Documente Academic
Documente Profesional
Documente Cultură
Donald S. McLusky
University of Stirling, Scotland, UK
Michael Elliott
University of Hull, England, UK
THIRD EDITION
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D.S. McLusky 1981
Blackie & Son Ltd. 1989
Oxford University Press 2004
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Preface
For the inhabitants of many of the worlds Users of the previous editions were kind
major cities and towns, estuaries provide their enough to welcome the framework of the
nearest glimpse of a natural habitat; a habitat book, which first outlined the estuarine envi-
which, despite the attempts of man to pollute ronment and the physical and biological fac-
it or reclaim it, has remained a fascinating tors, which are important within it. We then
insight into a natural world where energy is examine the responses of the animals and
transformed from sunlight into plant material, plants to these factors, consider the problems
and then through the steps of a food chain is of life in estuaries and why so few species
converted into a rich food supply for birds have adapted to estuaries, and then propose
and fish. The biologist has become interested a food web for an estuary. Thereafter we
in estuaries as areas in which to study the shall examine each trophic level in the food
responses of animals and plants to severe web in turn, first the primary producers
environmental gradients. Gradients of salinity, (plants and detritus), then the primary con-
for example, and the problems of living in sumers (herbivores and detritivores), and
turbid water or a muddy substrate, prevent finally the secondary consumers (carnivores).
most animal species from the adjacent sea or These chapters have been fully revised in this
rivers from entering estuaries. In spite of these third edition, to reflect our latest knowledge
problems, life in estuaries can be very abund- in these topics.
ant because estuarine mud is a rich food In the period since the publication of the
supply, which can support a large number of previous editions of this book, a vast amount
animals with a large total weight and a high of new information on pollution in estuaries
annual production. Indeed estuaries have has accumulated. It has been widely recogn-
been claimed to be among the most produc- ized that although the worlds seas are huge
tive natural habitats in the world. and may appear capable of receiving unlim-
When the previous editions of this book ited quantities of mans waste, such waste is
appeared, biologists were beginning to realize often discharged first into the confined waters
that the estuarine ecosystem was an ideal of estuaries. Many international experts have
habitat in which to observe the processes con- stated that, while the open oceans may not
trolling biological productivity. In the inter- be generally polluted, the coastal waters of the
vening period, several more estuaries and sea and especially the waters of estuaries are
their inhabitants have been studied inten- widely polluted. Thus in practice, marine pol-
sively, and it is now possible to answer many lution is often essentially estuarine pollution.
of the questions posed by the earlier edition, To reflect this large impact of mankind on
and to pursue further the explanation of high estuaries, and to consider how mankind may
productivity in estuaries and of energy util- either destroy or enrich the estuarine ecosys-
ization at different trophic levels within the tem, new chapters have been prepared in this
estuarine food web. edition. These consider pollution in estuaries,
vi P R E FA C E
and the diverse uses and abuses of the estuarine Estuaries are the most protected habitat in
habitat by man, as well as the methods used to the United Kingdom and elsewhere (meas-
study human induced changes in estuaries and ured as the percentage of the total habitat that
the ways in which estuarine management can is subject to protection orders), and for most
either monitor, control or prevent pollution or students on aquatic biology are their most
destruction of the estuarine ecosystem. accessible marine habitat. While few UK,
This latest edition therefore retains the con- European, or American Universities have
cept of the study of the ecosystem as the basis ships for access to the sea, many are located on
for our understanding of the natural world, the shores of estuaries. Thus for many
and shows that estuaries are ideal habitats for University courses, their local estuary is the
such studies. The new content and chapters marine habitat, which can be studied, while
reflect our attempts to recognize both the the open seas have to be studied only in
problems of pollution in estuaries and the films or video. Throughout the book exam-
solutions, which estuarine management can ples are drawn from Britain, Europe, and
offer, as estuarine ecosystems come under America, as well as other areas, which have
increasing pressure from a wide range of been well studied such as South Africa and
demands made by an increasing world popu- Australia.
lation. The topic of estuarine pollution, estuar- May we thank the following authors and
ine uses and abuses, and their management publishers for allowing us to use copies of
has moved on considerably in the past decade. their figures: Kai Olsen (Ophelia; 3.3, 4.1),
As many of the problems of point source Colin Moore (Royal Society of Edinburgh; 4.8),
pollution have been attended to, different Sven Ankar and Ragnar Elmgren (Ask
problems of diffuse pollution have become Contributions; 4.10), Peter Burkhill (Plymouth
apparent. In addition, while some pollution Marine Laboratory; 4.13), Tom Pearson, and
problems are seen as capable of being solved, Rutger Rosenberg (Oceanography and Marine
habitat loss and degradation is more of a Biology, Annual Review; 6.6, 6.7), Steve
threat. Hence the third edition concentrates on Colclough (Environment Agency, London; 6.9,
these as well as including sections on the 6.26), John Beardmore (Swansea University;
effects of fisheries, dredging, structures, aqua- 7.5), Blackwells Scientific Publishers (6.10,
culture, etc. on estuarine ecosystems. The 6.11, 6.23, 6.24, 7.4). Jan Elliott and Krystal
third edition seeks to include considerably Hemingway (IECS, University of Hull) are
more applied material and present case stud- thanked for preparing the complex flowcharts
ies of estuarine change. (the horrendograms!) in Chapters 6 and 7
Estuaries can be perceived as either the and Sue Boyes (IECS, University of Hull)
originators of pollutants, or more often as the is thanked for providing information for
recipients of pollution that originate on land, Chapter 8. The authors prepared other figures.
freshwater, or from the sea. The problems of Sources of information that are not mentioned
estuarine pollution and management are those in the text are listed in the reading list, which
dealing with a sheltered environment that acts has been fully updated for this edition.
as a trap for sediment and contaminants, and
as areas subject to intense pressure from Donald S. McLusky
mankinds activities such as land-claim (or Michael Elliott
reclamation). 2004
Contents
2 Life in estuaries 19
2.1 Distribution of estuarine organisms 19
2.2 Problems of life in estuaries 22
2.3 The estuarine food web 27
3 Primary producers: plant production and its availability 34
3.1 Introduction 34
3.2 Salt marshes 36
3.3 Mangroves 39
3.4 Intertidal plants and macroalgae 39
3.5 Microphytobenthos 42
3.6 Phytoplankton 43
3.7 Detritus 46
3.8 Summation of plant and microbial production in estuaries 49
4 Primary consumers: herbivores and detritivores 53
4.1 Introduction 53
4.2 The mud dwellersbenthic deposit feeders 56
4.3 The surface dwellersbenthic suspension-feeders
and algal grazers 60
4.4 Meiofauna 61
4.5 Zooplankton 64
4.6 The primary consumer community 68
4.7 Role of primary consumers in the estuarine ecosystem 70
Index 209
CHAPTER 1
1
2 T H E E S T UA R I N E E C O S YS T E M
turbidity of the water column, or in chemical For Pritchard it is the upstream limit of salt
composition including changes in nutrients, penetration, and for Fairbridge it is the
dissolved gases, and trace metals. upstream limit of tidal penetration, which in
The English word estuary is of sixteenth an un-modified estuary will always be further
century origin, derived from the Latin word inland. The Fairbridge definition also usefully
aestuarium meaning marsh or channel, which emphasizes the gradient of conditions that
is itself derived from aestus, meaning tide or may be found in a normal estuary.
billowing movement, related to the word aestas The salinity of seawater is approximately
meaning summer. A widely used definition 35, tending to be lower (33 in coastal seas and
of an estuary has been given by Pritchard higher 37) in tropical waters. The salinity
(1967) as: an estuary is a semi-enclosed of freshwater is always less than 0.5. Thus
coastal body of water, which has a free con- the salinity of estuarine waters is between
nection with the open sea, and within which 0.5 and 35. This range is generally termed
sea water is measurably diluted with fresh brackish, as distinct from marine or fresh-
water derived from land drainage. This defi- waters. Salinity is a measure of the salt content
nition of estuaries excludes coastal lagoons or of the water, and is expressed throughout
brackish seas, although we will discuss them this book using the Practical Salinity Scale (see
briefly later in this book as useful compar- Box 1.1). Whereas marine and freshwaters are
isons. Coastal lagoons, for example, do not characterized by stable salinities, estuarine
usually have a free connection with the open water is extremely variable in its salinity.
sea and may only be inundated with seawater An estuary is thus emphasized as a dynamic
at irregular intervals. Brackish seas, such as ecosystem having a free connection with the
the Caspian Sea, may have salinities com- open sea through which seawater normally
parable to some parts of estuaries, but they do enters according to the twice-daily rhythm of
not experience the regular fluctuations of the tides. The seawater that enters the estuary is
salinity due to the tide. The definition of then measurably diluted with freshwater flow-
semi-enclosed serves to exclude coastal ing into the estuary from rivers. The patterns of
marine bays and the definition of freshwater dilution of the seawater by freshwater vary
derived from land drainage serves to exclude from estuary to estuary depending on the vol-
saline lakes with freshwater from rainfall only. ume of freshwater, the range of tidal amplitude,
While many estuarine scientists have used and the extent of evaporation from the water
Pritchards definition, studies in the tidal within the estuary and may be used as a basis
freshwater regions of estuaries have sug- for the classification of estuaries.
gested that the definition of Fairbridge (1980) Three main types of estuaries can be recog-
is more suitable, namely that nized, namely positive, negative, or neutral
an estuary is an inlet of the sea reaching into a river estuaries. In positive estuaries the evaporation
valley as far as the upper limit of tidal rise, usually from the surface of the estuary is less than the
being divisible into three sectors: a) a marine or volume of freshwater entering the estuary
lower estuary, in free connections with the open from rivers and land drainage. In such a posi-
sea; b) a middle estuary subject to strong salt and tive estuary (Fig. 1.1) the outgoing freshwater
freshwater mixing; and c) an upper or fluvial estu- floats on top of the saline water, which has
ary, characterized by freshwater but subject to entered the estuary from the sea, and water
strong tidal action. The limits between these sectors gradually mixes vertically from the bottom to
are variable and subject to constant changes in the the top. This type of estuary, which is the most
river discharges
typical in the temperate parts of the world, is
The principal difference between the two thus characterized by incoming saltwater
definitions is in determining the upper limit. on the bottom, with gradual vertical mixing
T H E E S T UA R I N E E N V I R O N M E N T 3
Throughout this book salinity is expressed on For the past two decades salinity has been
the practical salinity scale, using what are called defined solely in terms of electrical conductivity.
Practical Salinity Units (PSU). Salinity is a scale Conductivity units are unfamiliar to some marine
and is thus a dimensionless unit (like pH). In scientists and therefore have been converted to a
earlier editions of this book, and many other older practical salinity scale. The salinity scale without
publications, salinity was expressed as parts per units has now been adopted as the practical scale
thousand (often abbreviated to ppt, , or g kg!1) since the numerical values are closely similar to
being the total concentration of salts in grams the values previously expressed in parts per
contained in 1 kg of seawater. The salts thousand of salt, the only obvious difference
are principally sodium and chloride ions, being the lack of any symbol. Since salinity is now
supplemented by potassium, calcium, magnesium, expressed as a scale, it is correct to say The
and sulfate ions, plus minute or trace amounts of salinity of seawater is 34, but it is incorrect to
many other ions. say 34 PSU, ppt, , or g kg!1.
30 20 10 1 Evaporation
Sea River 30 1
Sea River
Figure 1.1 Positive estuary. Freshwater runoff is greater Figure 1.2 Negative or Inverse estuary. Freshwater runoff is
than evaporation. The arrows show the pattern of circulation less than evaporation. The arrows show the pattern of
with denser marine water entering the estuary along the circulation with both seawater and freshwater entering the
bottom and then gradually mixing vertically with the estuary at the surface. Within the estuary, evaporation
outgoing surface stream of fresher water. Salinity is produces denser high-salinity water that sinks to the bottom.
expressed in PSU (see Box 1.1). Salinity is expressed in PSU (see Box 1.1).
leading to an outgoing stream of fresher causes the surface salinity to increase. This
surface water. saltier surface water is then denser than the
In negative (or inverse) estuaries the oppo- water underneath and thus sinks. The circula-
site situation exists, with evaporation from the tion pattern is thus opposite to that of a positive
surface exceeding the freshwater runoff enter- estuary, because in a negative estuary the sea
ing the estuary. This type of estuary is mostly and freshwater both enter the estuary on the
found in the tropics, for example, the Laguna surface, but after evaporation and sinking they
Madre, Texas, or Spencer Gulf and Gulf leave the estuary as an outgoing bottom
St Vincent, South Australia, although it can also current (Fig. 1.2).
occur in temperate regions where the fresh- Occasionally the freshwater input to the
water input is very limited such as the Isefjord estuary exactly equals the evaporation and in
in Denmark. The Mediterranean and Adriatic such situations a static salinity regime occurs
Seas are regarded as prime examples of inverse (Fig. 1.3). Such an estuary is termed a neutral
estuarine systems, as well as the Arabian Gulf, estuary, but they are rare, as evaporation and
the Red Sea. In negative estuaries evaporation freshwater inflow are almost never equal.
4 T H E E S T UA R I N E E C O S YS T E M
Fjord 2 1
Fjard 5 6
Ria 3 2
Figure 1.3 Neutral estuary. Freshwater runoff is equal to Coastal plain 35 31
evaporation. This situation is intermediate, and possibly Bar-built 6 8
transitory, between a positive estuary (Fig. 1.1) and a
Complex 18 17
negative estuary (Fig. 1.2). Salinity is expressed in PSU
(see Box 1.1). Barrier beach 2 3
Linear shore 4 6
Embayment 25 26
In this book the emphasis throughout will Source: Davidson et al. (1991).
be on the commoner positive type of estuary.
In general, estuaries are a habitat intermediate
between the sea, the land, and freshwaters: A increased as ice caps have melted, so in some
habitat, which is a complex dynamic mixture areas the land has also subsided and in other
of transitional situations and is almost never areas the land that was formerly glaciated has
static and where physical and chemical factors sprung up as the weight of ice on it has melted.
show marked variations. Many of these fac- Many estuaries of South Africa and
tors are closely linked to the salinity distribu- Australia are of the bar-built type, but in these
tion patterns mentioned above, such as the countries the sandbars may seal the estuary
strength of currents, tidal amplitude, wave from the sea during the drier months of the
strength, and deposition of sediments as well year. The estuary is thus only opened when
as temperature, oxygen, and the supply of the sand bar is breached either by high river-
nutrients. ine discharge or by artificial means (such as a
Just as the physical and chemical factors dredger). During the often lengthy periods
within estuaries are in a continual state of when they are closed these so-called blind
change, so even the topography of estuaries is estuaries do not have a free connection with
continually changing. Most estuaries have the open sea, considered to be crucial for the
developed on the low-lying ground that forms Pritchard and Fairbridge definitions of an
the coastal plains around landmasses. Here estuary. In order to accommodate the peculi-
estuaries typically develop in river valleys arities of these systems Day (1980) redefined
that become drowned by the sea. In the an estuary as a partially enclosed coastal
drowned valleys extensive deposition of sedi- body of water which is either permanently or
ments carried into the estuaries have pro- periodically open to the sea and within which
duced the estuaries that we see today. The there is a measurable variation of salinity due
various types of estuaries are described in to the mixture of sea water with freshwater
Box 1.2, with their frequency in the British derived from land drainage.
Isles, as an example, seen in Table 1.1. Many As siltation of estuaries occurs and salt
estuaries are drowned river valleys that have marshes extend and consolidate the land, so
become estuaries since the last Ice Age, which the waters of the estuary may be pushed
is within the last 1015,000 years. The height seawards. Small changes in sea level due to
of the sea in relation to the land is continually long-term climatic change could either drown
changing, and while the mean sea level has or expose many of our present estuaries,
T H E E S T UA R I N E E N V I R O N M E N T 5
Box 1.2 Types of estuaries (after Fairbridge and Davidson et al. 1991)
Fjords. Drowned glacial troughs. Fjord-type drowning so that they have a characteristic bar
estuaries occur where valleys have been deeply across their mouths, for example, Alde, England;
eroded by glaciation. Characterized by deep inner Ythan, Scotland. Barnegat Bay, New Jersey,
basins linked to the sea by shallow entrance sills, Laguna Madre, Texas, Albufeira, Portugal, and
for example, Sea lochs in West of Scotland, Fjords most estuaries of North CarolinaFlorida coast.
in Norway, Sweden, Alaska, British Columbia, In many estuaries in South Africa and Australia
New Zealand, such as the coasts of Norway, the bar may seasonally close the estuary, creating
Western Scotland, Alaska, and New Zealand. closed or blind estuaries.
Fjards. Typical of glaciated lowland coasts. More Complex estuaries. Drowned river valleys of
complex than fjords, with a more open and complex origin, typically a mixture of glaciation,
irregular coastline, for example, Solway Firth river erosion, and sea level rise, for example,
England/Scotland, eastern Canada, and Scottish Firths: Solway, Moray, Dornoch, Tay, and
New England. Forth. San Francisco Bay is a complex estuary
Rias. Drowned river valleys, formed by subsidence created by tectonic activity (land movement due
of land and/or a rise in sea level. Deep, narrow to faulting).
channels with a strong marine influence, for Barrier beaches. Open coast system where a
example, Estuaries of Cornwall, England and bar or barrier develops offshore, and an estuary
Brittany, France. is thereby created behind the barrier. For
Coastal plain estuaries. Formed by the flooding of example, North Norfolk Coast, Lindisfarne,
pre-existing valleys. Unlike Rias, these estuaries England.
are often very shallow and filled with sediment so Linear shore sites. Formed where the shore is
that extensive mudflats and saltmarshes occur. sheltered, for example, by barrier islands. Usually
Commonest type of estuary in United Kingdom, considered as a subdivision of a complex estuary.
for example, Severn, Dee, Humber, Thames, For example Essex and North Kent coast, England.
