trapped around the outfall. The highest
quantity and quality of particulate matter on the concentrations were at the outfall station.
distribution of benthic organisms; and 4) to During the period of high sediment accumulation
determine if and how the sewer outfall influenced (summer), the direct sedimentation of copper with
particulate matter in the study area. particles from the sewage effluent appeared to be
Particulates were trapped over time in plastic responsible for the gradient. However, during the
tubes mounted vertically on the bottom at stations fall, copper accumulating in the detached Ulva
located within and outside of the sewage field as appeared to be responsible.
defined by coliform levels. The contents were The importance of these findings to future
subjected to various analyses. environmental surveys and the location of sewer
Total sedimentation (grams of particulate matter outfalls is discussed. The significance of the Ulva
trapped per unit time per unit area) fluctuated phenomena as a potential pathway for toxins to get
through time in a seasonal cyclic manner, increased into the food chain is also considered.
in the fall, was at its maximum in the winter and
decreased again in the spring. This pattern was Weigelt, M. and H. Rumohr (1986) Effects of
observed at 11 different stations separated by a wide-range oxygen depletion on benthic
maximum distance of 1400 meters. At any one fauna and dernersal fish in Kiel Bay
point in time, however, large differences (up to 20 1981-1983. Meeresforschung 31:124-136.
fold) in sedimentation were observed between (abundance; biomass; benthic; macrofauna;
individual stations. oxygen; anoxia; hydrogen sulfide)

112

In late summer 1981 an extraordinary and wide influence oxygen depletion through two
ranging oxygen depletion killed most of the benthic mechanisms. (1) In bottom waters, uptake rates per
fauna below the halocline (>20m) in Kiel Bay. unit volume are low, but bulk uptake is a
This catastrophic period lasted for several weeks, significant factor in oxygen depletion because of
and even areas were affected where such events had the large volumn of water involved. (2) The
never been observed before. Fishermen reported intermediate zone, where respiratory uptake is also
mass catches of moribund polychaetes and significant, is strategically located between the
migrations of fishes into nearshore waters. Only a surface zone of oxygen renewal and the bottom
few macrofauna species survived (Arctica, Astarte, zone of depletion, where it constitutes an active
Corbula, Halicryptus). filter which reinforces the pycnocline as a barrier to
About 30,000 t of macrofauna died in Kiel Bay, vertical oxygen dispersion. The magnitude of direct
but. recolonization in the following months was oxygen removal in the water column relative to
rapid. Stomach analyses of demersal fish revealed removal by sediment oxygen demand and the
significant changes in food composition of cod and potential effects of this biological filtering
dab. mechanism are important considerations for
In late summer of 1983 a similar, but not so understanding eutrophic dynamics and managing
severe oxygen depletion caused the total breakdown Long Island Sound. Dynamic models which (1)
of the Syndosmya alba stocks below the halocline underestimate the role of water column uptake and
as well as that of other species. (2) incorporate only the two-zone characteristics of
physical stratification will tend to (a) overestimate
Welsh, B. L. and F. C. Eller (1991) the contribution of sediments to summertime
Mechanisms controlling summertime oxygen deficits and (b) overestimate rates of vertical
oxygen depletion in western Long Island dispersion and reventilation of bottom waters.
Sound. Estuaries 14:265-278.
(Long Island Sound; oxygen; mechanism; Weston, D. P. (1990) Quantitative
depletion; hypoxia) examination of macrobenthic community
Physical profile data (salinity, temperature, changes along an organic enrichment
oxygen, and downwelling irradiance) and in situ gradient. Marine Ecology Progress Series
incubation of light and dark bottles were used to 61:233-244.
characterize vertical structure and elucidate fabundance; biomass; macrobenthos; community;
mechanisms controlling summertime hypoxia in trophic; size; richness; distribution; Puget Soundl
western Long Island Sound. The period of oxygen Organic enrichment, both natural and
depletion corresponded with the period of thermally- anthropogenic, is one of the most common forms
controlled stratification. Bulk density differences of disturbance in the benthos. The effects of organic
between surface and bottom waters were only 1.2 to enrichment on the benthos in the vicinity of a large
2.7 sigma-t units; but they were apparently mariculture facility were examined as a general
sufficient to resist destratification by winds and model for enrichment, without the confounding
tides. Thus oxygen depletion was a cumulative effects of toxicants often associated with
process through the summer. During the anthropogenic inputs. Stations nearest the facility
stratification period, net oxygen production were subject to continuous input of fish feed and
(measured using light BOD bottles) was confined to fecal matter, and gross structural changes in the
a narrow surface zone of 1.9-4.5 m. Below this macrofaunal community (e.g. reduced species
zone was an intermediate zone of high net oxygen richness, dominance of opportunistic species) were
uptake, beneath which was a subpycnoclinic zone similar to those commonly reported for other
where oxygen uptake was very low. Rates of total enriched sites. More complex community and
oxygen uptake (dark bottles) were greatest in the population responses were indicated by changes in
surface layer and diminished with depth. There was body size, vertical distribution of infauna and
close coupling between physical conditions and patterns of trophic dominance. Enriched areas are
metabolic structure. Vertical structure patterns of generally assumed to be characterized by
oxygen production and removal were strongest in macrofauna with small body sizes, and mean
calm weather. The location of the intermediate zone individual size did, in fact, decrease with proximity
corresponded with that of the oxycline. The to the farm. At stations farthest from the farm,
thickness of the zone and the steepness of the however, size distributions were skewed by a few
oxyclinc were detem-iined by the depth and intensity large individuals, indicating that trends in mean
of both physical stratification and biological individual size are susceptible to the
production and respiration. The biological structure methodological limitations inherent in adqeuately
was weakened by physical mixing in the upper sampling rare individuals. While interspecific
water column, and the intermediate zone disappeared measures of animal size decreased with increasing
with fall destratification. We hypothesize that enrichment, intraspecific measures indicated a
biological uptake within the water column tendency for larger individuals to occur at the most