England. Chesapeake Bay, Charleston Harbor, Embayments. Large natural areas formed between
Delaware Bay, USA. rocky headlands that naturally fill with soft
Bar-built estuaries. Also drowned river valleys, but sediments. For example, Carmathen bay, Wales,
recent sedimentation has kept pace with their Morecambe bay, The wash, England.
although of course new ones may be formed 2. Fjord. ExamplesNorwegian fjords, West
elsewhere. The various types of estuary, of Scotland sea lochs. Silver Bay (Alaska),
however formed, are all characterized by the Alberni Inlet (British Columbia)
variability of their environmental parameters, 3. Partially mixed. ExamplesJames River
most notably in their water circulation and (US), Mersey, Thames, Humber, Forth (UK),
their sediments. Elbe (Germany)
4. Homogeneous. ExamplesDelaware, Raritan
1.2 Estuarine circulation (US), Solway Firth (UK), Mandovi-Zuari (India)
Within the commonest positive estuaries
In the first of these, the highly stratified or salt
four main types can be recognized depending
wedge, the freshwater flows seawards on the
on the tidal amplitude and volume of fresh-
surface of the inflowing saltwater. At the inter-
water flow (Dyer 1997):
face between the salt and freshwater, entrain-
1. Salt wedge or Highly stratified. Examples ment (mixing) occurs and saltwater is mixed
Mississippi (US), N. Esk (UK), Vellar (India) into the outflowing freshwater (Fig. 1.4). For
6 T H E E S T UA R I N E E C O S YS T E M
1 2 3 4 1 2 3 4
Fresh Fresh
Salt
Salt
1 2 3 4 1
Depth
4
Depth
3
2
will dominate the right-hand side. In the Depending on their tidal range estuaries can be
Southern Hemisphere, this pattern is reversed. classified as microtidal, mesotidal, macrotidal,
When the estuary is very wide the Coriolis or hypertidal (Box 1.3). The tidal amplitude
force will cause a horizontal separation of the may considerably influence the mudflats and
flow, with outgoing flow on the right-hand side vegetation of an estuary. While microtidal estu-
in the Northern Hemisphere, and ingoing flow aries have only limited intertidal areas, mesoti-
on the left-hand side. Thus the circulation in dal estuaries often develop extensive intertidal
such a homogeneous estuary will be across the areas, which are covered in vegetation, such as
estuary from left to right, rather than vertically
as in the other types (Fig. 1.7).
The range of tidal amplitude varies in a con-
stant pattern in the seas of the world and the Fresher Salt
tides within estuaries are due to the tidal wave
at their mouth. Tidal amplitude varies not
only from place to place but also varies at any
particular locality according to the neap and
Fresh
spring tide cycle, as the range of tides fluctu-
ates from a maximum rise and fall at spring
tides to a lesser rise at neap tides. The full Right Left
cycle from spring to neap tides occurs each
lunar month (28 days). In the waters around
Denmark and Sweden, for example, the tidal
amplitude is very small and the many estuar-
Salt
ies of Denmark are not subject to strong tidal
currents and do not develop large intertidal Northern hemisphere
areas. At the other extreme the estuaries of the
Figure 1.7 Salinity distribution in a large homogeneous
Canadian Bay of Fundy region experience max- estuary in the northern hemisphere. Due to the Coriolis force,
imal tidal amplitude producing strong tidal horizontal separation occurs, and the circulation is across the
currents as well as large intertidal areas. estuary rather than vertically. (Modified after Dyer 1997.)
Spartina, whereas macrotidal and hypertidal Table 1.2 Classification of estuarine divisions
estuaries more typically have bare mudflats
without large plants, but with microscopic Division Tidal Salinity Venice system
(1959)
microphytobenthos.
In all estuaries the strength of tidal and river River Non-tidal "0.5 Limnetic
currents is dissipated as the main inflow cur- Head The highest
rents collide and mix with each other, and point to
compared to both the sea and rivers, estuaries which tides
are quiet places. Due to the shelter of the land reach
surrounding the estuary, wind-driven waves Tidal fresh Tidal "0.5 Limnetic
are much smaller than in the sea. This absence Upper Tidal 0.55 Oligohaline
Inner Tidal 518 Mesohaline
of large waves, coupled with the mixing of
Middle Tidal 1825 Polyhaline
currents is vital for the deposition of estuarine
Lower Tidal 2530 Polyhaline
sediments. Mouth Tidal #30 Euhaline
The river flow, tidal range, and sediment
distributions in estuaries are continually Source: McLusky (1993).
changing and consequently estuaries are never
really steady-state systems. With increased
river flow, or during neap tides, the extent of inflow and deep outflow also occurs for 20% of
tidal intrusion of seawater diminishes, while the year, and four other patterns occur for the
with spring tides or decreased river flow the rest of the time in response to climatic events.
tidal intrusion increases. The concept of flush- The salinity at any particular point of an
ing time is one attempt to relate the fresh- estuary depends on the relationship between
water flow to the tidal range, and the volume the volume of tidal seawater and the volume of
of the estuary. The flushing time (T) is the time freshwater entering the estuary, as well as the
required to replace the existing freshwater in tidal amplitude, the topography of the estuary,
the estuary by the river discharge, and is cal- and the climate of the locality, but in general
culated by dividing the total volume of river it is possible to recognize various zones or
water accumulated in the estuary (Q) by the divisions within an estuary (Table 1.2)
river flow (R) (Dyer 1997) thus: T $ Q/R. While the salinity of the water bodies of
In the Forth estuary for example, the flush- estuaries is important for fish and planktonic
ing time under mean river flow conditions is organisms living in the water column, it is of
12 days, but in summer with reduced river less direct importance for the majority of estu-
flow it may be up to 10 weeks, and following arine animals, which live buried within the
severe rainfall it may be down to 6 days. These muddy deposits. Far more important for these
values need, of course, to be calculated indi- benthic animals is the interstitial salinity,
vidually for each estuary. Their calculation can which is the salinity of the water between the
be of great value in predicting the impact of mud particles. It has been consistently shown
polluting discharges on estuaries, as an estu- that the interstitial salinity varies much less
ary with a short flushing time is generally bet- than the salinity of the overlying water, due
ter mixed and better able to accept effluents. to the slow rate of interchange between them.
Within particular estuaries the pattern of On the intertidal mudflats, where the most
circulation may also vary throughout the year abundant populations of estuarine animals are
in response to meteorological events. The often found, the interstitial salinity matches
Potomac estuary (US) is a partially mixed estu- that of the high-tide salinity when it covers the
ary for 43% of the year, with surface outflow mudflats, which may be considerably higher
and deep inflow, but the reverse with surface than the salinity of the estuarine water at low
T H E E S T UA R I N E E N V I R O N M E N T 9
tide at the same location. Because of this estuary, these banks are the main source of
phenomenon it is usually possible for marine estuarine sediments. In the estuaries of the
animals living buried in the sediment to pene- Loire (France), Vigo (Spain), Apalachicola
trate further into estuaries, than for marine (US) and Yellow River (China), rivers carrying
animals living planktonically. large quantities of clay are the main source of
estuarine sediments.
Whatever the source of the sediments the
1.3 Estuarine sediments
deposition of it within the estuary is con-
Fine sedimentary deposits, or muds, are a trolled by the speed of the currents and the
most characteristic feature of estuaries, and particle size of the sediments. The relationship
indeed the estuarine ecosystem has been between current speed and the erosion, trans-
defined as a mixing region between sea and portation, and deposition of sediments is
inland water of such shape and depth that the shown in Fig. 1.8. It can be seen that for
net resident time of suspended (sedimentary) a sediment with pebbles l04 %m (1 cm) dia-
materials exceeds the flushing (Hedgpeth meter erosion of the sediment will take
1967). Sedimentary material is transported place at current speeds of over 150 cm s!1. At
into the estuary from rivers or the sea, or is current speeds between 150 and 90 cm s!1 the
washed in from the land surrounding the pebbles will be transported by the current,
estuary. In most North European and North but will be deposited at current speeds less
American estuaries the main source of sedi- than 90 cm s!1. Similarly for a fine sand of
mentary material is the sea, and the material is 102 %m (0.1 mm) diameter erosion will occur at
carried into the estuary either as suspended speeds greater than 30 cm s!1, and deposition
sediment flux or as bedload transported in the will occur at speeds less than 15 cm s!1. For
bottom inflowing currents that characterize silts and clays with particles of 110 %m dia-
salt wedges. In the Tay estuary, Scotland, UK, meter a similar relationship occurs, except that
for example, 70% of the sediments accumulat- consolidation of the sediment means that for
ing on the tidal flats are of marine origin. erosion to occur faster current speeds are
However, in some French estuaries, where needed as the consolidated sediment behaves
fine material is available on the banks of the as if it were composed of larger size particles.
1000
Mean velocity 15 cm above surface (cm s1)
50
Erosion
100
70
80 ti on
sp ort a
90 Tra n
10
Deposition
Figure 1.8 Erosion, transportation, and
deposition velocities for different grain particle
sizes. The diagram also illustrates the effect of
1 the water content of the sediment on the
1 10 102 103 104 degree of consolidation, which in turn modifies
Particle diameter (m) the erosion velocities. (After Postma 1967.)
10 T H E E S T UA R I N E E C O S YS T E M
The consequence of these relationships for to fall and settle within one tidal cycle.
estuaries is that in the fast-flowing rivers and Consequently the waters of estuaries tend to be
strong tidal currents at either end of the estu- very turbid as the silt and clay particles in sus-
ary all sizes of sedimentary particles may be pension are carried about the estuary until they
eroded and transported. As the currents start eventually settle to form the mudflats that are
to slacken within the estuary so the coarser so characteristic of estuaries. The speed of set-
pebbles and sands will be the first to be tlement can be slightly faster than portrayed in
deposited, and the finer silts and clays will Table l.3 as salt flocculation can occur. This
remain in suspension. In the calmer middle process is caused by the clay particles in salt-
and upper reaches of an estuary, where the water tending to adhere to each other, and as
river and tidal currents meet, and especially in they form larger particles so they will tend to
the slack water at high tide overlying the fall faster, for example, particles of 1.5 %m in
intertidal areas, only then will the currents be the deflocculated state form flocculated parti-
slow enough for mud to be deposited. cles, which behave as if they were 7 %m in
The rate of deposition, or the settling velo- diameter. Sea salts have two roles to play in the
city, of sediments is related to the particle aggregation of fine particles. First, even a small
diameter. For coarser particles (medium sand amount of salt is sufficient to make suspended
and larger) the settling velocity is determined particles cohesive, and second, the collisions
by the size of the particle and varies as the between particles prior to aggregation occur
square root of particle diameter. This relation- most frequently in the density gradients pro-
ship, known as the impact law, is represented duced by the mixing of salt and freshwaters in
by the equation: estuaries. The properties of sediments may also
be greatly altered by its inhabitants. In partic-
V $ 33 d
ular the sediments may become cohesive,
where V is the settling velocity (cm s!1) and which especially occurs when microscopic
d is the diameter of a spherical grain of quartz algae (microphytobenthos) inhabit the surface
(cm). For finer particles (silts, clays, and fine of the mud. The biological production of these
sands) the viscous resistance of the fluid in important microscopic plants is discussed later,
which the particle is settling determines the but here we should note the important role
settling velocity and Stokes law applies.
Particle shape, concentration, density, and effi-
ciency of dispersion all influence the precise Table 1.3 Settling velocities
values for Stokes law. Stokes law may be
Material Median diameter Settling velocity
represented for quartz particles in water at
(!m) (m day!1)
16 &C as:
Fine sand 250125 1040
V $ 8100d2
Very fine sand 12562 301
(V, d as above).
Silt 31.2 75.2
The results of the impact and Stokes law as Silt 15.6 18.8
settling velocity of various sizes of sediments Silt 7.8 4.7
are presented in Table 1.3, from which it can be Silt 3.9 1.2
Clay 1.95 0.3
seen that sands and coarser materials settle
Clay 0.98 0.074
rapidly in water, and any sediment coarser
Clay 0.49 0.018
than 15 %m will settle within one tidal cycle. Clay 0.25 0.004
However, for finer sediments, settling velocities Clay 0.12 0.001
are much slower and clay and silt particles less
than 4 %m diameter will certainly be unable Source: King (1975).
T H E E S T UA R I N E E N V I R O N M E N T 11
that they play in binding the fine particulate end of the estuary. The inward flow of marine
sediments of estuaries together. water and its sediment moves along the bot-
The middle and upper reaches of estuaries tom of the estuary until it reaches a stagnation
are thus characterized by very turbid water point where inward flow ceases. At this stag-
with poor light penetration. Within many nation point the seawater and its sediment rise
estuaries the suspended matter in the estuar- to mix with the surface fresher water. The
ies develop so-called turbidity maxima. The position of this stagnation point, and the con-
presence and magnitude of turbidity maxima sequent turbidity maximum, also varies with
are controlled by a number of factors, includ- the strength of river flow, moving upstream
ing the amount of suspended matter in the with low river flow, and downstream with
river or seawater, the estuarine circulation, high river flow.
and the settling velocity of the available mate- In many estuaries the maximum concentra-
rial. As Fig. 1.9 shows for the Ems estuary the tion of suspended sediment occurs at low
turbidity maxima is located at the meeting tide, as the ebbing tide washes sediment off
point of the river and tidal currents. The sedi- the intertidal areas and allows the sediment
mentary material in the central and upper in suspension to remain in the low water
reaches of the estuary is continually added channel. As the tide rises, the concentration of
to by river discharge and tide inflow, and suspended load is reduced as the flooding
remains largely in suspension, before finally tide increases the volume of water in the estu-
settling. The position of the turbidity max- ary, and the sediments are carried over the
imum is generally determined by the contribu- intertidal areas, where settlement may occur,
tion of suspended sediment from the seaward at high tide. In northern latitudes, ice break-up
Ems
Emden
Dollart
S Delfzijl
30 s.m P
mgl1 g- at l1
Suspended
150 15 matter
Salinity Total P
100 10 Figure 1.9 Turbidity maximum in the
15 Ems-Dollard estuary, on the border
between The Netherlands and the
Federal Republic of Germany. The
50 5 suspended matter is expressed as mg l!1,
salinity in practical salinity units, and
total phosphorus as %g-at l!1.
Observations were taken at the surface
of the estuary at the stations indicated
Stations (above- ) on the map. (After Postma 1967.)
12 T H E E S T UA R I N E E C O S YS T E M
in the spring is the most significant factor United States, in the Patuxent River estuary,
affecting muddy intertidal sedimentation. sediment collects at a rate of 3.7 cm year !1,
During summer, organic processes dominate and in the large estuarine system of Long
as organisms feed on, pelletize, and bioturbate Island Sound, near New York, it has been
the sediment, while plants may stabilize shown that since the estuary was formed 8000
the sediment. In autumn (fall) seasonal years ago the volume of marine mud supplied
storms increase, and from then into winter to the estuary greatly exceeds the volume
most sediment erosion and transportation will supplied by the rivers feeding the area.
occur. The changes in the particle size composition
Along with the sediments being carried into of sediments in estuaries noted above also
estuaries are usually carried particles of cause changes in other chemical and physical
organic debris derived from the excretion, properties of the sediment, which may influ-
death, and decay of plants and animals. Once ence the animals and plants living there. These
the dissolved and particulate organic matter changes are: to the water content and intersti-
reaches estuaries from fresh and saltwater it tial space, with higher water retention in well-
tends to remain there as it is deposited and sorted fine-grained sediments; to temperature
incorporated into the estuarine ecosystem and salinity where they change much more
along with fine inorganic matter. The organic slowly within the sediments than in overlying
matter in all aquatic ecosystems is convention- water or air; and to organic content and oxygen
ally divided into two fractions by filtration content, where fine-grained sediments are asso-
through filters with an average pore diameter ciated with greater organic content, and due to
of 0.5 %m. The fraction passing through the fil- the biological processes within them, with
ter is termed dissolved organic matter (DOM) lower oxygen content. The change from an aer-
even though it will contain very fine partic- ated surface sediment layer to deeper anoxic
ulate matter in addition to truly dissolved layers takes place closer to the surface in fine-
matter. The fraction retained by the filter is grained muds than in coarse-grained sands.
termed particulate organic matter (POM). This change, which may be approximated to
The sedimentation of both inorganic and by measuring the redox potential (Eh), of the
organic suspended material leads to the devel- sediment, serves to limit the macrofauna in
opment of mudflats and other areas of deposi- estuarine sediments to species that can form
tion within estuaries. Attempts have been burrows or have other mechanisms to obtain
made to summate the sources of suspended their oxygen from the overlying water.
material, and calculate the rate of deposition of
sediments. For example, within the upper 1.4 Other physico-chemical factors
Chesapeake Bay regions (US), the river sup-
plies 83% of the suspended matter (3/4 inor- It is tempting to regard estuarine water simply
ganic, 1/4 organic). In the middle reaches of as diluted seawater. However, there are dan-
the same estuary, shore erosion provides 52% gers in this approach, as evidenced by Fig. 1.10.
of the suspended matter, with production of It can be seen that the concentration of bicar-
organic matter within the estuary providing bonate ion drops only slightly from the sea to
another 22%. Within the whole Chesapeake freshwater. The concentration of other ions
Bay system, including the lower reaches, deviates markedly from the hypothetical dilu-
deposition of sediment occurs at a rate of tion line at low salinities (i.e. at less than l0),
0.8 mm year!1, with 56% of the sediment and certain ions such as phosphate, nitrate, or
supplied by rivers, 31% by shore erosion, and silicate may even be more abundant in river
12% from tidal inflow. Elsewhere in the water than seawater, and thus will show a
T H E E S T UA R I N E E N V I R O N M E N T 13
li
so
lu
1 v
s ol
al
T ot al
he
e
yp
o
C hl
low ionic strength, results in nonlinear chem-
0.1 ical perturbations that are amplified in the
0.1 1 10 100 low salinity waters of the FSI. The chemical
% seawater changes, which occur at the FSI contribute to
Figure 1.10 The effect of the dilution of seawater by river the food supply of zooplankton, as is discussed
water on the concentrations of bicarbonate and chloride in Chapter 3, and may also be responsible
ions, and the concentration of total dissolved solids. for changes in the composition of the estu-
Concentrations expressed as percentage of the arine fauna at low salinities, as discussed in
concentration in seawater. Note the logarithmic scales. Chapter 2.
(After Phillips 1972.)
The waters flowing into estuaries will con-
vey large quantities of oxygen into the estuary,
and additional oxygen will be supplied
decrease in concentration from rivers to the through the surface of the water and by plant
sea. Dissolved organic matter may also be photosynthesis. However, the many organ-
higher in river water than the sea, and the isms living within estuaries, especially in the
concentrations of most trace metals are similar bottom deposits, rapidly consume the oxygen
in river and seawater and therefore tend to be and thus many sediments are anoxic, except
relatively constant within estuarine water. for a thin surface layer. Where excessive
The distribution of trace metals, such as iron, organic enrichment occurs the multiplicity of
manganese, cobalt, nickel, copper, zinc, and microorganisms so produced may also con-
mainly the distribution and transport of sume all the oxygen within the water body as
suspended particulate matter, especially the well. The temperature regimes within estuar-
position of the turbidity maximum, control ies are often more variable than adjacent
cadmium in estuaries. waters due to the shallowness of estuaries,
Most of the biogeochemical processes which expose the waters to both heating and
involved in the meeting of freshwater and cooling, and also due to varied inputs of water
seawater in estuaries occur at very low salin- to the estuary, which may be at different tem-
ities ("1), where a region known as the peratures. A detailed account of estuarine
freshwaterseawater interface (FSI) has been chemistry is beyond this book and here we
identified as an important site for chemical shall restrict ourselves to the nutrient chem-
and biological reactions. The FSI has also been icals that may control the production of plant
termed the FBI (freshwaterbrackish water material, and thus serve as the foundation for
interface). It should be noted that changes in the estuarine ecosystem.
14 T H E E S T UA R I N E E C O S YS T E M
Estuaries are generally rich in the nutrients likely to be trapped within the estuaries.
needed for plant growth, especially nitrogen So, for example, 75% of the riverine input of
and phosphates, as supplies from rivers, the nitrogen to the North Sea derives from the
sea, and the land are continually replenishing Rhine and Elbe rivers, having passed through
the supply of these nutrients. Within the Ems their respective estuaries. The nitrogen balance
estuary, Netherlands Germany border, for of the Westerschelde in the Netherlands shows
example, it has been found that the freshwater little cycling within the system. Although den-
supplies about 92% of the nitrogen entering itrification or burial loses 25% of the nitrogen
the estuary and 71% of the phosphorus. entering the system, most nitrogen is oxidized
Despite processes within the estuary, mostly from ammonia to nitrate and then leaves the
burial and denitrification, 60 and 72%, respect- system to enter the North Sea. The nitrogen
ively, of the nitrogen are then transported model shows a net annual export to the North
from the estuary to the adjacent North Sea. Sea of 51,000 tonnes of nitrogen from the
The presence of large quantities of sus- Westerschelde, which is about 3% of the total
pended sediment in estuaries may cause sub- nitrogen input to the North Sea (1.5 ' 106
stantial modifications to the supply of nutrients tonnes N year!1), whereas the contribution of
for plant production. In the Tamar estuary, freshwater discharge from the Westerschelde is
England, the phosphate supply is the main lim- about ten times lower. Soetaert and Herman
itation on the growth of phytoplankton, but the (1995a,b) calculate the nitrogen load of
suspended sediments in the water exercise North Sea coastal waters off the mouth of
some form of buffering action such that they Westerschelde is increased due to tidal flows in
remove phosphate from the water when the and out of the estuary by about 14% per day.
concentration is high and release it when it is Because the estuary mineralizes nearly all
low. Thus as the phytoplankton assimilate allochthonous and in situ produced carbon, it
phosphate from the water, so there will be a is a large net nutrient exporter. Despite the
tendency for more phosphate to be released denitrification process that removes part of the
from the sediment. The turbidity of estuarine nitrogen, the amount of dissolved nitrogen
waters may also limit phytoplankton produc- that is flushed to sea is some 25% higher than
tion by restricting the penetration of light. the amount that enters from rivers or from
Studies by Heip et al. (1995) have shown that wastes.
it is the relationship between the benthic and Comparison of nutrient cycling and food
pelagic environment that critically controls the webs of six large Dutch estuaries shows that
entire functioning of the estuarine ecosystem all the estuaries experience increased nutrient
and they conclude that estuaries are effectively loading but, because of turbidity, increased
light limited systems where excess nutrient production of primary producers may not
input cannot lead to increase in primary pro- necessarily occur. The uptake of dissolved
ductivity. The overall recycling efficiency of inorganic nitrogen (ammonia and nitrate) in
nutrients in estuaries leads to three or fourfold several European estuaries (including the
re-use of any nutrients brought into they sys- North Sea estuaries of the Ems, Rhine, and
tem. Since the pelagic system is light limited by Scheldt) has been recently described by
turbidity, the role of the benthic flora is crucial Middelburg and Nieuwenhuize (2000), show-
notwithstanding this recycling of nutrients, ing that most nitrate entering or produced in
there is still a major outflow from estuaries into estuaries flushes through, and enters the sea,
the sea, and on balance as many nutrients may unless denitrification and/or burial in the
leave an estuary as enter it. sediment occurs. Ammonium and particu-
Nutrient inputs from rivers to the sea are for late nitrogen are efficiently recycled within
a variety of reasons, as discussed above, less the estuary.
T H E E S T UA R I N E E N V I R O N M E N T 15
The distribution of sediments and salinity For the biologist looking at the estuarine
within estuaries, along with the distribution of ecosystem certain characteristic zones emerge,
oxygen, temperature, and organic debris are each with typical sediments and salinity. In
all interdependent. The pattern of distribution particular estuaries one zone may occupy a
of these factors varies from estuary to estuary large proportion of the area, and the other
depending on topography and the volumes of zones may be compressed, but a clear sequence
water involved. Inherent in the nature of true can always be seen:
estuaries around the world are certain ecolog-
1. Head. Where freshwater enters the estuary,
ical common denominators, or factor com-
and river currents predominate. Tidal but very
plexes. They are:
limited salt penetration. FSI salinity "5.
Sediments become finer downstream.
(1) the presence of well-aerated, constantly
2. Upper reaches. Mixing of fresh and saltwater.
moving, relatively shallow water, which is
Minimal currents, especially at high tide, lead-
mostly free from excessive wave action or
ing to turbidity maxima. Mud deposition.
rapid currents;
Salinity 518.
(2) a salinity gradient from near zero up to
3. Middle reaches. Currents due to tides.
over 30, and accompanying chemical gradients;
Principally mud deposits, but sandier where
(3) a range of sedimentary particle sizes from
currents faster. Salinity 1825.
colloids to sands and detritus, resulting from
4. Lower reaches. Faster currents due to tides.
land weathering, water transport, and estuar-
Principally sand deposits, but muddier where
ine processes; and
currents weaken. Salinity 2530.
(4) complex molecular interactions in both
5. Mouth. Strong tidal currents. Clean sand or
water and sediments, taking place in an abun-
rocky shores. Salinity similar to adjacent sea
dance of dissolved and particulate organic
(#30).
matters, microorganisms, and fine sedimen-
tary particles. The relationship of these zones to the distribu-
tion of animals and plants will be discussed in
Romao (1996) has summarized the habitat of
Chapter 2.
an estuary as
The downstream part of a river valley, subject to 1.5 Brackish seas and coastal lagoons
tide and extending from the limit of brackish
waters. River estuaries are coastal inlets where, So far we have mainly examined the positive
unlike large shallow inlets and bays there is gen- estuaries, which are the commonest for
erally a substantial fresh water influence. The mix- those working in the estuarine environment.
ing of fresh water and seawater and the reduced Mention, however, needs to be made of the
current flows in the shelter of the estuary lead to range of brackish waters, whose inhabitants
the deposition of fine sediments, often forming
face similar problems to those encountered by
extensive intertidal sand and mudflats. Where the
tidal currents are faster than the flood tides mist
estuarine organisms. These areas, such as the
sediments deposit to form a delta at the mouth of Baltic Sea, Mediterranean Sea, Black Sea,
the estuary. Caspian Sea, and a variety of coastal and hyper-
saline lagoons cannot be classified as estuaries,
What this and similar descriptions have in since the definitions of Pritchard and Fairbridge
common is the recognition that the estuarine (Section 1.1) emphasize the importance of tides
environment is a complex and variable environ- as a distinguishing feature of estuaries and thus
ment characterized by salinity, tide, currents, explicitly exclude the Baltic Sea and other
and sediments, which is intimately linked as an brackish seas from being designated as estuar-
ecological unit with its surrounding habitats. ies. Indeed we have seen that the origin of the
16 T H E E S T UA R I N E E C O S YS T E M
word estuary is from aestus meaning tide. meaning than Estuary. All waters with salini-
Whether it is necessary for all estuaries to be ties between sea and freshwater can be called
tidal, or whether it is accepted that transitional brackish, whether they are large seas, closed
waters waters in all areas, irrespective of hav- lagoons, or tidal estuaries. An estuary is char-
ing tides can be called estuaries is still a matter acterized by tidal mixing and flushing as well
for debate. To avoid this problem legislators as a salinity gradient, whereas a brackish
unable to define estuaries are increasingly using water area is characterized principally on the
the term transitional waters. basis of salinity. Thus all estuaries are brack-
Waters with salinity lower than seawater, ish, but not all brackish waters are estuarine.
and higher than freshwater are called brackish The Baltic Sea is a major brackish water
waters (a word derived from the Dutch word sea area, but because of its lack of tides
brac, meaning salty). It should be made clear cannot classified as an estuarine. The Baltic
that the term Brackish waters has a wider Sea (Fig. 1.11) extends from Copenhagen in
4
5
lf
Finland
Gu
i
e
nk
inn
5.5 2 1
3
lsi
4
m
He
ar
Oslo 5
Tv
Sweden
Fin land
f
Stockholm lf o
Gu Tallinn
6
7
30
The 20
Sound Kalmar Riga
Denmark
15
Copenhagen
Bel
t Se 8
Kiel a
10
Gdansk
Germany Poland
Figure 1.11 Map of the Baltic Sea, showing average surface salinity. (Redrawn from Segerstrale.)
T H E E S T UA R I N E E N V I R O N M E N T 17
Denmark to Finland, and into the Gulf of the ecological master role of salinity also
Bothnia and the Gulf of Finland reaching emphasizes that much of the biological pat-
St Petersburg. Its maximum depth, off Gotland terns seen in estuaries are due to the muddy
is #200 m, and it is connected to the North Sea sediments and the widely varying nature of
via the shallow Oresund (The Sound) and the currents, tides, and associated physical and
Belt Sea. On the seaward side of the Sound the chemical factors.
salinity is high (#30). The salinity of the Sound Similarly the Mediterranean Sea is an
is variable depending on factors such as wind important sea area, which is almost closed
and water level. Saltwater entering the Baltic except for the Straits of Gibraltar, but can-
sinks owing to its greater density and becomes not be considered estuarine due to its full
virtually stagnant in the deeper areas of the salinity and lack of tides. Nevertheless some
Baltic. The fresher water from land drainage Mediterranean River mouths are commonly
and snow melt flows outward on the surface. called estuaries. The Black Sea was once part
The salinity pattern that develops is relatively of the great Tethys Sea, along with the Sea
stable with a salinity of 810 between Denmark of Azov, the Caspian Sea, and the Aral Sea
and Germany, around 6 between Stockholm (Fig. 1.12). This marine sea contained a typical
and Tallinn, and "2 in the innermost parts of Mediterranean fauna and during the Miocene
the Gulfs. Unlike estuaries this salinity gradient era (1520 million years ago) was split into
is relatively stable over hundreds of kilometers, distinct basins. During later years gradually
so that the organisms living in the Baltic expe- lowering of salinity occurred, the basins
rience a stable salinity regime coupled with a became established as the seas of today and
wide variety of substrates, ranging from rock to in time their salinity regime became stabi-
soft-sediment. As a consequence many animals lized. The salinity of the Black Sea is typically
are able to penetrate to lower salinities than 1719. The salinity is uniform and not in a
they do in estuaries, which while it confirms salinity gradient as in the Baltic. The fauna is
Sivash
Don
Vo
gal
Sea of Azov
Aral
Black Sea
Sea
Cas
Kara
pia
Gol
ea
Mediterranean 300 km
Sea
Figure 1.12 Map of the Black Sea, Sea of Azov, Caspian Sea, and Aral Sea.
18 T H E E S T UA R I N E E C O S YS T E M
Life in estuaries
2.1 Distribution of estuarine organisms while also providing nursery grounds for
marine fish and feeding areas for migratory
Estuaries are important to humans and marine
birds.
life. The tidal sheltered waters of estuaries sup-
The life of the plants and animals living in
port unique communities of plants and animals
estuaries will be examined in detail in later
that live at the margin of the sea. The estuarine
chapters, but first we need to consider the gen-
habitat experiences a steep gradient in salinity
eral features that are common to all of them.
conditions, coupled with a high degree of tur-
All the plants and animals in estuaries origin-
bidity, which leads to the deposition of muddy
ate from either the sea or freshwaters. They
intertidal areas. This habitat is rich in wildlife
either enter estuaries as part of a movement or
and for many people on this planet is the near-
migrate during their own lifetimes, or else
est thing to a natural habitat that they ever see.
their ancestors entered estuaries and succes-
It is a habitat that man has exploited and often
sive generations have become settled within
destroyed, but the estuarine habitat remains as
estuaries, so that it is now the only habitat for
one of the most resilient habitats on earth,
some species. The species living in estuaries
maintaining their attractiveness for wildlife,
fall into several categories (Box 2.1)
despite industrialization and land-claim. It is a
Many studies of the distribution and abun-
habitat that can provide unique ecosystem serv-
dance of animals and plants in estuaries have
ices to help the sea and mankind, for example,
shown that the number of species within
by trapping contaminants in its sediment,
Oligohaline organisms. The majority of animals minimum salinity that it can live in. Many disappear
living in rivers and other freshwaters do not by 18, but a few can survive at salinities down to 5.
tolerate salinities greater than 0.5, but some, the
Stenohaline marine organisms. These are marine
oligohaline species persist at salinities of up to 5.
organisms that only occur in the mouths of
True estuarine organisms. These are mostly estuaries, at salinities down to 25.
animals with marine affinities that live in the
Migrants. These animals, mostly fish and crabs,
central parts of estuaries. Most of them could live
spend only a part of their life in estuaries, with
in the sea but are apparently absent from the sea
some such as the flounder (Platichthys flesus) or
probably due to competition from other animals.
the shrimp (Pandalus montagui) residing and
Most commonly at salinities of 518.
feeding in estuaries. Others such as the salmon
Euryhaline marine organisms. These constitute the (Salmo salar) or the eel (Anguilla anguilla) are
majority of organisms living in estuaries with their purely migrants that use estuaries as routes to
distribution ranging from the sea into the central and from their breeding areas in rivers or the
parts of estuaries. Each species has its own open sea.
19
20 T H E E S T UA R I N E E C O S YS T E M
estuaries is less than the number of species categories of estuarine organisms a clear pat-
within either the sea or the freshwaters. Data tern emerges as shown, for example, in Tay
from a survey of the benthos of the Tay estu- estuary data (Fig. 2.1).
ary (Scotland), for example, shows a clear In the major groups of species living within
decrease in the number of species from the estuaries a clear generalized pattern of declin-
mouth of the estuary inward (Table 2.1). If we ing diversity can be seen as one enters the estu-
also consider the abundance of the various ary from either end. The number of oligohaline
species is high in true freshwaters, but declines
within estuaries. If the number of species
Table 2.1 Number of benthic species recorded from within freshwater is expressed as 100%, then by
transects in the Tay estuary a salinity of 5 the number of oligohaline
Transect Locality Number Substrate Salinity
species has fallen below 5%. The numbers of
number of species stenohaline marine species similarly falls as
one enters estuaries from the seaward end, so
I Buddon Ness 51 Sand 3233 that of the stenohaline species present in full
II 53 Sand strength seawater, virtually none are present
III 36 Sand 1132 below a salinity of 25. The numbers of species
IV Dundee 23 Mud/rock 630 of euryhaline marine organisms also declines
V 22 Sand 0.220.6
progressively from the sea into estuaries,
VI 21 Mud/sand
although in this case some survive down to a
VII 8 Mud 010.7
VIII 7 Mud salinity of 5. To offset this pattern of declining
IX Newburgh 7 Mud/sand 00.3 diversity of species, the numbers of species of
true estuarine animals in the sea or freshwaters
Source: Khayralla and Jones (1975). is low or nil, but increases within estuaries
120
North sea
100
Dundee
80
Number of species
Newburgh
ts
River Earn
n
la
60
ep
in
ar
M
40
100
Freshwater animals
50
%
Brackish-water animals
with a maximum number of estuarine species Table 2.2 Faunistic components in the Avon-Heathcote
in the 5 to 18 salinity region. The total number estuary (New Zealand) expressed as species number
of true estuarine species, as well as the number
Mouth Middle Head Total
of migratory species, is, however, relatively
low compared to the numbers of oligohaline or Salinity range 2534.3 8.430.3 0.215.7
marine species. Thus when the numbers of Freshwater 0 2 8 8
species of all groups living within estuaries is Estuarine 2 10 8 10
combined in a single diagram, as first drawn Euryhaline marine 69 67 10 72
by Remane and now known as Remanes Stenohaline marine 74 3 0 74
curve (Fig. 2.2), it can be seen that estuaries Total species 145 82 26 164
are characterized by having fewer species than Source : Knox and Kilner (1973).
adjacent aquatic environments.
Salinity determines the distance that a
Table 2.3 Number of benthic macroinvertebrate species in
species is capable of penetrating into the estu- the Potomac estuary
ary, but the full potential of any species to enter
the estuary can only express itself when suit- Salinity Number of benthic
able substrates are present. Data from the Avon invertebrate species
Heathcote estuary (New Zealand) (Table 2.2),
Zone
shows how small the estuarine and freshwater
Tidal fresh 00.5 70
component of the fauna is, with only 10 and 8 Oligohaline 0.55 104
species each, compared to 146 marine species, Mesohaline 518 184
of which 72 are stenohaline and 74 are Polyhaline 1830 199
euryhaline. Data from the Potomac estuary
(USA) is summarized in Table 2.3, where the Source : Lippson et al. (1981).
sequence of species within the estuary can also
clearly be seen. these two physical factors often makes it diffi-
In most estuaries there is a close connection cult for the biologist to distinguish their effect.
between salinity distribution and substrate The ecological factors of salinity and substrate
type, with reduced salinity associated with are closely interwoven in explaining the distri-
finer substrates. The close connection between bution of estuarine animals.