113

enriched sites. Enriched areas may represent Wilcox, W. H. (1979) The effect of nitrogen
increased food resources, and thus increased and phosphorus enrichment on salt
potential growth and attainment of larger body sizes marsh. Dissertation for Ph.D., The University of
in those species capable of exploiting such Tennessee.
habitats. Organically enriched areas are generally (salt marsh; biomass; dominance; periphyton;
considered to be characterized by infauna living at composition; Floridal
or near the sediment-water interface. Increasing Fertilization of a Batis maritima and Salicornia
organic enrichment resulted in the loss of large, virginica dominated salt marsh in Florida, with a
deep-dwelling species and dramatically altered the combined nitrogen and phosphorus fertilizer applied
vertical biomass profiles, but, because these at four rates (50:8; 100: 16; 150:24 and 200:32
individuals were numerically few, had little effect kg/ha/yr N:P) produced the following responses: 1)
on the vertical abundance profiles. Changes in dominance shifted to Batis, 2) Batis aerial biomass
trophic dominance did occur along the trophic increased from 428 g/m2 (control) to 1062 g/m2
groups and assign species to them. The utility of (highest rate) while Salicornia showed no positive
trophic grouping approaches in identification of response to fertilization, 3) highest fertilization
enrichment-induced disturbances is thus limited. plots had a 9% higher decomposition rate than
control plots (0.00728/day versus 0.00668/day), 4)
Whiteledge, T. E. (1985) Nationwide Review no luxury consumption of nutrient was measured,
of Oxygen Depletion and Eutrophication 5) Batis stem biomass reached an asymptote of 750
in Estuarine and Coastal Waters. NOAA, g/m2 (295 g/m2 control) after 2 years fertilization
loxygen; estuary; coastal; nutrients; production; at 150:24 and 200:32 kg/ha/yr N:P, 6) Batis
United States; hypoxia; depletion} biomass at 328 g/m2 (133 g/m2 control) had not
No abstract reached an asymptote after two years fertilization.
Widbom, B. and R. Elmgren (1988) Response Measurement of initial changes in snail numbers
of benthic meiofauna to nutrient and periphyton biomass and species composition,
enrichment of experimental marine in response to fertilization could not be repeated,
ecosystems. Marine Ecology Progress Series probably due to a drying trend in the marsh and a
42:257-268. drastic reduction in snail and periphyton numbers.
(benthic; abundance; composition; meiofauna; Wittberg, M. and W. Hunte (1992) Effects of
nematode; polychaete; harpacticoid; bivalve; eutrophication and sedimentation on
ostracod; kinorhynch; MERQ juvenile corals. 1. Abundance, mortality
The long-term (2.4 yr) response of benthic and community structure. Marine Biology
meiofauna to eutrophication of experimental marine 112:131-138.
ecosystems was studied at the Marine Ecosystems (sedimentation; corals; abundance; mortality;
Research Laboratory, Graduate School of community structure; reef; size)
Oceanography, University of Rhode Island (USA). This study investigated effects of eutrophication
Ammonium, phosphate and silicate were added and sedimentation on juvenile abundance, juvenile
daily in the mesocosms in a logarithmic mortality and community structure of scleractinian
progression (OX, IX, 2X, 4X, 8X, 16X, and 32X), corals on fringing reefs on the west coast of
with the IX addition being N = 2.89, P = 0.225, Si Barbados, West Indies, in 1989. Juvenile abundance
= 0.205 mmol m-2 d -1. Phytoplankton production was lower on eutrophic/high-sediment reefs than
and biomass in the tanks increased with increasing less cutrophic/l ow- sediment reefs, but juvenile size
nutrient enrichment. The benthic community gave was larger on the former. The larger size could
a quantitatively less marked response to the gradient result from size-selective mortality against smaller
of nutrient input. The meiofauna showed juveniles on the eutrophic reefs, or from lower
remarkably little response in terms of biomass and recruitment to the eutrophic reefs, or from faster
abundance, but significant effects were found on growth on the eutrophic reefs. Juvenile mortality
major taxa, leading to a changed meiofauna was higher on the eutrophic reefs than the less
community structure. Nematode and juvenile eutrophic reefs and may result from increased
polychaete abundance increased with increasing smothering of corals by eutrophic reefs, probably
nutrient input, especially in early summer, whereas in response to elevated nutrients and/or because
kinorhynchs, ostracods, harpacticoids and juvenile grazers (Diadema antillarum; herbivorous fish) were
bivalve decreased. The lack of a positive biomass less common on eutrophic reefs. Juvenile
response of the total meiofauna in the enriched community structure on all reefs was dominated by
tanks suggests that the meifauna was limited not Type I corals (high recruitment, high natural
only by the availability ot food, but also by biotic mortality), but Type 2 corals (low recruitment, low
interactions. natural mortality) became more common in adult
communities on the less eutrophic reefs. This
transition in community structure did not occur on