22 T H E E S T UA R I N E E C O S YS T E M
By contrast to the decline in the diversity of a productive environment for those species
species within estuaries, it must be emphas- that have entered it from the river, or from the
ized that the abundance of individual species sea. The estuarine habitat is thus characterized
often increases markedly within estuaries. by relatively few species, but these species
Thus a picture emerges of a greater number of may be very abundant.
individual animals, but of fewer species. If a
comparison is made between the numbers
2.2 Problems of life in estuaries
and weight of animals living in the bottom
deposits of marine beaches, freshwaters, and Estuaries are characterized by having abund-
estuaries at similar latitudes, it becomes clear ant populations of animals, but with relatively
that the estuary has the highest number and few species. What features within estuaries
weight of organisms. serve to restrict the diversity of species, and
In addition to the physico-chemical factors, explain the observed decrease within estuar-
discussed in Chapter 1, which control the dis- ies? The stability-time hypothesis compares
tribution and abundance of estuarine organ- the diversity of animals living within different
isms, Wildish identified three major biotic bottom deposits of the sea and estuaries.
factors that exercise some control over the In this approach the number of species pres-
numbers of animals inhabiting estuaries: ent within a sample is plotted against the
(1) food supply, (2) supply of colonizing number of individual animals present in the
larvae, and (3) interspecific competition. The same sample. When several environments
richness of the food supply, especially in the are compared (Fig. 2.3) a series of curves are
form of detritus, will be discussed later. produced. For estuarine samples the curve is
The supply of larvae within estuaries does always nearer the x-axis indicating fewer
present some problems, as planktonic larvae species in relation to the number of animals
may get carried out of the estuary by flushing present than occurs with marine samples,
currents. Many estuarine animals have thus which confirms the situation for estuaries of
suppressed the plankton larval stage, and only low diversity but high abundance. It is sug-
have bottom-living larval forms. Competition gested that estuarine environments are phys-
occurs in all ecosystems, but the relatively ically controlled communities, where physical
harsh estuarine environment, which chal- conditions fluctuate widely and are not rigidly
lenges the physiological mechanisms of all predictable. Thus organisms are exposed to
species (see Section 2.2) excludes many species, severe physiological stress and or the environ-
and thus the estuary represents for those ment is of recent past history. By contrast,
species, which can adapt to the environment, samples from the deep sea are described as
an escape from competition in the sea or fresh- biologically accommodated communities
water. Thus those species that can adapt to with a large number of species per unit num-
estuaries are able to utilize the rich food sup- ber of individuals, where physical conditions
ply available there, and it is this food supply, are constant and have been uniform for long
which becomes the main factor controlling the periods of time. From such data the stability-
biomass and productivity of estuarine animals. time hypothesis (as shown in Fig. 2.4) can be
The estuarine habitat is thus not a simple developed, with species number diminishing
overlapping of factors extended from the sea as physiological stress increases.
and from the land, but a unique set of its own Two main hypotheses thus emerge to
physical, chemical, and biological factors. The explain the paucity of estuarine species. The
estuarine habitat has provided the environ- first explanation is that of physiological stress,
ment for the evolution of some true estuarine and the second that the estuarine environment
organisms, but more importantly has provided is of recent past history and that there has
L I F E I N E S T UA R I E S 23
Gradient of
physiological stress
been insufficient time for a large number of physiological stress can be many and varied,
species to develop within estuaries. These two ranging from adaptations of behavioral pat-
hypotheses are not mutually exclusive and terns, and the modifications of particular organ
both could be operative. systems, through to the evolution of new races
The general term physiological stress or species adapted to the estuarine ecosystem.
covers a range of problems that confront the These physiological adaptations may take the
estuarine organism. The most conspicuous and form of the ability, or otherwise, of an animal to
probably best-studied physiological factor is tolerate and thrive in the estuarine environ-
salinity, since estuarine organisms are exposed ment, such as the tolerance of particular tem-
to variable and fluctuating salinities in contrast perature, salinity, or substrate conditions.
to the stable salinities, which characterize both The effects of salinity on estuarine organ-
marine and freshwater habitats. Other physio- isms are varied. The impact of salinity, as with
logical problems, which impinge on the life of other environmental factors, is often multi-
an estuarine organism, are the nature of variate. Thus, for example, temperature may
the muddy substrate, both in terms of the fine interact with salinity, and the response of an
particulate matter, which can clog delicate animal to a change in salinity may be different
organs, and of the virtually anoxic conditions at different temperatures. Salinity may affect
which occur within muds. The responses to an animal through changes in several chemical
24 T H E E S T UA R I N E E C O S YS T E M
properties of the water. First, the total osmotic of osmoregulation. For most stenohaline marine
concentration of seawater reduces as it is animals the former situation applies. When
diluted with freshwater. Second, the relative living in the sea their internal environment is
proportion of solutes within estuarine water isosmotic (osmotically similar) with seawater,
varies with the salinity, especially at very low thus although the ionic composition of the
salinities as described earlier. Third, the con- internal environment may be different from
centration of dissolved gases varies with the seawater, the total salt concentration is similar
salinity, with freshwater containing more to the seawater. As such an animal enters
oxygen than seawater at the same temper- an estuary the osmotic concentration of the
ature. Finally, the density and viscosity of internal environment falls to maintain equality
water varies according to salinity, with fresh- with the less saline estuarine water. This pas-
water lighter than saltwater. Thus when it is sive tolerance of osmotic change is, however,
reported that an animal responds to a change only possible down to a salinity of 1012
in salinity it is important to check whether the because below this salinity the cells and tissues
animal is responding to the change in salt con- of the animal may cease to function.
centration, or to any of the above factors As an alternative to the passive tolerance
instead. The effects of salinity on an animal of osmotic dilution, an animal may attempt
may also differ at different stages in the life to osmoregulate and maintain the internal
cycle. In general it appears that animals are environment at a different osmotic concen-
more sensitive to extremes of salinity at the tration from the outside. Thus, for example, an
egg stage, when newly hatched, or when in oligohaline, or true estuarine animal, which is
adult breeding condition, than they are at living in salinities below 10, must keep the
intermediate stages of growth. internal concentration at a level greater than
Salinity stress may invoke several responses. the outside. Such an animal, termed a hyper-
Confronted by an abnormal salinity an animal osmotic regulator, will strive to maintain its
may seek to escape or otherwise reduce con- blood concentration greater than a salinity of
tact with the water. Such behavioral responses 1012, even though the external environment is
are common in many estuarine animals. For lower, and thus enable the cells and tissues to
actively swimming animals, such as fish, it is function.
relatively easy to escape from an adverse salin- An animal, which hyperosmoregulates
ity, but sedentary animals such as barnacles when living in low salinities, is faced with two
can only respond by reducing contact and options when living at high salinities. Either it
attempt to seal themselves inside their shell. can become isosmotic with the seawater, or it
For a burrowing animal it is often possible to can seek to minimize the fluctuations of the
retreat into the burrow, or digs deeper, when external salinity regime by maintaining the
confronted by an abnormal salinity, but such a blood less concentrated, or hypo-osmotic,
solution is only useful if the abnormal salinity than the external salinity. In the latter case the
is of a short duration. If it is not possible to animal may thus live in salinities ranging
escape or reduce contact with the abnormal from near zero to over 35, while the blood con-
salinity, then the organism must rely on physio- centration may only rise from about 12 to 22
logical responses. Either it can allow the salinity. Such an animal, termed a hyper-
internal environment (blood, cells, etc.) to hypo-osmoregulator can thus achieve a much
become osmotically similar to the external more stable internal environment than is the
environment (sea or estuarine water); or else it case with a hyper-iso-osmoregulator, which
can attempt to maintain the internal environ- has still achieved greater stability than the
ment at an osmotic concentration different completely iso-osmoregulator. These patterns
from the external environment by the process of osmoregulation are presented in Fig. 2.5.
L I F E I N E S T UA R I E S 25
ype
H know. It has been suggested that the life his-
o-
e
lin
Baltic Sea and the Belt Sea, whereas H. ulvae temperate estuaries occurs in both unpolluted
dominates in the fluctuating salinities that and polluted estuaries alike, although the
typify many estuaries. Two species of cockle decrease will be more marked in polluted sit-
occur in European estuaries, Cerastoderma uations (Chapter 6). The size of estuaries as
($Cardium) edule and Cerastoderma glaucum a reason is rejected because the decrease in
($C. lamarcki). In Danish estuaries C. glaucum species occurs in both small coastal estuaries as
is the principal cockle at salinities below 18, well as large brackish seas, such as the Baltic.
while both species occur at salinities of 2025. Furthermore, small freshwater bodies have
At 3035, C. edule is the principal species, but greater species diversity than larger estuaries
at salinities of 60 in the Sea of Azov C. glaucum nearby. The turbidity of estuarine waters may
occurs. It thus appears that the brackish- inhibit marine species with delicate gill struc-
water cockle C. glaucum can tolerate a range tures, but the decline of species in brackish
of salinities wider than C. edule, but in normal waters occurs in both turbid estuaries, as well
seawater the edible cockle C. edule is the as estuaries with lower sediment loads, and
dominant species. non-turbid brackish seas. The critical salinity
The two main Corophium species in estuaries hypothesis of Khlebovich (1968) suggested that
appear to be separated by both substrate and for many purposes estuarine and brackish
salinity. Corophium volutator is more tolerant of waters with more than a salinity of 5 may be
low salinities, while Corophium arenarium is regarded as dilutions of seawater maintaining
more abundant at high salinities and with a constant ionic ratio, but that below a salinity of
sandier substrates. The importance of osmo- 5 many chemical properties of estuarine water
regulation as a means of limiting the penetra- change. Accordingly he suggested that these
tion of species into estuaries has been shown chemical peculiarities of estuarine water are
in the isopods of Jaera albifrons group. Jaera responsible for the observed eco-physiological
ischiosetosa can survive for several days in barrier at a salinity of 5 dividing the marine and
freshwater and is able to maintain a steady freshwater components of the estuarine fauna.
internal osmotic pressure even in very dilute This hypothesis is, however, invalidated by
seawater, and is the species, which penetrates many important estuarine animals, which live
furthest into estuaries. Jaera forsmanni and across the 5 salinity divide, by the fact that the
Jaera praehirsuta have lower survival rates in reduction in species occurs progressively all
dilute salinities and are less capable of main- the way from the mouth of an estuary, and
taining a steady state in dilute seawater, and by the fact that the major ionic perturbations
are found at the seaward end of estuaries. occur at very low salinities, the FSI at "1, at
J. albifrons is intermediate in both osmoregula- which salinity species diversity is increased
tory ability and in its ecological distribution. compared to 5.
These physiological differences coupled with If the above reasons are rejected as unsatis-
the observed ecological difference show how factory explanations for the reduction of
genetic isolation between the species can occur. species in brackish and estuarine waters,
In considering the possible causes for the what can be suggested? The main hypothesis
brackish water minimum of species at a salin- that remains is that of physiological stress.
ity of 5, the so-called Remanes minimum The salinity conditions of estuaries present,
shown in Fig. 2.2, Wolff (1973) and Deaton as we have already seen, many challenges to
and Greenberg (1986) reject pollution, size the physiological processes of the estuarine
of estuaries, turbidity, or the critical salinity habitants. The main challenge of living in a
hypothesis of Khlebovich, as satisfactory range of salinities rather than the constant
explanations. Pollution is rejected as an expla- salinities of the sea or freshwaters is met by
nation because the decrease in species within the range of adaptations both genetic and
L I F E I N E S T UA R I E S 27
behavioral that occur in brackish-water species. estuarine food web we need to study and quan-
It appears that few of the many animals living tify the life of each member of the food web.
in the sea have these osmoregulatory abilities, The simplest means of quantification is to
and the decline of species within estuaries identify, count, and weigh the organisms
can be seen as a progression of species each within particular parts of the estuary. The iden-
reaching their own physiological limit. tification of the organisms is of course the first
Studies of the fauna of brackish seas, such step, and can generally be performed with the
as the Baltic, where there is a progressive aid of guides to the seashore. The estimation of
decrease in salinity, reveal an accompanying the number of organisms depends on choosing
decrease in species diversity just as seen in a unit area, usually 1 m2, and counting the ani-
typical estuaries. The rate of decrease of mals and plants within the area. For small
species is, however, generally less marked in organisms it is usually impracticable to count
brackish seas than in estuaries. This difference everything in 1 m2, so one would count those
in the rate of decrease may be explained by the within a smaller area such as 10 cm2 and mul-
stability of the salinities in each area. In the tiply the result by an appropriate factor to pro-
Baltic it is found that more species are able to duce a result in terms of 1 m2. Such counts
live at lower salinities in the stable low salin- need to be repeated several times and the
ities of the Baltic than they are able to in the mean result determined. If the animals are
salinity regimes of estuaries, which vary with buried in the sediment, as will be the case in
each tide as well as seasonally. It thus appears many estuarine species, then the sediment
that the fluctuations of salinity within estuaries must be sieved through an appropriate sieve to
enhances the rate of decrease of species, but extract the animals. A 1-mm sieve is suitable
since a decrease occurs in both stable as well as for bivalve molluscs, but for most other estuar-
fluctuating situations instability cannot be an ine animals a 0.25 or 0.5 mm ($250500 %m)
exclusive explanation. A further factor that sieve is needed to be certain that a true estim-
permits survival of fauna at lower salinities in ation of abundance is obtained.
brackish seas than in estuaries is that low salin- The typical results of an exercise in counting
ity areas in estuaries are invariably associated estuarine animals is shown in Fig. 2.6, from
with high turbidity and fine sediments. which a clear pyramid of numbers can be seen,
Whereas in brackish seas the waters are less with many small animals, decreasing to fewer
turbid due to their lack of tides and there may larger animals. If, however, the animals are
often be a wider variety of habitats ranging weighed the reverse picture can be seen,
from rocky shores through to soft-sediments. Fig. 2.7, with the total weight of the smallest ani-
mals considerably less than the total weight of
the larger animals, the pyramid of weight. The
2.3 The estuarine food web
name given to the weight of the organisms is
The estuarine food web is dependent on the the biomass. The measurement of biomass alone,
input of energy from sunlight and the trans- important though it may be in comparing
portation of organic matter into the estuary the immediately available standing crop from
by rivers and tides. Within the estuary the one part of an estuary to another part, is quite
plants, or primary producers, convert these inadequate for the purpose of estimating the
inputs into living biological material. As amount of food, which one trophic level, such
the plants grow they are consumed by the herb- as the plants, can make available to the animals
ivores, or primary consumers, which are in turn feeding on them. In order to understand
consumed by the carnivores, or secondary con- these feeding, or trophic, relationships we
sumers. In order to understand the conversion need to know the rate of production of organic
of material as it passes through the links of the matter by the various members of the estuarine
28 T H E E S T UA R I N E E C O S YS T E M
Weight
Component Abbreviation Value
(mg)
Mean annual B 203 kcal m!2
< 10,000
biomass
< 1000 95 Change in biomass (B 40 kcal m!2 year!1
< 100 48 Mortality E 20 kcal m!2 year!1
Production P 124 kca1 m!2 year!1
< 10 88
Assimilation A 600 kcal m!2 year!1
<1 15 Gamete production G 64 kcal m!2 year!1
< 0.1 10 Respiration R 476 kcal m!2 year!1
Egesta F 388 kcal m!2 year!1
< 0.01
Consumption (A ) F ) C 988 kcal m!2 year!1
50 100 Production/biomass P : B ratio 0.61
Wet weight (g m2) ratio
Assimilation efficiency A ' l00/C 61%
Figure 2.7 The pyramid of weight. The weight of the fauna
Net growth efficiency P ' l00/A 21%
(m!2) of Niva Bay, Denmark, divided into seven weight
categories. Compare with Fig. 2.6. (After Muus 1974.) Ecological efficiency E ' 100/C 2%
ecosystem. The distinction between the rate of The production of a population may be
production (or productivity) of organic matter derived by one of two approaches. Either all
by an organism or community, and the biomass the growth increments of all the members of
(or standing crop) consisting of the organism the population under study are added for
itself or the community, is fundamental. 1 year, or else growth is ignored or production
A summary of the terms and units used in is derived from the addition of the change in
productivity studies is given in Box 2.2. The the biomass over 1 year plus the mortality due
derivation of various units of biomass and to predation etc. over the same period. An
its conversion into energy units is given in energy budget for an intertidal population of
Box 2.3. A summary of approximate conver- the bivalve Scrobicularia plana may be taken as
sions from one unit to another is given in an example of the utilization of energy by an
Box 2.4 and examples are given in Table 2.4. estuarine invertebrate (Table 2.5). Note that
L I F E I N E S T UA R I E S 29
Box 2.2 Terms used in productivity studies, which are utilized in this book
Note: Assuming the conservation of energy we may derive the following equations:
Consumption C $ P ) R ) G ) U ) F $ (total energy budget)
Absorption Ab $ C ! F $ P ) R ) G ) U
Assimilation A $ C ! (U ) F) $ P ) R ) G
Derived from the above is the P : B ratio, which is the ratio of the production of an organism to the mean
biomass of that organism over 1 year.
Source: Crisp (1984).
79% of the energy assimilated was utilized in important measure of the transfer of energy
respiration, which is a typical value for ener- within an ecosystem than is the biomass. The
getics studies, and only 2% of the energy P : B of estuarine animals from temperate areas
consumed by Scrobicularia was consumed by may vary from less than 1, to well over 5.
the next trophic level. The significance of this ratio is that it is a
The relationship between production and clear indication of the ecological performance
mean annual biomass is known as the of a population. Two populations may, for
turnover ratio, or P : B ratio. Comparison of example, have similar biomasses, but differ-
various temperate benthic invertebrates ent P : B ratios, thus the one with the high
shows that the P : B ratio varies only between ratio will produce more organic material to
2.5 and 5, with a mode of about 2.5. This value be made available to predators than the one
shows that production over a study year is 2.5 with the lower ratio. In comparing differ-
times greater than the mean biomass, and ent animals at one site, smaller animals will
emphasizes that production is a much more have higher P : B ratios than large animals.
30 T H E E S T UA R I N E E C O S YS T E M
Box 2.3 Units used in productivity studies, which are utilized in this book (After Crisp 1984)
Biomasses expressed as gram carbon or gram nitrogen, like gram ash-free dry weight, are intended
as measures of the living tissue present in a sample, and provide a useful alternative to the determination
of the energy content of the material.
The above biomasses (14) can also be expressed in kilogram or milligram. However, the values derived take
no account of the energetic value of the tissue, which may vary from species to species.
Reasonable estimates of annual production many productivity studies, the P : B ratio has
and its distribution among size groups in ben- been extensively investigated over the past
thic communities can be derived from knowl- 30 years. Our understanding has passed from
edge of biomass and the size structure of the a realization that body size and P : B are linked
community (Fig. 2.8). As a consequence of to one where the relationship between the two
L I F E I N E S T UA R I E S 31
Box 2.4 Conversion of units to and from each other. Conversions of units of biomass into
units of energy vary from species to species, and from season to season, however, certain
approximations can be used as indicated below (Data from various sources)
1 gC approx. $ 10 kcal
1 gC approx. $ 2 g ash-free dry wt
1 g ash-free wt approx. $ 21 kJ
1 g organic C approx. $ 42 kJ
1 litre oxygen $ 4.825 kcal (oxycalorific equivalent)
1 g carbohydrate approx. $ 4.1 kcal
1 g protein approx. $ 5.65 kcal
1 g fat approx. $ 9.45 kcal
1 cal $ 4.184 J
Some selected examples:
1 g flesh dry wt C. edule $ 18.5719.46 kJ
1 g ash-free dry wt C. edule $ 19.9921.50 kJ
1 g flesh dry wt C. volutator $ 13.2616.82 kJ
1 g ash-free wt C. volutator $ 18.6620.88 kJ
1 g flesh dry wt M. edulis $ 19.9520.83 kJ
1 g ash-free dry wt M. edulis $ 21.7122.51 kJ
1 g flesh dry wt M. balthica $ 16.1019.04 kJ
1 g ash-free dry wt M. balthica $ 19.3821.46 kJ
1 g flesh dry wt N. diversicolor $ 16.4419.70 kJ
1 g ash-free dry wt N. diversicolor $ 20.8722.42 kJ
Thus 1 g flesh dry wt estuarine invertebrate approx. $ 18 kJ
Thus 1 g ash-free dry wt estuarine invertebrate approx. $ 21 kJ
102
Bacteria
101
Annual P : B
Meiofauna
Macrofauna
10
Figure 2.8 Relations between annual P : B
(production : biomass ratio) and mean individual body
mass (as kilocalorie equivalents) for benthic organisms.