114

the eutrophic reefs, adult community structure Channel community was dominated by the crab
continuing to be dominated by Type I corals. The Portumus hastatoides Fabricius and the gast.opod
fact that the pattern of relative abundance of species Nassarius crematus (Hinds). Along an increasing
in the juvenile community is maintained in the gradient of organic pollution, Nassarius crematus
adult community on the eutrophic reefs suggests was gradually replaced by the crab Charybdis
that juvenile mortality rates of different species are vadorum (Alcock), and a community dominated by
similar on eutrophic reefs, and hence that Portumus hastatoides and Charybdis vadorum was
differences in adult community structure between found in the polluted inner Harbour. The abundance
eutrophic and less eutrophic reefs may be largely and dominance of predatory gastropods also showed
explained by interspecifics in juvenile mortality a decrease from the Channel to the inner Harbour,
becoming smaller on eutrophic reefs. reflecting changes in the trophic structure of the
community in relation to pollution. No significant
Wolff, W. J. (1988) Impact of pollution of change in the percentage of deposit-feeders was
the Wadden Sea. In: W. Salomons, B. L. found along the pollution gradient. Summer
Bayne, E. K. Duursma and U. F6rstner (ed.) mortality of benthos appeared to occur regularly in
Pollution of the North Sea. An Assessment. inner Tolo Harbour, and was attributed to summer
Springer-Verlag, Berlin. oxygen depletion resulting from eutrophication.
(phytoplankton; production; microphytobenthos; The benthic community was, however, soon
Dutch Wadden Sea; biomass; macrobenthos; restored to its original state by rapid winter
review) recolonization. It is postulated that the cyclic
No abstract phenomenon of summer mortality followed by
winter recovery may be a common characteristic in
Wong, P. S. (1987) The occurrence and subtropical benthic communities subjected to a
distributition of red tides in Hong Kong - high level of organic pollution.
Applications in red tide management.
In: T. Okaichi, D. M. Anderson and T. Nemoto Wulff, F. and A. Stigebrandt (1989) A time-
(ed.) Red Tides: Biology, Environmental Science, dependent budget model for nutrients in
and Toxicology. Elsevier, New York. the Baltic Sea. Global Biogeochemical Cycles
fred tide; bloom) 3:63-78.
The occurrence of red tides in Hong Kong is fnitrogen; phosphorus; nutrient; loading; model;
reviewed based on reports received by the budget; Baltic}
Aquaculture and Fisheries Department. Interesting Overall budgets for nutrient and humus are
patterns of distribution of some common red tide described for the Baltic Sea as well as for the
species in Hong Kong are obtained. Data on subsystems, i.e., the Baltic proper, the Bothnian
seasonal occurrence and distribution of each species, Bay and the Bothnian Sea. The residence times for
particularly the toxic ones, would enable better total phosphorus, total nitrogen, silicate and humus
planning of preventing measures or management are 13.3, 5.5, 11.2, and 9.6 years respectively,
actions in different coastal regions. Knowledge of compared to 21.8 years for a conservative substance
toxicity of different species would facilitate the (salt). About 90% of the nutrient losses are due to
initial assessment of risks involved. Further biogeochernical sinks within the Baltic Sea. Thus
suggest the possibility of development a system for only about 10% is exported to external areas (the
forecasting red tide occurrences. Kattegat/Belt Sea). For humus the corresponding
figures are about 75 and 25%, respectively. This
Wu, R. S. S. (1982) Periodic defaunation means that the Baltic Sea to a large extent can be
and recovery in a subtropical epibenthic regarded as a closed system and perturbations in the
community, in relation to organic water exchange with the North Sea should have
pollution. Journal of Experimental Marine little effect on the nutrient budgets. This sinks are
Biology and Ecology 64:253-269. parameterized by an expression borrowed from
(benthic; defaunation; Tolo Harbour; Tolo channel; limnology where the net nutrient loss is a function
Hong Kong; abundance; biomass; composition; of the winter surface concentration. A budget model
oxygen; recovery; diversity) is run in a prognostic, hindcast mode with the
A monthly survey was carried out for two years assumed time-dependent phosphorus and nitrogen
on the epifaunal. community in Tolo Harbour and loading of the Baltic proper. The computed
Tolo Channel, Hong Kong, a subtropical development of the winter surface concentrations of
embayment subjected to a gradient of organic total P and total N for the period 1950-1988
pollution. The number of animals and species, appears quite realistic. The possibility of having
biomass and species diversity (Shannon's function variable sinks which are functions of the surface
H' and eveness J), were higher at less polluted winter concentrations of nitrogen and phosphorus is
stations in the outer Harbour and Channel than at described using calculations based on data from the
the polluted stations in the inner Harbour. The different Baltic subareas. Such sinks should

115

significantly decrease the winter N:P ratio in the phosphate in June and August, 1981 and September
surface water when the nutrient loading increases 1982. Despite positive relationships between
with time. With better description of in particular nutrients and chlorophyll a, phytoplankton growth
the pools of nutrients in the sediment, it would be is not likely to be limited by these nutrients in
possible to model future changes of nutrient Jinhae Bay. The role of growth stimulators in
concentrations in the water column in relation to initiating the extensive blooms of red tide
loading. organisms in suggested.

Wulff, F., A. Stigebrandt, and L. Rahm (1990) Yi, S. K., J.-S. Hong, and J. H. Lee (1982) A
Nutrient dynamics of the Baltic Sea. study on the subtidal benthic community
Ambio 19:126-133. in Ulsan Bay, Korea. Bulletin of the Korea
(nitrogen; phosphorus; nutrient; Baltic) Ocean Research and Development Institute 4:17-
A prerequisite for understanding the large-scale 26.
eutrophication of the Baltic Sea is an understanding (benthic; Ulsan Bay; polychaete; abundance;
of the factors responsible for regional and long-term composition)
variations of nutrients. This article summarizes A quantitative investigation of the soft-bottom
recent studies on the changes in overall total macrobenthic community in Ulsan Bay was carried
amounts and the distribution pattern of nutrients. out from June, 1980 to March 1981. Of the 127
The total amounts of phosphorus and nitrogen have species of benthic organisms found, Polychaeta was
increased over the last few decades. On the other the most dominant taxonomic group with 67
hand, silicate levels have been decreasing, species which accounted for 88% of the total
indicating a higher net primary production and number of individuals. The number of species and
sedimentation of diatoms. A series of models has individuals decreased from the inner bay toward the
been used to test our present understanding of the outer bay stations. At station 1, a sharp drop in the
critical processes controlling nutrient and oxygen number of individuals was noted from June to
conditions. It is shown that a few morphometric September, and this drop was mainly due to
factors are essential, like shallow sill depth at the decrease in three dominant polychaetes, Cirratulus
entrance of estuarine-like sea and shallow mean cirratus, Lumbrineris longifolia, and Tharyx sp.
depth. It is also shown that phosphorus, nitrogen Based on interstational species similarity, the
and silicate differ in terms of residence times and study area can be divided into two zones; Station
the fate of these nutrients take place in the 1,2,3, and Station 4,5. The first zone was
sediments. A model showing the long-term considered to be under an organic enrichment
behavior of this system in response to external condition. This was supported, to a certain extent,
loading illustrates the importance of these by hydrological conditions and species
processes. composition.