101
Geometrical mean functional regressions are shown
1010 108 106 104 102 10 for bacteria, meiofauna, and macrofauna. (After
Mean individual body mass (kcal equivalents) Schwinghamer et al. 1986.)
is routinely accepted for productivity calcula- discrepancies when used in tropical situa-
tions. The most commonly used relation- tions. Similar relationships can be shown
ships are shown in Table 2.6. While they work between the annual respiration (R) and annual
well in temperate situations, there are some productivity (P) (Table 2.7).
32 T H E E S T UA R I N E E C O S YS T E M
Table 2.6 Productivity as a function of biomass (B), body Table 2.7 Productivity (P ) as a function of respiration (R )
size (M, kcal equivalents) and temperature (T, &C)
Group Units Relationship
Group Relationship
Poikilotherms kcal P $ 0.64*R 0.85
Bacteria P : B $ 0.696*M !0.208
Macrofauna kcal Log10R $ 0.367 )
Meiofauna P : B $ 0.73*M!0.337 0.993*log10P
Macrofauna P : B $ 0.525*M!0.304 Estuarine fauna gC m!2 year!1 Log10R $ 0.081 )
Benthic invertebrates P : B $ 0.65*M!0.37 1.02*log10P
Benthic invertebrates Log10P $ !0.473 ) Estuarine Birds kcal Log10P $ 0.79*log10R !
1.007*log10B ! 0.274*log10M 1.055
Benthic invertebrates P $ 0.0049*B 0.80T 0.89
Source: Wilson (2002), and references therein.
Source: Wilson (2002), and references therein.
Salt marsh
Producers
&
Detritus algae Phytoplankton
Zooplankton
Detritus Plant Eurytemora
10 Consumers
Invertebrate
predator
Nephtys
20 Consumers
Carcinus
Birds Fish
Waders Flounder
Ducks Goby Figure 2.9 An estuarine food web. The
arrows indicate the feeding relationships of a
Death typical northern hemisphere estuary. The
dotted lines indicate the division into
producers, primary consumers, and secondary
Microbial decomposers consumers used in this book.
Organisms within the estuarine ecosystem within a spacetime unit of any magnitude,
may be assigned to trophic (or feeding) levels and thus includes the biotic community of
following the trophicdynamic approach to organisms plus its abiotic environment. Within
ecology. The ecosystem may be defined for- the biotic community we can assign organ-
mally as the system composed of physical isms to trophic levels, which are the groups of
chemical and biological processes active organisms, which share a common method of
L I F E I N E S T UA R I E S 33
obtaining their energy supply. The first trophic some consumer animals, for example, feeding
level is the producers, which are the plants on prey from different trophic levels. In the suc-
obtaining their energy by photosynthesis. ceeding chapters of this book we shall examine
Following them are the primary consumers, the estuarine ecosystem as a series of trophic
which are herbivorous animals feeding on levels, but the reader should be cautioned that
plant material. Following these are the it is not always easy to assign a particular
secondary consumers, which are carnivorous organism to a particular level. In Fig. 2.9 an
animals feeding on the primary consumers. attempt is made to show some of the links
Eventually we reach the tertiary consumers within a typical north European estuary. Such a
feeding on the secondary consumers. At each food web is derived from an understanding of
stage in this trophic sequence energy is con- the biology and feeding habits of the various
sumed, and while some of it is rejected as members of the estuarine ecosystem. The
waste, or converted into bodily growth, the studies of ecosystem processes and the factors
majority of the energy is dissipated as heat fol- controlling energy flow, either at the species or
lowing respiration. The loss of energy due to the community level are the basic building
respiration is progressively greater for higher blocks of our understanding of the estuarine
levels in the trophic sequence, as is the effi- ecosystem. Initially many studies have con-
ciency of utilization of the food supply. Due to centrated on single species, measuring their
the losses of energy in food chains there are consumption, growth, and the production of
rarely more than five trophic levels. Apart energy. In time such studies can be combined
from the loss of energy from each trophic level in order to build up food webs or trophic
due to respiration and predation, other mate- diagrams, and eventually we can measure the
rials such as excreta or dead organisms pass to trophic structure, cycling, and the ecosystem
the decomposer trophic level where micro- properties of estuaries.
organisms break down the material until it too Throughout this book we shall attempt to
is lost as heat or else reutilized as detritus by explain both the energy sources, which support a
the detritivores, which are regarded as mem- particular organism, as well as the contribution
bers of the primary consumer level. that organisms make to the functioning of the
This model of trophic transfer is essentially a estuarine ecosystem of which it is a member.
food chain with one level linking directly to the We shall consider the ecology of each trophic
next. In practice, however, when examining level in turn, before considering how the
ecosystems we are looking at a food web with systems operate together.
CHAPTER 3
34
P R I M A RY P R O D U C E R S 35
generally called particulate organic matter Estuaries are heterotrophic systems, in which
(POM), without regard to its exact origin. consumption exceeds production, and it is the
In this chapter, therefore, we shall examine overall organic loading that controls primary
both the primary production of the salt and secondary production (Heip et al. 1995).
marshes and algae (benthic or planktonic), In terms of loading, estuaries receive large
and the limitations placed upon this pro- quantities of allochthonous inputs, that is
ductivity by nutrient availability. Also the organic matter generated outside the system
fate of the plant material as it is frag- and transported into the estuary where it is
mented and decomposed, and thereby becomes then available for heterotrophic consumption.
more available to consumer animals. The term There are three principal sources of allochtho-
primary production is generally viewed as nous input, namely tidal import from the sea,
the assimilation of inorganic carbon and nutri- riverine sources and sewage and waste dis-
ents into organic matter by autotrophs. posal. Tidal imports vary with the size of the
Hence primary production is a rate. Although tide and can be linked directly to the volume of
this definition also includes production water exchanged on each tide, but these have to
by chemoautotrophs, this is not normally be balanced against the corresponding export
measured, because most primary production of material on the ebb. It is this balance, which
measurements on phytoplankton (and other determines whether or not tidal movements
aquatic plants) are made with the 14C method, function as a net import (flood dominated) or
and with this method the dark-bottle meas- export (ebb dominated) of material.
urements are usually subtracted from the The concept of ebb or flood dominated tidal
light-bottle values to obtain a true photosyn- flows and interest in the transport of materials
thesis rate. Community ecologists generally in estuaries resulted in the Outwelling
use the term gross primary production as Hypothesis of Odum (1968). This stated that
organic carbon production by the reduction of marshestuarine ecosystems produce more
CO2 as a consequence of photosynthesis. Net organic material than can be utilized or stored
primary production is then defined as gross within the system and that the excess material is
primary production minus autotrophic respi- exported to the coastal ocean where it supports
ration. Within the literature there may, how- near coastal ocean productivity. Since this was
ever, be confusion regarding these terms due, formulated several investigations of material
in large part, to the wide variety of techniques transport have been carried out. It has been
in use and the reader is referred to Under- found in general that in European systems there
wood and Kromkamp (1999) for fuller details is usually a net import of particulate materials
of definitions and techniques used to measure while in North American estuaries, export of
primary production in estuaries. material is normally observed (Table 3.2).
Table 3.2 A summary of material (particulate and dissolved) transport in estuarine systems
Table 3.3 Production, respiration, net ecosystem production, burial, and exchange of American salt marsh systems
(gC m!2 year!1)
Note: The mass balance export requires extrapolation to the entire salt marsh, and direct flux is based on samples taken directly
from tidal creeks.
Source: Heip et al. 1995 and references therein.
P R I M A RY P R O D U C E R S 37
first studies to present a complete energy flow Spartina production was also assimilated
for any ecosystem, and he showed that the salt directly by herbivorous insects.
marsh under study received 600,000 kcal m!2 Nitrogen is a key nutrient in the productivity
year!1 of sunlight, of which 8295 kcal m!2 of coastal ecosystems, and salt marshes that
year!1 became net primary production within receive increased amounts of nitrogen show
the salt marsh. Twenty percent of this net increased rates of primary production. The
primary production was due to benthic algae, nitrogen budget of Spartina-dominated salt
with 80% of the net primary production due marshes on the Atlantic coast of the United
to Spartina grass. The algae were utilized by States has been investigated in detail, where it
consumer animals directly, but most of the has been shown that increased nitrogen supply
Spartina became detritus and was subject to not only increases the productivity of the
decomposition by bacteria, with much of the plants, but also leads to increased biomass in
Spartina production dissipated as bacterial the detritus feeding invertebrates dependent on
respiration (Fig. 3.1). A small amount of the the salt marsh (Table 3.4). Over a 2-year period
Sunlight
600,000
305 81 28
Insects Spiders
Spartina
8205 5
& algae 224 53 23
gross 1 prod
36,380 7900 48 3671 Net export
4010 35
Detritus
28,175 462 65
Nematodes
Bacteria
& crabs
3890 397
Figure 3.1 Energy-flow diagram for a Georgia salt marsh (units are kcal m!2 year!l). (After Teal 1962.)
Table 3.4 Nitrogen budget for Great Sippewissett Marsh. Units are kilogram per year
for the entire 0.48 km2 marsh
groundwater flow from small underground detritus was exported in 5 h from a 0.36 km2
springs, rainfall, tidal exchange, and the salt marsh. Not all American estuaries receive
amounts of ammonium, nitrate, nitrite, dis- such large quantities of detritus from Spartina.
solved organic nitrogen, and particulate nitro- In the salt marshes of the Patuxent river
gen within each were measured, as well as the estuary less than 1% of the Spartina produc-
fixation of nitrogen by both free-living bacteria, tion reaches the estuary as detritus. This is in
bacteria associated with the roots of marsh great contrast to the 2045% reported for other
plants, and by algae. Measurements were estuaries, which is attributed to greater
made of the loss of nitrogen from the system degrees of tidal flooding elsewhere. Dissolved
due to sedimentation, denitrification, and har- organic carbon (DOC) is also released from the
vesting of shellfish. Finally, account was made leaves of S. alterniflora into the estuarine water,
of the input of nitrogen from bird faeces. A which rhythmically inundates the salt
remarkably good agreement between the marshes. It has been calculated that the DOC
measured input and the output of nitrogen is released from Spartina is 61 kgC ha!1 year!1.
seen in Table 3.4. The 11% difference is small Although this represents only a few percent of
considering the many possible sources of error the total production, the DOC can be readily
in the calculation, and in general it seems that metabolized by the microbial populations in
this ecosystem is in balance. Most of the the water and thus becomes available to con-
nitrogen budget is controlled by the physical sumer animals.
factors of the tide, supplemented especially by Within British and other north European
groundwater flow. Considerable changes do, estuaries the salt marshes are typically found
however, take place within the salt marsh. For only in the region above the point of the low-
example, 64% of the nitrate, which enters the est neap high tide. In this region they are not
marsh, is intercepted, and ultimately leaves covered by the tide every day, but are covered
the marsh in the form of particulate ammo- periodically by spring high tides. Salt marshes
nium and nitrogen. An amount of particulate thus occur in the upper intertidal area and the
organic matter equivalent to about 40% of plants that occur there must be able to tolerate
the net annual above-ground production of being covered occasionally by saline estuarine
the marsh is exported from this marsh, pro- water. Salt marshes display a clear zonation,
viding a rich food supply for the detritus or successional sequence, from low to high
feeders. The highly productive salt marsh elevations. The plant most typical of the outer,
studied has achieved a balanced steady state, or shore, end of the salt marsh is Salicornia
which supports the estuarine ecosystem (glasswort or marsh samphire). The classical
mainly as a source of particulate organic sequence is then Glyceria maritima, Suaeda
matter, and as a means of converting and recy- maritima, or Aster tripolium, above these are
cling nitrogen. Limonium vulgare (sea lavender), then Armeria
Relatively little of the Spartina is consumed maritima (sea pink), followed by Atriplex
directly by animals, and instead the net prim- species, and Festuca rubra and Juncus maritimus
ary production of Spartina mostly reaches the toward the top of the salt marsh.
estuarine ecosystem in the form of fragments The salt marsh habitat is recognized as a key
broken off the grass. These fragments form the component of the estuarine ecosystem, and is
basis for detritus, as bacteria progressively often specifically protected under legislation.
decompose them. By trapping the detritus in There are, for example, 643 km2 of salt marsh
the tidal creeks of salt marshes, it has been on European North Sea coasts, over half of
found that periodic storms are responsible for which (55%) is on the Wadden Sea coasts,
the export of large quantities of detritus from while 26% is on Britains east coast and 7% is
salt marshes. After one storm over 2000 kg of in the Dutch Delta region. The ecosystem role
P R I M A RY P R O D U C E R S 39
of salt marshes is often identified as a habitat cold currents. Tropical estuaries grade into
for animals such as juvenile shrimps or fish, subtropical systems beyond the tropics of
and as a roosting and breeding site for birds. Cancer and Capricorn where a winter water
For example about 20,000 pairs of Redshank temperature low of about 12 &C marks their
(Tringa totanus), being about 60% of the total southern and northern limits. Although many
British breeding population, breed on salt estuaries of the south and south east United
marshes. It is because of their importance to States have been described as tropical or
the functioning of the major estuarine systems subtropical, this is not really the case in a
and their survival of the large populations world context, with the possible exception of
of winter feeding birds (see Chapter 5) that south Florida mangrove-lined systems. Winter
a very high proportion of salt marshes are water temperatures in many US Gulf of
protected in Europe. This is not to denigrate Mexico estuaries fall as low as 5 &C.
their other interests, but their significance as a Like salt marshes, the main contribution of
roost site for birds is a much more obvious mangroves to the estuarine ecosystem is
manifestation of their conservation value. through the abundant supply of plant litter,
which is either used directly or in various
degraded forms by many animals (Fig. 3.2).
3.3 Mangroves
Many of the worlds great estuaries are in
3.4 Intertidal plants and macroalgae
the tropics. The Amazon, Orinoco, Congo,
Zambezi, Niger, Ganges, and Mekong are all Sea grasses are true flowering plants, and
very large and receive drainage from enormous several sea grass species inhabit estuaries,
catchments. Tropical estuarine environments including Thallasia, Posidonia, and Cymodocea
range in size from tiny seasonally flowing sys- in warm and tropical waters, and Zostera,
tems of 12 km2 to the estuaries of some of the Ruppia, Potamogeton, and Zannichellia in tem-
worlds largest rivers. They also encompass perate areas. Sea grasses, like other estuarine
extensive coastal lakes and the reduced salinity organisms, show greatest species diversity at
estuarine waters extending along the coast in the seaward and freshwater ends of estuaries,
parts of southeast Asia, South America, and and reduced diversity within the central parts
Africa. In all these tropical and subtropical of an estuary. The eel-grass or widgeon grass,
estuaries mangrove trees occupy a similar habi- Zostera spp. is the commonest sea grass on the
tat to the salt marshes of temperature estuaries, intertidal estuarine flats in many temperate
fringing the banks of the estuaries with dense estuaries growing on sandy and muddy sub-
stands of vegetation and forming the dominant strata, and occurring subtidally down to 1-m
intertidal vegetation. depth (Fig. 3.3). The annual net production is
Mangroves generally match the 20 &C about twice the maximum biomass and ranges
isotherms in both hemispheres, suggesting from 58 to 330 gC m!2 year!1, and exception-
that water temperature is the most significant ally up to 1500 gC m!2 year!1.
influence. It has been shown that the presence In warmer waters, such as in Florida or
of mangroves correlates with areas where the Puerto Rico, Thalassia becomes the main sea
water temperature of the warmest month grass, with biomasses of 208100 g dry wt m!2
exceeds 24 &C; also that their northern and and productivity values of 100825 gC m!2
southern limits correlate reasonably well with year!1. These high productivity values will
the 16 &C isotherm for the air temperature of often be supplemented by 2030% epiphytic
the coldest month. Latitudinal ranges are plants, that is smaller plants growing attached
greater on eastern continental margins than on to the Thalassia. As for other plants, the energy
western sides due to the presence of warm or may be utilized by animals, not so much by
40 T H E E S T UA R I N E E C O S YS T E M
Mangrove tree
Crabs
Protista Molluscs
Detritus (POM)
Beach with wrack masses Upper slope Lower slope (Zostera meadows) Mud bottom
Figure 3.3 A typical eel-grass (Zostera) community of Danish fjords and land-locked brackish waters, in an area not subject
to destruction. (From Rasmussen 1973.)
grazing the sea grass, but rather through the waters, and coastal lagoons with favorable
detritus route. The distribution of submerged light conditions are the primary sites for the
vascular plants is determined principally by development of submerged sea grasses.
the presence of shallow (sandy) sediments The production of seaweeds (macroalgae)
and the turbidity of the water. Shallow, quiet such as Fucus and Ascophyllum can be high
P R I M A RY P R O D U C E R S 41
on marine rocky shores. In estuaries, however, summer season, and decline in the autumn.
populations of seaweeds tend to cover a very At the end of the growing season large popu-
small proportion of the total area, being con- lations of heterotrophic bacteria, and sub-
fined to rocky outcrops, quays, and piers. The sequently denitrifying bacteria, develop on
seaweed Fucus ceranoides is confined to estu- the rotting algae. The Enteromorpha can be the
aries, in contrast to other Fucus species that main means of accumulating nitrogen from
tend to occur only on fully marine coasts. In a the waters that flow into the estuary, and as the
comparison of the estuarine F. ceranoides with algal mats decay the nitrogen is made available
the marine Fucus vesiculosus it has been found to other parts of the ecosystem. The mats
that the distribution of the species are limited may also smother the animals living within the
by salinity, with low salinity unfavorable for mudflats, and as the mats decay they may
F. vesiculosus and high salinity unfavorable utilize much of the available oxygen, to the
for F. ceranoides. The seaweeds of Florida estu- detriment of the animals.
aries, which are tolerant to a wide range of Macroalgae can have high rates of primary
temperature, light, and salinity and even short production within the areas where they occur
exposures of freshwater, can continue to photo- (Table 3.5) and on an estuary-wide basis can con-
synthesise while both covered and exposed by tribute up to 27% of total primary production.
the tide. Highest biomass and production values seem
The mudflats of estuaries, which receive to occur when hydrodynamic energy is rela-
high nutrient (especially nitrogen) inputs from tively low, such as in lagoons and tidal inlets.
inland areas, for example, the Eden, the Ythan, In open river dominated estuaries, especially
or Chichester harbour in the United Kingdom, in funnel shaped estuaries, biomass seems to
may become covered in profuse growths of the be lower, although wherever hard substrates
green alga Enteromorpha (mainly Enteromorpha occur in the intertidal, locally high densities of
prolifera), which develop as mats during the macroalgae can be found.
mgC m2 day1 C
25
600
Primary
400 production
15
Mudflat Figure 3.4 Average primary production of
temp. microphytobenthos on a tidal flat in the
200 Wadden Sea, near the island of Texel, The
5 Netherlands. Production is expressed as
mgC m!2 day!1, and the temperature of the
0 upper sediment layer of the same mudflat is
J F M A M J J A S O N D also shown. (After Cadee.)