Wyatt, T. and J. Horwood (1973) Model which Zambianchi, E., C. Calvitti, P. Cedcamore,
generates red tides. Nature 244:238-240. F.D'Amico, E. Ferulano, and P. Lanciano (1992)
(red tides; model; bloom; dinoflagellates) The mucilage phenomenon in the Norther
No abstract Adriatic Sea, summer 1989: A study
carried out with remote sensing
Yang, D. B. (1987) Nutrient and techniques. In: R. A. Vollenweider, R.
chlorophyll a variations during the red Marchetti and R. Viviani (ed.) Marine Coastal
tides in Jinhae Bay, Korea. In: T. Okaichi, Eutrophication. Elsevier, Amsterdam.
D. M. Anderson and T. Nemoto (ed.) Red Tides: [Adriatic Sea, gels)
Biology, Environmental Science and Toxicology. Observations are presented of the mucilage
Elsevier, New York. phenomenon which affected the Northern Adriatic
fred tides; nutrient; abundance; chlorophyll) Sea in the summer of 1989. The observations were
Tidal cycle time series distributions of nutrients made with data acquired by different spatial and
and chlorophyll a were measured at a fixed station temporal resolution satellite and airborne sensors.
located at mid-channel of Jinhae Bay, Korea. High The results of our study are presented, together with
nitrate concentrations were observed at the time of data from other sources, and analyzed in order to
low tides whereas high phosphate concentrations contribute to the understanding of the spatial and
occurred occasionally at the time of high tides. temporal evolution of the phenomenon.
Anoxic bottom water appeared to be the major
source of high phosphate in the outer bay. Zdanowicz, V. S., D. F. Gadbois, and M. W.
Chlorophyll a concentrations were positively Newman (1986) Levels of organic and
correlated with nitrate concentrations in April 198 1, inorganic contaminants in sediments and
May 1982 and June 1983. However, chlorophyll a fish tissues and prevalences of
concentrations were also positively correlated with pathological disorders in winter flounder

116

from estuaries of the Northeast United
States, 1984. Oceans '86 578-585,
Jorganic; inorganic; contaminants; sewage;
pathology; winter flounder; disease}
The Benthic Surveillance Project of NOAA's
National Status and Trends Program seeks to
determine prevalences of pathological disorders in
demersal finfishes and levels of organic and
inorganic contaminants in sediments and fish
tissues and evaluate possible correlations between
biological effects and contamination. More than
fifty of the Nation's estuaries and other coastal
water bodies are under study. Preliminary findings
from the first year of sampling in the Northeast
region of the U.S. reveal that sediments from
Raritan Bay, Boston Harbor and Salem Harbor
contained substantially elevated concentrations of
primary sewage related organic and inorganic
contaminants. PCB levels in livers of winter
flounder exceeded 10 ug/g in specimens from
Boston Harbor, but were 3.2 ug/g or less
elsewhere. Distributions of certain pathological
conditions, such as the presence of gigantic cells in
the tubular epithelium of winter flounder kidney,
paralleled the distributions of sediment
contaminants, whereas distribution of other
pathological conditions, such as winter flounder
pancreatitis, did not,

Zingone, A., M. Monitrcsor, and D. Marino (1990)
Summer phytoplankton physiognomy in
coastal waters of the Gulf of Naples.
P.S.ZN.I.- Marine Ecology 11:157-172,
[phytoplankton; Gulf of Naples; nutrients;
abundance-, diversity; bloom)
Thirteen sampling cruises were conducted at
weekly intervals in the inner part of the Gulf of
Naples in the summer of 1983 to investigate the
effects of excess nutrient inputs on phytoplankton
communities. High surface phytoplankton
concentrations (up to 1.15 x 108 cells 1-1) were
recorded, particularly near Naples harbour and along
the eastern coast, two locations that receive most of
the area's sewage and industrial discharge,
Phytoplankton populations were generally
dominated by small species, mainly diatoms, which
were associated with small phytoflagellates.
Species diversity values were relatively high
(W:53.62) in most samples, Throughout the
sampling period a high spatial and temporal
variability for phytoplankton abundances and
species composition was observed.

117
Appendix: Keyword List
abiotic Black Sea decomposition
abnormal Blanca Bay decrease
abnormalities bloom defaunation
Abra blue-green algae demand
abundance Bohai Bay demersal
acid Boston Harbor denitrification
acidity Brachionus density
activity brown tide deoxygenation
Adriatic brown tide bloom depletion
Adriatic Sea budget deposit-feeder
Aegean Sea Burgas Bay depth penetration
Alexandrium Buzzard Bay desert
algae Cadiz Bay development
algal bloom California diatom
algal growth capacity diatoms
alkaline phosphatase Capitella Dictyocha
Ammonia toxicity carbon diel
ammonia 14C production dinoflagellate
ammonium causes dinoflagellates
ammonium/nitrate ratio change Dinophysis
Ampelisca abdita Charlotte Harbor disappearance
amphipod Chesapeake Bay discharge
annelids chironomonids disease
anoxic chla dissolved oxygen
aquaculture chlorophyll distribution
aquatic vegetation chlorophyll a diversity
archipelago Chrysochromulina Dogger Bank
archipelago sea ciliates DOM
Arkona Cladophora dominance
Arkona Sea classification dredging
Asia climate Dunaliella
assay C02 Dutch Wadden Sea
assemblage coast dynamics
assessment coastal dystrophic
Atlantic menhaden coccolithophores ecosystem
Atlantic shelf colored tides ecosystem health
atmosphere community eelgrass
auhwuchs community structure effects
Aurococcus comparison effluent
Australia concept effluents
autoecology contaminants egg
bacteria copepod egg production
bacterioplankton copepods Eh
Baltic copper Elefsis Bay
Barbados coral Ems estuary
beach coralreef energetics
benthic corals energy flow
benthic algae Corophium enrichment
benthic communities Crassostrea environment
benthic fauna criteria environmental condition
benthos cnistacea epifauna
Berrow Flat culture epiphytes
bioassay cyanobacteria epiphytic
biogeochemical cycling cycle Erie
bioindicator cycles estuaries
biomass cycling estuarine ponds
bird cyst estuary
bivalve decqpod eutrophic
bivalves eutrophic marine ecosystems

118
eutrophicated heterotrophic uptake macrophytes
eutrophication historical macrozoobenthos
evidence Hong Kong Madracis
excretion Hudson River management
Exuviaella Hudson River estuary mariculture
Far East humic marine
fate hydrogen sulfide Marsdiep
fats hydrogen sulphide marsh
fauna hydrographic mats
faunal.composition hydrographic dynamics Matsushima Bay
feeding hydrography maturation
filamentous hydromedusae mechanism
Finland hypoxia Mediomastus ambiseta
fish impacts Mediterranean
fish farming increase meiobenthos
fish kill index meiofauna
fish yield indicator MERL
fishkill Indo-Pacific meroplankton
infauna mesocosm
@ord
flagellates inorganic mesoplankton
.61
floating inorganic nutrients mesozooplankton
Florida interface metabolism
flounder intertidal methane
fluorescence Irish Sea method
flux iron methods
food Italy Mexico
food web Japan microalgae
Forth estuary Kanaohe Bay microflora
fouling Kastela Bay microheterotrophs
fi-amework Kattegat microlayer
freshwater Kelatella micronutrient
Fucus kelp micronutrients
Furcellaria Kiel Bay microphytobenthos
gels Kiel Bight microplankton
kinorhynch microzooplankton
general model
German Bight lagoon mining
gonadindex Lake Kinneret model
gonads Lake Ontario modeling
Gonyaulax lake modelling
Gotland Sea lakes mollusc
grazing Laminatia morphotype
Greece larval development mortality
green algae Lebanon mucous aggregates
green tide length mudflat
Greifswald Bay life cycle mussel
growth life history Mytilus
growth form light N/P ratio
growth potential limitation N20 production
growth rate loading Nahant Bay
Gulf of Maine lobster nanoplankton
Gulf of Naples log-normal Narragansett Bay
Gulf of Patras Long Island nematode
Gymnodinium Long Island Sound Nephrops norvegicus
H2S Louisiana net ecosystem metabolism
habitat Macoma net ecosystem production
harpacticoid. macroalgae Netherlands
Hawaii macroalgal mats New Jersey
health macrobenthos New York Bay
Helgoland macrofauna New York Bight
herring macronutrient nitrate