P R I M A RY P R O D U C E R S 43
Table 3.6 Microphytobenthic primary production from intertidal sediments in different estuaries
Sources: Underwood and Kromkamp (1999), Heip et al. (1995), and MacIntyre et al. (1996).
energy) decreases benthic algal production, is relatively low, which might be due to two
and for this reason chlorophyll biomass and factors, shallowness, and turbidity. The shal-
hence primary production of microphytoben- low nature of the estuaries studied, which is
thos seems to be positively correlated to the typical of most estuaries, meant that the mean
clay content of the sediment. That is, fine sed- depth of the water, at 1.18 m, was 1.7 m less
iments indicate quiet wave conditions that than the optimum depth for producing max-
favor the microphytobenthos. imum net photosynthesis. The penetration of
light in estuarine waters is severely limited by
the turbidity of the water, due to suspended
3.6 Phytoplankton
sediments and POM, which will again limit
The phytoplankton is an integral part of the the production of the phytoplankton. Within
estuarine ecosystem; however, it does not the Lower Hudson estuary, USA, dissolved
have such a dominant role as, for example, inorganic nutrients are high throughout the
the phytoplankton in marine ecosystems or year, but large blooms of phytoplankton do
freshwater lakes. Despite the abundance of not occur despite this availability of nutrients.
nutrients in estuaries, other factors may limit It is suggested that this is due to the flushing
the production of estuarine phytoplankton. rate of the estuary, whereby the populations of
The photosynthesis and respiration of phyto- phytoplankton are carried out to sea before
plankton has been measured in a 400-km2 sys- their growth rates permit the development of
tem of estuaries near Beaufort, North Carolina. phytoplankton blooms. Those peaks of phyto-
The net production was 52.5 gC m!2 year!1. plankton that do occur are related to the
While the daily rate of production of phyto- incursion of marine water carrying plankton
plankton could be quite high, the annual rate into the estuary.
44 T H E E S T UA R I N E E C O S YS T E M
Research workers studying different estuar- than the flushing rate of the estuary. Table 3.7
ies have come to widely different conclusions summarizes the results of measurements of
regarding the role of phytoplankton, some phytoplankton primary production from
claiming that primary production of phyto- several estuaries worldwide. The productivity
plankton is insignificant, while others regard of estuarine phytoplankton may be lower than
phytoplankton production as being of central production values for coastal phytoplankton,
importance to the estuarine ecosystem, res- and much lower than values for salt marsh
ponsible for approximately 85% of total production, but the high productivity in rela-
ecosystem primary production. Although tion to the biomass is pronounced, and the
nutrients appear to be available for the pro- phytoplankton is for many animals a richer
duction of large quantities of phytoplankton, source of food than plant fragments prior to
maximal production is apparently rarely decomposition. The ciliate protozoa, with
achieved due to three factors. First, turbidity their rapid turnover time (2 days) may often
can limit the penetration of light, second, the be the main consumers of phytoplankton in
shallowness of many estuaries means that brackish waters, consuming more than the
blooms may not develop, and third, the heavier, but slower-growing, copepods of the
growth rate of the phytoplankton may be less zooplankton.
Table 3.7 Phytoplanktonic primary production in different estuaries. Where different areas within
estuaries have been studied, values are given from central areas
United States
San Francisco 150 2.23.3 1070
Bay (South Bay)
San Francisco 9398 1.114.6 "0.55 2050
Bay (Suisun)
Tomales Bay 360810 511 39 1583
Neuse River 360531 2.53.4 40 #50
Chesapeake Bay 337782 4.010.5 15
Hudson River 70220 1822 1.2 3055
Bash Harbor 11 2.6 12 11
Marsh
Delaware 296 40
Peconic bay 162191 2.93.2 2.14.6 14
Europe
Oosterschelde 242406 3.06.6 48.5 1636
Westerschelde 184197 8.511.1 1.23.7 2633
Bristol Channel 49 210
Ems-Dollard 26154 4.69.9 1.64.5 1345
Marsdiep 250390 615 56.5 3842
(Wadden Sea)
Asia
Pearl River 20 35
There is a large interplay of variables influ- concentrations and ratios may influence
encing the rate of phytoplankton photosynthe- species composition of phytoplankton, which
sis (nutrient or light limitation, osmotic stress) might have profound ecological implications.
and factors influencing biomass such as graz- Primary production generally increases
ing, washout, resuspension, and deposition. toward the mouth of an estuary, indicating
Phytoplankton in estuaries may experience that the decreases in nutrients are more than
rapid changes in the type of limitation compensated for by the increased water trans-
(nutrients, light) and different physical parency. This fact alone demonstrates that
environments (mixing, salinity) and these primary production is not determined solely
changes may influence species composition. by nutrient input and availability. Table 3.8
Interannual variability in primary production shows some recently published values of
can to a large extent be explained by changing annual production in different regions of
watershed conditions and changing land use, several estuaries, and Table 3.9 shows phyto-
as the watershed and rainfall determine the plankton biomass. These tables show a trend
nutrient and sediment input into estuaries for increased annual production toward the
from the land. These inputs can both stimulate outer part of the estuary, but that maximal
primary production when the system is biomass may occur in inner areas. Light
nutrient limited or when the light conditions and nutrient availability mainly influence
improve, or it can decrease primary produc- phytoplankton growth. Freshwater input can
tion as turbid sediment-laden water can have negative effects by cresting osmotic
decrease the light availability or flush out the stress, by flushing out populations and by
populations. There is a wealth of evidence increasing turbidity, but can have positive
that, due to increased land use and the associ- effects as a major source of nutrients and by
ated nutrient load, many estuaries have creating a stratified water column, which can
undergone eutrophication. The effect may to improve light conditions and thus initiate
a large extent be dampened out when grazing phytoplankton blooms. In view of these con-
by suspension feeders is important. In some flicting forces, single environmental variables
cases eutrophication can lead to harmful algal are poor predictors of phytoplankton primary
blooms in the phytoplankton. Even in this production in estuaries. Primary produc-
case, total primary production will not neces- tivity of estuarine phytoplankton is mainly
sarily change, but the changes in nutrient controlled by three variables: Phytoplankton
Table 3.8 Estimates of annual production of phytoplankton (gC m!2) in several estuaries,
arranged as freshwater, inner, central, or outer
biomass (i.e. chlorophyll concentration), inci- In the context of an estuarine energy budget,
dent irradiance, and turbidity according to detritus is referred to as allochthonous sources
Heip et al. (1995). Nutrient concentration, of particulate organic carbon, to distinguish
grazing, transport, sedimentation, tempera- it from autochthonous material, being that
ture, and daylengtth seem to be of minor produced by the primary producers through
importance. Others caution against over- photosynthesis. Allochthonous sources can be
emphasizing the role of turbidity and under- divided into (a) riverine, (b) marine, (c) atmos-
estimating the effects of nutrients and suggest pheric, and (d) erosion inputs, as well as
that while turbidity probably controls pro- (e) direct domestic and industrial inputs. The
ductivity in macrotidal systems or in river- relative importance of the various sources
dominated reaches, nearly all estuaries depends on factors such as river discharge,
experience some degree of nutrient limitation tidal amplitude, estuarine morphology, land
near their seaward boundaries. usage, and human population as well as the
geology of the area. Inevitably the proportions
of the different sources will vary from estuary
3.7 Detritus
to estuary. In the context of allochthonous
Detritus has already been defined as all types organic matter it is important not only to rec-
of biogenic material in various stages of ognize the source and amount of organic mat-
microbial decomposition. Much of this bio- ter but also its quality. Labile allochthonous
genic material may be fragments of plants, organic matter refers to material, which can be
such as Spartina. In estuaries without large salt readily degraded and hence made available to
marshes the main sources of detritus are frag- consumers, whereas refractory matter is obstin-
ments of dead plants and animals from the ate and resistant to degradation and hence
sea, from rivers, or from the estuary itself, as may be unavailable to consumers.
well as the faeces, and other remains of the Spartina and other plant detritus is rela-
estuarine animals. All the types of primary tively indigestible to the consumer animals
production described in the preceding sec- and thus much of the flux of organic matter to
tions of this chapter can supply material that detritivores must involve the conversion of
becomes detritus, and it is clear from many the particulate detritus to soluble compounds
studies that most primary production in estu- and their assimilation by microorganisms,
aries is not consumed directly by herbivores, which can then be consumed by detritivores.
but rather is converted into detritus before The nutritive value of Spartina increases as
consumption by detritivores. the detrital fragments become enriched with
P R I M A RY P R O D U C E R S 47
microbial populations. In Fig. 3.5 it can be become smaller the protein content increases
seen that living Spartina has a content of 10% to 24%. Thus the detritus, which is rich in pro-
protein. Dead leaves entering the water have tein, may be a better food source for animals
about 6% protein, but as the plant fragments than the grass tissue that formed the basis for
the particulate matter. Similar results have
been described for leaves of the tropical estu-
Spartina Detritus arine salt-marsh plant, red mangrove, with
6.1% protein in leaves on the tree, 3.1% protein
at leaf fall, and 22% protein after decomposi-
tion in estuarine water for 12 months. The
main decomposers of plant material in seas
60 and estuaries are bacteria, as shown in Fig. 3.6.
The detritus, composed of the decaying
50 Carbohydrate remains of plant primary production, and
microbes, has a valuable role in stabilizing the
% Ash-free dry wt.
100 1.5
% Original carbon
(%)
(%)
100
No. g1 1010
7.0
% Original nitrogen Bacteria
Figure 3.6 Changes in percentage
(%)
50 3.5
original carbon, original nitrogen, and total
nitrogen concentration, and numbers of
bacteria, during the decomposition of
0 80 160 0 80 160 submerged leaves of the Mangrove
Days Days Avicennia marina. (After Robertson 1988.)
48 T H E E S T UA R I N E E C O S YS T E M
provide a larger surface area for microorgan- activities of these suspension-feeding bivalves
isms, and so accelerate the processes of decay. may be profound. In South San Francisco Bay
The activities of animals that consume the estuary, for example, the suspension-feeding
entire sediment, lead to a continual mixing of bivalves filter a volume equivalent to the total
the organic and inorganic particles in the sedi- volume of the area each day, and this grazing
ment, in a process known as bioturbation. This may be the primary mechanism controlling
will tend to distribute detrital material phytoplankton biomass. In Chesapeake Bay,
throughout the surface layers of the sediment, USA, the detritus averages 77% of the total
and so enable material, which has settled on the organic particles in the water column, and the
surface of the sediment to organically enrich phytoplankton 23%. The numbers of detritus
the sediment to a depth of several centimeters. particles in the water showed little seasonal
It has been calculated that the biomass of bac- variation, whereas the phytoplankton in this
teria within estuarine sediments may be of the area showed considerable seasonal variation.
same order of magnitude as the biomass of Estimates of the global organic inputs into
the animals in the sediment. Apart from the estuaries indicate that the major sources are
immediate surface layer, estuarine sediments primary production from both wetlands (salt
tend to be anaerobic, as the bacteria and other marshes) and planktonic and intertidal algae,
microorganisms consume all the available along with organic matter carried into the
oxygen. Much detritus therefore undergoes estuary from rivers. To these values must
anaerobic metabolism, with hydrogen sulfide, be added mans discharges of sewage, oil
methane, or ammonia produced, as well as products, food products, and wood pulp and
dissolved organic carbon compounds that can a quantity of organic matter entering the estu-
be utilized by aerobic microorganisms living on ary from the sea. All these sources of organic
the surface. These aerobic microorganisms may matter are utilized by microorganisms within
also be consumed by detritivores. the estuaries of the world, to become detritus.
It should not be assumed that all detritus is Along with the POM that forms detritus in
the same for detritivore animals. Studies on estuaries, there may be considerable quantities
the utilization of detritus by the polychaete of dissolved organic matter present, derived
Capitella capitata have shown that detritus from plant exudation, animal excretion, and
derived from salt marsh and sea grass plants from the products of decomposition. The
that contain a high percentage of unavailable capacity for the uptake of dissolved organic
energy is consumed as the products of micro- matter by animals is widespread, but despite
bial decomposition and protein enrichment, this it seems likely that estuarine animals get
whereas detritus derived from seaweeds may the vast majority of their food from POM. The
be consumed directly. dissolved organic matter will mainly be
The primary consumers that ingest POM metabolized by bacteria, and some estimates
from the water column do so without regard show that bacterial production utilizing dis-
to whether it is phytoplankton, suspended solved amino acids can amount to 10% of
detritus, or microbial organisms. One study algal production. The bacteria, consuming the
attempted to segregate these components, and dissolved organic matter, themselves become
came to the conclusion that the bay scallop, part of the particulate matter in the estuary.
Argopecten irradians, ingests 20% phytoplank- In parallel with the decomposition of organic
ton and 80% detritus and bacteria, emphasizing matter there is liberation of carbon dioxide,
the much greater availability of detritus within phosphorus, nitrogen, and other nutrients. This
the estuarine ecosystem, even though the recycling of nutrients, referred to as mineraliza-
growth rate of the scallops would have been tion, is a prerequisite for the new production of
higher on a diet of phytoplankton alone. The organic matter by autotrophs.
P R I M A RY P R O D U C E R S 49
Table 3.10 Net primary production of particulate material in various estuaries, expressed as percentage of total
production (gC m!2 year!1)
of detritus, and it may be the supply of detritus in the water and sediments (25.4 ' 106 kgC
derived from the breakdown of the primary year!1).
producers, which is the feature of most impor- The study of the Dollard estuary clearly
tance to the success of the primary consumers. shows that primary production within an estu-
Measurements have been made in the ary is inadequate to support the large number
Dollard estuary on the DutchGerman border of detritus feeders inhabiting the mudflats,
in an attempt to quantify all sources of organic and the detritus feeders must rely on the
input to this estuary. Eighty percent of the importation of organic debris from outside the
estuary is composed of tidal sand mudflats. estuary. What primary production does take
The total amounts of organic carbon entering place is due overwhelmingly to the benthic
and leaving the Dollard estuary are shown algae, rather than the phytoplankton whose
in Table 3.11. This preliminary attempt at a production is inhibited by the turbidity of the
carbon budget has clearly revealed a large dis- water. Thus the basic biological processes cre-
crepancy between the measured inputs and ating energy for the primary consumers in this
outputs, which is believed to be mainly due to estuary are concentrated on the mud surface
unquantified export of dissolved carbon from with the primary production of the benthic
the estuary. Nevertheless, several valuable algae, and the transformation of organic debris
points emerge from this study. First, the main into more digestible material by bacteria.
sources (75%) of carbon are outside the estuary The high productivity of the Dollard, and
in the river, the sea, and an industrial plant many similar estuaries, is thus seen to be due
(potato flour mill), which discharges effluent. to the position of the estuary as a collecting
Second, due to the turbidity of the water the area for organic matter, supplemented by the
primary production from phytoplankton is primary production of the benthic algae on the
only 7.5% of the primary production from intertidal flats. The high productivity of this
benthic algae such as diatoms and blue-green estuarine ecosystem is thus because it is subsi-
algae. Furthermore, the total primary produc- dized by the transfer of energy from other
tion of 10 ' 106 kgC year!1 is considerably ecosystems.
less than the carbon consumed, or utilized The Grevelingen estuary, in the Netherlands,
was studied intensively prior to the implemen-
Table 3.11 Organic carbon budget for the Dollard estuary tation of the Delta Barrage scheme, which is
(units are ' l06 kgC year!1 for the entire area of almost described in Chapter 6. This estuary covered
100 km2) 140 km2, of which 81 km2 was covered at all
times, 55 km2 was intertidal sand and mud-
Import/ Export/utilization flats, and 4 km2 was salt marsh. A detailed
production of organic carbon
food budget for the period before it was
Particulate C from 37.1 Dissolved C to 7 dammed is shown in Table 3.12. The levels of
North Sea ) North Sea primary production are dominated by the pro-
River Ems duction of phytoplankton, supplemented by
From potato 33.0 Utilization in water 7.2 benthic microalgae. This is the reverse of the
flour mill situation in the Dollard estuary, and is due to
From salt marshes 0.5 Utilization in 18.2 the less turbid waters in the Grevelingen, cou-
sediment pled with the smaller proportion of intertidal
Primary production 0.7 Buried in sediment 9.9
area. The total contributions from all sources
Phytoplankton
within the estuary to the carbon budget is,
Primary production 9.3 Bird feeding 0.26
Microphytobenthos however, exceeded by the material carried in
Total import 80.6 Total export 35.56 on each tide from the adjacent North Sea.
The carbon budget for Barataria Bay,
Source: van Es (1977). Louisiana, USA shown in Table 3.13 reveals
P R I M A RY P R O D U C E R S 51
Source: Wolff (1977). Figure 3.7 The Forth estuary, eastern Scotland, UK.
A typical European-type estuary with large intertidal mudflat
areas, bare of macrophyte vegetation. Microphytobenthos is
a main primary producer in such habitats, in the foreground,
Table 3.13 Carbon budget for Barataria Bay, Louisiana, the plant Salicornia can be seen colonizing the uppermost
(gCm!2 year!1) assuming that 1 gC is equal to 2 g dry areas of the mudflat.
organic wt
Source Sink
In the European-type estuary much of the within the estuary, excess material remains,
primary production within the estuary is per- which is carried out of the estuary to fertilize
formed by large populations of microscopic the adjacent sea.
benthic algae living on the surface of the mud For both types of estuary, and those inter-
supported by phytoplankton in the water col- mediate between the two extremes, we can
umn. The extent of the primary production of conclude that the high levels of production
the phytoplankton depends on the turbidity of within estuaries are due to a plentiful supply
the water. However, in these estuaries the of nutrients supporting the primary produc-
majority of the energy within the primary pro- tion of benthic algae, phytoplankton, and salt
ducer trophic level is derived from outside the marshes. This production is enhanced by the
confines of the estuary, and is in the form of import of POM into the estuary from either
organic matter, which is carried into the estuary, the sea or the margins of the estuary, which
usually from the sea, but also from land dis- undergoes microbial decomposition within
charges of river water or sewage. The estuary is the estuary to yield a rich food supply for the
thus a net recipient of energy, and the high pro- consumer animals.
ductivity that supports large populations of In all estuaries the gradients in concentra-
consumer animals is due to the position of the tions of nutrients and turbidity are steep and
estuaries as traps for both nutrients and POM. ecosystem studies have often emphasized the
In the American-type estuary the primary light limitations on pelagic systems imposed
production of benthic algae and phytoplank- by estuarine turbidity and that benthic prim-
ton is important for the productivity of the ary production can therefore be relatively very
whole ecosystem, but the dominating factor is important. Nevertheless, the estuaries are large
the much greater proportion of the estuary, net exporters of excess nutrients. Estuaries are
which is inhabited by rich beds of Spartina also large net importers of carbon (mainly as
grass. The Spartina is only consumed directly detritus). The primary transfers of energy
by animals to a small extent, and instead they within estuarine ecosystems therefore derive
rely on the fragments of Spartina forming the from organic detritus inputs plus microphyto-
substrate for large populations of bacteria, benthos to support benthic communities that
which form detritus, which is then ingested by in turn support the birds, fish, and shrimps, as
the animals. Despite high rates of consumption will be seen in the next chapters.
CHAPTER 4
53
54 T H E E S T UA R I N E E C O S YS T E M
Mollusca
C. edule Suspension 2605565 200300
M. balthica Suspension 120727 78500
and (Spat 2000)
A B Deposit
S. plana Mainly 1213 0
suspension
Figure 4.1 M. balthica The left individual is seen
feeding on suspended matter with the inhalent siphon Mya arenaria Suspension 217 3040
stretched up in the water. The right individual is seen H. ulvae Grazing 106014,000 400153,000
sucking in bottom material with the inhalent siphon. For and
both, the exhalent siphon rejects waste material. deposit
(From Olsen, in Rasmussen 1973.) Annelida
L. conchilega Suspension 3571
Nephtys sp. Carnivore 5877 "160
within a specific size range (e.g. #2 %m for Hediste Carnivore 235425 36750
Mytilus) and it is unable to distinguish ($Nereis) sp. and
between phytoplankton and floating detritus Suspension
Crustacea
of similar size. Among deposit-feeders, the
C. volutator Deposit 173368 151985
typical example may be the lugworm, Arenicola
marina, which ingests large quantities of mud, Compiled from various sources.
and having digested off any organic material,
rejects the bulk of the mud in the familiar
worm cast. Between these clear-cut examples
are many animals that can both suspension- because the food supply for suspension-feeders,
feed and deposit-feed. The Baltic tellin, which is phytoplankton and suspended POM,
Macoma balthica lies buried in the mud with its varies seasonally during the year, or even from
siphons protruding (Fig. 4.1). When food is day to day. Whereas the food supply for
available in the water overlying the mud, it is deposit-feeders, which is benthic algae or
able to raise the incurrent siphon and draw in detritus and the microbes growing on it, will
water and particulate matter. When detritus vary less throughout the year. Furthermore
particles are available on the surface of the the biomass, but not necessarily the abun-
mud surrounding its buried location, it can dance of suspension-feeders shows a gradation
reach out across the mud and collect the parti- within the estuary, with a greater biomass of
cles by using its siphon as a vacuum cleaner. suspension-feeders near the seaward end of the
Within the animal the same gills provide a estuary, in order to feed on material brought in
size-limited filtration for the food particles, so by the tides (Fig. 4.2).
that Macoma, for example, can spend 1040% As an alternative to the classification of the
of its life suspension feeding and 6090% benthos into suspension-feeders and deposit-
deposit-feeding. feeders, the system of epifauna (or surface
The numbers (Table 4.1) of suspension- fauna) and infauna (or buried fauna) may be
feeding benthic animals appear to fluctuate suggested. Typical epifauna are animals such
more, than those of deposit-feeders do. This is as mussels (Mytilus) on a mussel bed, barnacles
P R I M A RY C O N S U M E R S 55
7000
10
6000
5000 8
4000 6
3000
4
2000
2
1000
0 0
1 2 3 4 1 2 3 4
Salinity region Salinity region
Relative dominance of different feeding guilds Relative dominance of different feeding guilds
100 100
90 90
80 80
Abundance (%)
70 70
60 Biomass (%) 60
50 50
40 40
30 30
20 20
10 10
0 0
1 2 3 4 1 2 3 4
Salinity region Salinity region
Suspension-feeder Surface deposit-feeder Deposit-feeder Omnivore
Figure 4.2 Absolute and relative dominance (abundance and biomass) of the different feeding guilds in the intertidal
zone of each salinity region of the Oosterschelde estuary. Region 1 is nearest the mouth, and Region 4 is nearest the head
of the estuary. (After Ysebaert et al., 2003.)
4.2 The mud dwellersbenthic as abundant as 3500 m!2, which is the highest
deposit feeders density recorded from North America. At
Minas Basin, the abundance of Macoma has
4.2.1 Molluscs
been compared with environmental factors
The bivalve M. balthica (Fig. 4.1) is one of the such as sediment grain size, tidal elevation,
most widespread of estuarine benthic animals. organic carbon and nitrogen, and bacterial
It can be a rather slow-growing animal, and in density. The sediment grain size was found to
some situations such as the St Lawrence in set limits for the animal, with none being
Canada may take 12 years to grow to 14 mm. found in sands coarser than 0.23 mm, or finer
In Massachusetts or the Wadden Sea it grows than 0.024 mm. Within these limits it was
faster, reaching 26 mm in 6 years. It is pro- found that the density of Macoma could be
posed that growth rate and longevity are a accurately predicted by bacterial density alone.
function of the hydroclimate; with warmer Macoma cannot survive by deposit-feeding on
temperatures a larger size and shorter life bacteria alone, and to survive in the large den-
span are observed. Typical productivities of sities reported at Minas Basin it must supple-
Macoma and other molluscs are shown in ment its diet by suspension-feeding.
Table 4.2 Macoma balthica in the Wadden Sea feeds as a
The Minas Basin, Bay of Fundy, Canada has deposit-feeding animal for up to 90% of the
tides of up to 17-m range. On the intertidal time that it is feeding. While deposit-feeding it
mudflats exposed by the tide, M. balthica is the is utilizing mainly benthic algae, which grow on
dominant organism, along with the amphipod mudflats. Macoma growth starts in the spring as
Corophium volutator, and adult Macoma can be soon as the production of microphytobenthos
Bivalves
M. balthica 5.03 Ythan, Scotland Mudflat
1.95 Bay of Fundy, Canada Mudflat
M. edulis 134 Ythan, Scotland At mussel bed
24.3 Ythan, Scotland Whole estuary
1875 Morecambe bay, England At Mussel bed
87.3 Dublin Bay, Ireland At Mussel bed
55.5 Wadden Sea, Netherlands Mussel bed
M. arenaria 5.8 Petpeswick, Canada Mudflat
Modiolus demissus 5.75 Georgia, USA Salt marsh
C. edule 40.3 Wash, England Sand flat
(in patches)
31.0 Baie de Somme, France Muddy sands
Crassostrea virginica 171.2 South Carolina, USA Oyster reef
S. plana 12.4 Bangor, Wales Mudflat
Gastropods
Hydrobia ulvae 6.4 Grevelingen, Netherlands Mudflat
Littorina saxatilis 0.75 Petpeswick Inlet, Canada Spartina marsh
reaches 100 mgC m!2 day!1 and continues Hydrobia ulvae (Fig. 4.4) is an important
throughout the spring and early summer as pri- feeder on detritus and algae on the surface of
mary production increases. The annual growing mudflats, although it retreats into the surface
season starts earliest in years that follow a layers of the mud when the tide recedes.
mild winter. As the temperature reaches 10 &C Hydrobia is a selective deposit-feeder that
spawning occurs, and growth ceases at temper- feeds mainly by grazing on diatoms growing
ature above 16 &C. Thus growth is restricted to a on the surface of particles of 20250 %m dia-
range of water temperatures between 4 and meters. Alternatively it can secrete mucus,
16 &C in spring. The variation in the Macoma which traps bacteria, and then it reingests the
population in the Wadden Sea over an 8-year enriched mucus. Eutrophication by sewage
period has been shown to be related to the level effluent has led to the development of exten-
of the food supply coupled with the time of sive algal mats over of the former open
immersion by water. Other studies agree that mudflats in several estuaries. The presence of
Macoma behaves most of the time as a deposit- the mat reduces the biomass and diversity of
feeder, but have shown that it depends for its the infauna, but leads to a great increase in the
food intake for the greater part on food present numbers and biomass of Hydrobia. In one
in the water column. The explanation for this study at Langstone harbor the biomass of
apparent contradiction is that, as shown in Hydrobia increased from 5.4 g m!2 to 27.4 g m!2
Fig. 4.3, much of the total food intake is rejected with abundances increasing from 9000 m!2 to
as being unsuitable, and is lost as feces (or 42,000 m!2 due to the algal mats, which pro-
pseudofeces). A greater proportion of the food vide a rich food supply for the Hydrobia. The
taken in by suspension-feeding, rather than hatching of embryonic snails of H. ulvae can
deposit-feeding, is regarded as suitable and occur at salinities from 8 to 60 and at temper-
hence it is this food, which forms the main basis atures up to 35 &C. The newly hatched snails
of their diet. attach to the surface of the adults shells. The
wide range of possible salinity and temper-
ature combinations for hatching coupled with
Total intake (258) the non-planktonic development and the vari-
able modes of feeding of the adults are all
Unsuitable
important reasons for the success of this species
as food 106 in estuaries.
186 Faeces
The survival strategies for estuarine mol-
luscs are, thus, a tolerance of low salinities, a
tendency toward non-pelagic larvae, and the
80
Suitable
as food
51 Respiration 1
72
Production
(mm)
21
3
adoption of a flexible mode of feeding Table 4.3 Productivity (gC m!2 year!1) of estuarine
designed to cope with the problems of a annelids
muddy environment. Many estuarine mol-
Species Productivity Location Habitat
luscs can utilize a number of different food
(gC m!2
sources, either at different times in their life year!1)
history, or at different phases of the tide. This
flexibility of feeding strategies is the hallmark N. ($Hediste) 27.2 Belgium Brackish
of a successful estuarine mollusc. diversicolor pool
N. diversicolor 5.45 Ythan estuary, Mudflat
($Hediste) Scotland
4.2.2 Annelids A. marina 3.13 Grevelingen, Fine
Netherlands sand flat
The polychaete annelid worms, commonly
Nephtys 3.68 Lynher estuary, Mudflat
known as bristle worms, are the most diverse hombergii England
group of worms in an estuary. Four Nereid Ampharete 1.16 Lynher estuary, Mudflat
polychaete species (Nereis ($Hediste) diversi- acutifrons England
color, Nereis ($Neanthes) virens, Nereis
($Neanthes) succinea, Nereis pelagica) com- Source: Wilson (2002) and McLusky (1989).
monly penetrate into estuaries, and show dif-
ferences in their physiological tolerances, and
in the pattern of their life cycles. The ragworm
(N. diversicolor) is able to tolerate the lowest Waste
salinities, appears to have only a brief ("6 h)
larval planktonic phase to assist in retaining the Sand
populations within estuaries, and is usually the
commonest worm in European estuaries. In
America, the clamworm (N. succinea) is the
most likely species of polychaete to be encoun-
tered. Typical productivities of Nereis and
other annelids are shown in Table 4.3 from
which it is clear that the polychaetes with their
shorter life spans and faster growth than, for
example, most molluscs are a key component
in the productivity of estuaries.
Figure 4.5 A. marina The position of the lugworm
The biomass of the lugworm, A. marina Arenicola in its burrow. Filled arrows (from waste end)
(Fig. 4.5), is apparently controlled by the indicate water currents, and open arrows (at head end)
amount of organic matter in the sediment, indicate the ingestion of sand. (After Kruger 1971.)
although it is not found in sediments finer
than a median particle diameter of 80 %m, as it
cannot maintain its burrow in such fine sedi- The sand-mason (Lanice conchilega) occurs on
ments. In a closely related species Abarenicola the sandier parts of many estuaries, where it
pacfica it has been found that a gardening forms burrows up to 30 cm deep but with a
strategy is adopted whereby the undigested small protrusion of about 2 cm standing clear of
sediment, which is passed out as feces is pop- the sand surface. For much of the time it is a sur-
ulated by microbial organisms, which increase face deposit-feeder, but it can spend up to 35%
the nitrogen content of the sediment by their of its time feeding on particles in suspension in
activity. The lugworm then reingests the the water. This combination of two modes
enriched feces. of feeding enables it to utilize alternative food
P R I M A RY C O N S U M E R S 59
algae, fungal spores, and plant fragments, all occupy 18% of the intertidal area of the Ythan.
of which can be consumed by Corophium. The level of production of the Mytilus aver-
aged over the whole estuary is 48.7 g flesh dry
4.3 The surface dwellersbenthic wt m!2 year!1, which is more closely compa-
suspension-feeders and algal grazers rable to other estuarine fauna (Table 4.2).
Large populations of the American oyster,
4.3.1 Benthic suspension-feeders
Crassostrea, are typical of the high salinity salt
The edible (or blue) mussel M. edulis develops marsh tidal estuarine systems common to the
large aggregated populations within estuaries southeastern United States, where they are
(Fig. 4.7). The accumulation of dead and live economically important. In the North Inlet
mussels typically leads to a mussel bed com- estuarine ecosystem near Georgetown, South
munity developing. In the Netherlands har- Carolina, the oysters form extensive bars in
vesting of mussels is a major industry. From a places completely lining the intertidal creek
140-km2 Dutch estuary, for example, mussel banks. Oysters may have a valuable role to play
harvesters collect an average annually of in increasing the productivity of the area in
19,225 tonnes of mussels. which they live by filtering out food from the
For all suspension-feeders the highest levels water column above them and depositing mate-
of production are only maintained in the mus- rial as feces and pseudofeces, which is available
sel or oyster beds, and average levels of pro- to the benthic organisms. The Pacific oyster,
duction for the whole estuary are much lower. Crassostrea gigas, can deposit material on the
For example, the highest level of production of bottom, equal to that resulting from the com-
the Mytilus population of 268 g flesh dry bined activity of plankton and gravitational
wt m!2 year!1 in the Ythan is only maintained sedimentation in a 45-m column of water, and
in the mussel beds of the estuary, which furthermore the biodeposited particles are
Figure 4.7 Mussel bed on the Ythan estuary, eastern Scotland. This suspension-feeding animal is often, as here, the main
inhabitant of the lower reaches of the estuary, feeding mainly on material brought in from the sea on the incoming tide.
P R I M A RY C O N S U M E R S 61
smaller and have a higher nutritive potential to In Spartina salt-marsh ecosystems, large
browsing organisms. One square meter of oys- quantities of plant fragments are potentially
ter bed can filter 282,720 1 year!1, and retain available to primary consumer animals.
about 11% of the food material in the water, However, studies of the mud-snails in the salt
amounting to 2570 kcal m!2 year!1. Much of the marshes of Georgia have shown that littler of
energy from the food filtered out from the water the Spartina detritus is utilized or available as
is utilized in metabolism and gamete produc- an energy source for the snails; instead they
tion, and small amounts of energy contribute to rely on the benthic algae or the bacteria, which
shell growth, but the bulk of the energy live on the plant fragments.
ingested, amounting to 1545 kcal m!2 year!1 is
rejected by the gills or passes through the gut
4.4 Meiofauna
undigested. This material is deposited on the
bottom around the oysters where it can support The meiofauna are those animals, which are
a larger number of small deposit-feeders. able to pass through a 0.5-mm (500 %m) sieve,
The edible cockle Cerastoderma ($Cardium) but are retained by finer sieves (usually
edule lives buried within the mud, with short 62 %m). They are thus defined mainly by their
incurrent and excurrent siphons protruding body size, and include chiefly animals with
above the surface when they are covered by small elongated bodies, which live intersti-
the tide. Its distribution in estuaries is gener- tially in sand, or in the loose upper layers of
ally patchy and production may vary annually mud. The most abundant of the meiofauna are
dependent on the success of the spat-fall. The the nematodes, and along with them may be
newly settled spat may achieve densities of found many groups of animals, especially
2400 m!2, but high rates of mortality soon harpacticoid copepods, and also Turbellaria,
reduce that number. Gastrotricha, Tardigrada, Archiannelida,
Smaal and Prins (1993) have shown that Coelenterata, and Annelida. Earlier studies of
suspension-feeders may generally be limited the meiofauna tended to consider their role in
by the food supply, which is directly affected by the energetics of marine and estuarine benthic
the residence time of the water in the estuary. ecosystems as being unimportant. Latterly
They found that the biomass of the suspension- there has been a realization that despite their
feeders was highest in estuaries where the small individual size and total biomass, the
water was being exchanged rapidly, and hence meiofauna can have a high productivity (high
the food supply was constantly being replen- P : B ratio) and may contribute substantially to
ished. In general suspension-feeders are con- the production of the estuarine benthos. The
strained by major hydrodynamic ($physical or meiobenthic community has been found to be
mixing) constraints, and therefore can only similar in many estuaries, being controlled
occur patchily on a limited fraction of the total more by predation and or interference from
surface area of the estuary. macrofauna than by food supply.
A comprehensive survey of the meiofauna
of the Forth estuary, Scotland, recorded a total
4.3.2 Grazers
of 172 different species, with a mean dry
The winkle Littorina littorea is a grazer on algal weight biomass for all the meiofauna of
films, utilizing its radula to remove this film 1.1 g m!2. Peak biomasses of about 4.0 g m!2
from rocky outcrops or off the surface of mud were recorded close to a sewage-works and an
and sand deposits. Within estuaries, L. littorea industrial effluent. Nematodes were found to
is very patchily distributed, and may often be be the numerically dominant taxon, supple-
found in salt marshes, eel-grass meadows and mented by oligochaetes in the upper estuary,
mussel beds. and copepods and polychaetes in the middle
62 T H E E S T UA R I N E E C O S YS T E M
50 Grangemouth Alloa 60
40 Total
% of total production
Number of species
40
30
Macrofauna
20
Nematodes 20
10 Meiofaunal nematodes
Copepods 0
0 0 10 20 30 40
A B C D F G H J K L N O P Q R S T U
Species number
Mouth Sites Head
Phytoplankton
& 183 54
benthic algae Planktivores Fish
52
n
ntatio 130
ime
Sed
Figure 4.10 Energy flow model (Odum circuit language) of the benthic ecosystem of the brackish Asko/Landsort area of the
Baltic Sea. Units are kilo joules per square meter per year. Figure to the left of the hexagons represent assimilation, figures
within hexagons represent biomass, figures to the right of the hexagons represent production, and those below the hexagons
represent respiration. (From Ankar and Elmgren 1976.)
and lower estuary. The total number of species Grangemouth area is apparently related to
was 50 per sample (2 ' 5.5 cm2) at the mouth industrial discharges at that locality.
of the estuary, and decreased steadily within The most abundant representatives of
the estuary to below 10 in the upper estuary, the meiofauna in estuaries are the free-
increasing to over 15 at the freshwater head living nematode worms. Figure 4.9 shows the
(Fig. 4.8). The dip in species number in the partitioning of annual production amongst
P R I M A RY C O N S U M E R S 63
secondary consumers such as shrimps, crabs, Table 4.6 Estimates of abundance of microzooplankton
and fishes. This may serve to explain why so (number m!3)flagellates ciliates and bacteria in the
pelagic zones of temperate estuaries
many carnivores from the open sea use shallow
estuarine areas as nursery areas for their Location Flagellates Ciliates Bacteria
youngest stages, as discussed in the next (" 109) (" 106) (" 1012)
chapter.