119

nitrification plankton secondary
nitrite Po River secondary production
nitrogen pollution sediment
Nitzschia seriata polychaete sediment transport
Nodularia polychaetes sedimentation
North Sea Polydora ligni sediments
Norway Pomatoschistus settlement
Nucula population sewage
Nucula annulata Portugal shading
nuisance precipitation shelf
nuisance bloom prediction shellfish
nutrient primary production silica
nutrient addition primary productivity silicate
nutrient enrichment processes silicoflagellate
nutrient limitation production silicon
nutrient ratios productivity similarity
nutrients protein size
oligochaetes Providence River size structure
opportunistic Puget Sound Skagerrak
organic pulp Skeletonema
organic carbon Pyrodinium sludge
organic enrichment ratio smelt
organic matter recolonization spatial
Oslofjord recovery spawning
ostracod recycling species
oxygen Red Sea species abundance
oxygen demand red tide species composition
oxygen uptake red algae species richness
oyster redox potential SPM
Palmico River estuary reef stability
paper regeneration stage
particulate relationship standing crop
Parvilucina remedial action standing stock
pathology remineralization Strait of Georgia
pelagic remote sensing strategy
pelagic biology reproduction stratification
pelagic ecosystem research needs striped bass
Peridinium reservior submerged
periphyton resource submerged plants
pH respiration submerged vegetation
Phaeocystis review subsurface
Phaeodactylum Rhine succession
Philippines richness sulfate reduction
phosphorus River Elbe survival
photosynthesis rocky shore susceptibility
photosynth *etic rate rotifer Sweden
Phycodrys runoff temperature
Phyllophora salt marsh temporal
physical energy salt pond Thailand
physiology San Francisco Bay tidal flat
phytoplankton sand goby tidal marsh
phytoplankton abundance sandflat fide
phytoplankton assemblage sandy beaches tissue
phytoplankton bloom Saronikos Gulf Tokyo Bay
phytoplankton species Scandinavian tolerance
composition Scottish coast Tolo channel
picoalgae Scottish water toxic
picoplankton seagrass toxic algal exudates
pigments seasonality toxicity
Pilayella seaweeds transition

120

transparency
trend
trophic
trophic state index
trophic status
turbidity
Ulsan Bay
ultrastructure
Ulva
United States
upwelling
Uronema
Vancouver Harbor
vascular plants
vegetation
Venice Lagoon
Victoria Harbour
Visakhapatnam Harbour
Vollenweider
Wadden Sea
wastewater
winter flounder
yield
Ythan estuary
Yugoslavia
zonation
zooplankton
zooxanthellae
Zostera

*U.S. Government Printing OFFICE- 1995- 386 -543/2 5406

OTHER TITLES IN THE
DECISION ANALYSIS SERIES

No. 1. Able, Kenneth W. and Susan C. Kaiser. 1994. Synthesis of Summer Flounder Habitat
Parameters.

No. 2. Matthews, Geoffrey A. and Thomas J. Minello. 1994. Technology and Success in
Restoration, Creation, and Enhancement of Spartina alterniflora Marshes in the United States. 2
vols.

No. 3. Collins, Elaine V., Maureen Woods, Isobel C. Sheifer and Janice Beattie. 1994.
Bibliography of Synthesis Documents on Selected Coastal Ocean Topics.

ATmo

@- V-@ 1-4 - -
sp/Y,S;g

36 80 034886