Sapelo island, USA 1.331 0.080.09
Georgia, USA 2.34 1629 217
4.5 Zooplankton Boston Harbor, USA 0.25 1030
Hudson Estuary, USA 0.91.7 35 1.97.6
The zooplankton is commonly defined as the
Clyde estuary, 0.550 0.00150 7.5
animals that float in the water column, and
Scotland
have only limited swimming capacity. Few Loch Striven, 0.550 0.00110 2.9
estimates of the biomass of the zooplankton Scotland
in estuaries have been undertaken and all
results are many times smaller than the rich Source: Heip et al. (1995) and references therein.
benthic populations that we have already
discussed. Estuarine zooplankton can be sub-
divided either by size into micro, meso, and Table 4.7 Grazing impact of protistans on bacteria and
macrozooplankton, or by the duration of the algae in estuarine waters
zooplanktonic stage. The permanent mem-
Location Protistans % of % of phytoplankton
bers of the zooplankton, such as the calanoid
bacterial production
copepods Acartia or Eurytemora, are known as production grazed
the holoplankton, and the temporary mem- grazed
bers, such as the larvae of benthic forms daily
(e.g. oysters or barnacles) whose life in the
Georgia, Flagellates 50
zooplankton is a dispersal phase, are known
USA and Ciliates
as the meroplankton. Chesapeake Flagellates 10023 5060
The microzooplankton is most typically Bay, USA
protozoans, such as flagellates and ciliates, Essex, Community 100
which graze on bacteria and phytoplankton. England
Bothnian Community 100
Abundances are in the order of up to 109 m!3
Sea,
as shown in Table 4.6. The impact of proto- Sweden
zoan herbivory on the size of the bacterial Hudson Community 321
populations can be very important, as shown estuary, USA
in Table 4.7. It can be seen that up to 100% Boston Ciliates 1590
Harbor, USA
day!1 of bacterial biomass and production
Narragansett Flagellates 100 1626 (annual)
and up to 60% of primary production by phy- Bay, USA
toplankton can be grazed by protozoans in
estuaries. Sources: Heip et al. (1995) and references therein.
The mesozooplankton is most typically the
calanoid copepods, including Eurytemora,
Acartia, and Pseudodiaptomus, with densities (Ctenophores, for example, Beroe) as well as
of up to 100,000 Ind. m!3, but usually less. true jellyfish (Scyphozoans, for example,
The macrozooplankton includes the mysid Aurelia).
shrimps (Neomysis, Praunus), also referred The mesozooplankton can be divided into
to as the hyperbenthos, as well as the large three main groups. (a) Allochthonous fresh (e.g.
gelatinous zooplankton, such as comb jellies Eurytemora) feeding on allochthonous material
P R I M A RY C O N S U M E R S 65
brought in to the estuary from the river; High population densities of endemic estuarine
(b) Autochthonous (e.g. Acartia) feeding on zooplankton are often observed when primary
phytoplankton produced within the estuary; production is small, showing that mesozoo-
and (c) Allochthonous marine (e.g. Pseudo- plankton are at least partly supported by detri-
calanus) feeding on allochthonous material tus as food. Because of their ability to graze on
brought into the estuary with the tides from the non-algal diets the endemic mesozooplankton
sea (Fig. 4.12). Estimates of estuarine copepod of estuaries may be able to maintain relatively
production are generally in the range of high stocks all year round. Through selective
830 gC m!2 year!1, which is close to values feeding the mesozooplankton can greatly influ-
reported from coastal marine areas (Table 4.8). ence the pelagic system of estuaries, thus in
Chesapeake Bay, it has been observed that on
average less than 50% of the phytoplankton
Zooplankton
biomass
was consumed by the mesozooplankton, but
b on one occasion the entire primary production
was grazed.
c Two features potentially limit the zooplank-
ton of estuaries. First by turbidity, which can
limit phytoplankton production and thus limit
a
the food available for the zooplankton, and
Salinity
second (and often more important) by cur-
rents which, particularly in small estuaries or
Figure 4.12 Idealized distribution of allochthonous and those dominated by high river flow, can carry
autochthonous zooplankton organisms along a salinity the members of the zooplankton out to sea.
gradient. (a) Allochthonous fresh, (b) Autochthonous, and The main solution to these problems is for the
(c) Allochthonous marine. (After Heip et al. 1995.)
zooplankton to stay near to the bottom and
utilize as far as possible the inflowing marine
Table 4.8 Yearly production of mesozooplankton in currents, but these are tidally intermittent, and
estuarine waters compared to coastal marine waters the best a species may achieve is to be carried
(gC m!2 year!1) to and fro about one location. Several species
have been shown to undergo selective vertical
Habitat Location Production P : B (year!1)
migration upward and downward during dif-
(gC m!2
year!1)
ferent phases of the tide, by means of which
they can maintain their location within the
Estuary Patuxent River, 9 71 estuary.
USA The catches of zooplankton in an estuary
Narragansett Bay, 1823 140150 can be variable. The most important factor
USA affecting variability is the tide, and therefore
Westerschelde, 11 45 to get accurate estimates of the zooplankton
Netherlands
it is necessary to take samples over a 24-h
Coastal Northumberland, 929 period. Taking this into account, Lee and
marine England McAlice (1979) estimated that there were
Scotia shelf, 8 9
23,882 copepods m!3 in the Damariscotta river
Canada
estuary, Maine, with 8571 Acartia tonsa m!3,
Southern 1223
North Sea, 7360 Eurytemora herdmanni m!3 and 3753
Dutch coast Acartia clausi m!3. The mean biomass of these
populations of zooplankton was 0.086 g dry
Source: Heip et al. (1995) and references therein. wt m!3, which with a P : B of 5 would only
66 T H E E S T UA R I N E E C O S YS T E M
create an annual production of 0.43 g dry The greatest constraint on a pelagic popula-
wt m!3. Production of zooplankton has been tion in an estuary is the lack of stability, result-
measured in Chesapeake Bay, and the neigh- ing from tidal mixing and high river flows.
boring Patuxent river estuary, and suggests Intuitively, a true planktonic fauna would
that zooplankton production is of the order of seem unlikely to develop in the upper estuary
510 gC m!2 year!1, with the highest values in where this instability is greatest. However, the
the upper estuarine reaches. copepod E. affinis, along with the hyperben-
A detailed study of the zooplankton of the thos, such as the mysid Neomysis integer,
Forth estuary identified a total of 135 taxa, thrives in such situations. Both these species
with 52 holoplanktonic taxa and 83 mero- may be regarded as epibenthic species, they
planktonic taxa. The commonest holoplank- both exhibit a marked shoaling behavior and
tonic species were the calanoid copepods they can apparently concentrate their distribu-
Acartia longiremis, Acartia bifilosa inermis, and tions very close inshore at slack water. These
Centropages hamatus, followed by the cyclopod behavioral features help them retain their
copepod Oithona similis and the chaetognath position in the uppermost parts of the estuary,
Sagitta elegans. Larvae of Littorina, Carcinas so that they can take advantage of a unique
maenas, and many polychaetes dominated the supply of food available there. This food sup-
meroplankton. Within the estuary a clear ply is available at, and downstream of the
sequence could be seen with the calanoid freshwaterseawater interphase (FSI). The
Eurytemora affinis totally dominating the upper processes involved are summarized in Fig. 4.13.
reaches, a complex of Acartia species in the As river phytoplankton and other material
middle and lower reaches, with Centropages, enter the head of the estuary they undergo
Oithona, and Pseudocalanus minutus at the plasmolysis, or rupture, due to osmotic stress,
mouth, and penetrating further into the estuary which liberates both soluble and particulate
in summer. organic material. These organic materials are
FreshwaterSeawater
Interphase Horohalinlcum
O2 CO2+H2O HCO3+H+
River-borne Osmotic stress NH3 NO2 NO3
biota Soluble organic
Bacteria
Plasmolysis material
Washout
Particulate Ciliates
organic material
Downstream
Copepods
Osmolysis
Advection of
stenohaline
marine biota
Figure 4.13 Summary of biological and related chemical mechanisms in low salinity regions of estuaries.
From P. Burkhill. (pers. comm.)
P R I M A RY C O N S U M E R S 67
utilized by bacteria. The bacteria are consumed until they find a suitable settlement site. The
by ciliate protozoa, which constitute the micro- role of estuaries in the transport of crab larvae
zooplankton and are mainly non-tintinnid has been investigated in the York river estuary,
ciliates. Eurytemora then feeds upon the cili- USA, where it was found that the larvae of
ates. Additional organic matter is carried into crabs, which are restricted to estuaries (e.g.
this region by inflowing bottom marine cur- Rhithropanopeus harrisii) were adapted to be
rents, and this material experiences osmolysis confined to the bottom waters of the estuary
in the low salinity waters of the FSI. The abun- so that any transport that occurred would
dant growth of bacteria in this zone, utilizing tend to be upstream and thus confine the ani-
the organic material that is liberated here, will mals to the estuary. Other more migratory
consume a considerable amount of oxygen, and crabs (e.g. Callinectes sapidus) produced larvae
may cause an oxygen sag in these waters. with no such adaptations so that the larvae
The concentration of zooplankton, albeit of tended to be carried out of the estuary into the
only one species, may thus often be greater in sea, and later on the animals underwent a
these uppermost reaches of an estuary, than in return immigration as juveniles or adults.
any other part of the estuary. The estuarine hyperbenthos are a group of
Within the Patuxent estuary, Maryland, animals that generally live close to the bottom
large quantities of detritus enter the water of and perform vertical migrations into the water
the estuary from the adjacent salt marshes. column. The commonest examples of estuar-
This occurs particularly in the months of ine hyperbenthos are the mysid shrimps,
January and February, when ice scour carries Neomysis and Praunus (Fig. 4.14). The mysids
the plant debris into the estuary. Within the are considerably larger that the copepods and
estuary the mysid shrimp Neomysis americana, in some cases may be the main predators upon
and also the planktonic copepod E. affinis use the mesozooplankton. Comparative studies of
the detritus, presumably in a similar manner European estuaries have shown how similar
to that described above for the FSI. The input is the pelagic community in all estuaries
of the detritus in the late winter has been examined, and that the hyperbenthos (mysids
found to generate a substantial production of etc.) in particular are an important link every-
Eurytemora of 1.38.65 g dry wt m!2 year!1. where, feeding upon the smaller zooplankton
Fish larvae, especially those of the striped bass and providing essential feeding for fish stocks.
(Roccus saxatilis), consume the Eurytemora in
turn. This simple food chain of detritus, to
Neomysis and Eurytemora, to striped bass lar-
vae is a rapid and efficient process, and it has
proved to be possible to correlate the landings
of striped bass with the severity of the winters,
with cold winters causing extensive ice scour
and export of detritus, leading to a large popu-
lation of Eurytemora and an increased popula-
tion of striped bass.
Oyster larvae are released into the zoo-
plankton where they selectively swim and
thus contribute actively to their retention
within the estuary. This swimming behavior is
connected to the increases in salinity that Figure 4.14 A typical Mysid crustacean (Opossum shrimp).
accompany a flood (incoming) tide, and thus These animals can be some of the largest animals within
helps maintain the oysters within the estuary the zooplankton of estuaries, being up to 3 cm long.
68 T H E E S T UA R I N E E C O S YS T E M
Other large members of the zooplankton are Table 4.9 Estimated biomass and production of four
the comb-jellies (Ctenophore) and the true jelly- communities on the Spurn Bight mudflats of the Humber
estuary in tonnes as an example of the calculation of total
fish (Scyphozoan). During the summer these
community production. Figures are given only for the main
gelatinous zooplankton can reach large num- species. Figures in parenthesis in kilogram per hact
bers through apparent population explosions.
Relatively little is known about these compo- Biomass Production (annual)
nent of the estuarine fauna, however, clogging (t) (t)
by jellyfish can be one of the commonest rea-
Spartina zone
sons for the cooling systems of estuarine
(91 ha)
located power stations being shut down. H. ulvae 14.1 14.1a
Hediste diversicolor 8.7 22.2
4.6 The primary consumer community Edukemius benedii 1.3
Total for Zone 24.4 36.4
Although large numbers of studies have been
(268 kg ha!1) (400 kg ha!1)
undertaken of the biomass and productivity of
Upper shore SILT
individual species within estuarine ecosys- (332 ha)
tems, fewer studies have been made of entire M. balthica 59.6 123.9
estuarine trophic levels or communities. H. ulvae 13.1 13.1
Overall estimates are compiled by adding the H. diversicolor 12.7 42.9
biomass and annual production for the differ- Total for Zone 91.5 182.3
ent communities or zones within the estuary. (276 kg ha!1) (549 kg ha!1)
In Table 4.9, as an example of such compila- Midshore mud
tions, the biomass and production of the Spurn (1747 ha)
Bight area of the Humber estuary is given. M. balthica 321.0 229.0
In Table 4.10 the total biomass and produc- C. edule 134.0 80.5
Nephtys spp. 64.5 147.0
tion of several estuaries are given. The levels
H. diversicolor 25.3 83.4
of production for estuaries tabulated in H. ulvae 19.0 19.0
Table 4.10 are much higher than reported for Retusa obtusa 5.1 24.0
marine areas, where 4.5 g m!2 year!1 has been Total for Zone 571.3 582.9
recorded for Loch Ewe, Scotland, and 1.7 g m!2 (327 kg ha!1) (334 kg ha!1)
year!1 has been recorded for the North Sea Downshore sand
benthos, off Northumberland. The highest (786 ha)
values recorded from freshwater localities are M. balthica 32.0 8.3
the studies of the eutrophic Loch Leven, Nephtys spp. 9.3 21.3
Scotland, where the bottom community pro- C. edule 3.9 2.4
Total for Zone 46.9 32.2
duced 525 J m!2 year!1 (equivalent to 26.3 g
(59.7 kg ha!1) (41 kg ha!1)
ash-free dry wt m!2 year!1).
It is the factors that control the distribution Total for whole 734.1 833.8
estuary (2956 ha) (248 kg ha!1) (282 kg ha!1)
of the individual species, which are respons-
ible for the assemblages that we observe in a
Productivity figures were not available for Hydrobia, this
estuaries, rather than some process of biolog- estimate assumes a P : B ratio of l and is, therefore, likely to be
ical organization. In assessing the total pro- a considerable underestimate. All these figures are liable to
ductivity of an estuarine ecosystem we should various degrees of under- or over-estimation and exclude Mya
thus remember that while it is useful to group and mysids etc. In general, the upper shore sites are likely to be
animals together in order to evaluate their underestimated whereas the lower shore figures are more
contribution to the next trophic level, explana- likely to be overestimates.
tions of the reasons for the distribution of a Sources: From Key (1983) in Jones (1988).
P R I M A RY C O N S U M E R S 69
Table 4.10 Biomass and production values for so estuarine macrobenthic assemblages,
all intertidal unless otherwise stated. Values as gram (ash-free dry weight) per square meter
per year, gram carbon per square meter per year
particular species should generally be sought abundance and diversity) in all estuaries
at the species level. examined. More marine macrobenthic com-
Studies of the benthic communities in sev- munities are found in the seaward part of all
eral large European estuaries have shown a estuaries, often dominated by filer feeders.
clear relationship to suspended matter con- It has been demonstrated in this chapter that
centrations and presumably silt deposition. In estuaries can sustain high levels of production,
sediments covered by turbid waters commu- especially among the benthic primary con-
nities dominated by deposit-feeders were sumers. These high levels of benthic produc-
found in the intertidal areas. In subtidal chan- tion are responsible for maintaining abundant
nels, tidal currents and the instability of the secondary consumers but at this stage we are
sediment became the limiting factors, leading led to ask how is it that estuaries can sustain
to very poor communities (in terms of biomass, high levels of benthic production. In Chapter 2,
70 T H E E S T UA R I N E E C O S YS T E M
on primary producers, we stated that two shallow nature of estuaries these sources of
distinct types of estuaries can be recognized. In food become rapidly available to the benthic
the case of American estuaries, especially those primary consumers. Sinking of surface produc-
studied in Louisiana, Florida, and Georgia, tion, coupled with tidal currents, may transport
which have small tidal amplitudes, large quan- phytoplankton from the surface to the filter
tities of detritus are derived from salt marshes, feeders within a matter of hours, and only a lit-
mangroves, or eel-grass beds within the estu- tle longer time is required before the organic
ary, which, supplemented by the primary pro- material sinks finally to the bottom where it
duction of phytobenthos and phytoplankton, becomes available to microbial decomposition
produce large quantities of organic carbon, and the detritus-feeding benthos.
which are available to the primary consumers From these considerations of the productivity
in the estuary. Currents from the estuary to the of the primary consumer populations within
adjacent sea export any excess organic carbon. estuaries it becomes clear that the high produc-
In the case of many north European estuaries, tivity is due to the large quantities of food mate-
especially those studied in The Netherlands or rial, which are potentially available. Abiotic
the British Isles, which have large tidal ampli- factors, such as currents, tidal exposure, sedi-
tudes, much of the detritus that supports the ments, or salinity regimes may limit the distri-
benthos is derived by importing currents from bution of individual species, and biotic factors,
the adjacent sea or river. Seventy-five percent especially competition between the macrofauna
of the particulate organic material in the and the small food web may limit the produc-
Wadden Sea, for example, is derived from the tion of any one group of organisms. We have
North Sea, and only 25% is produced within also seen in assessing the value of the primary
the estuary by the phytoplankton or phytoben- consumers as food for the next trophic level, the
thos. The most distinctive difference is that in secondary consumers or predators, that bio-
many American estuaries studied the marsh mass is often not a good guide to the impor-
grass Spartina alterniflora occupies about 2/3 tance of a species. Many slow-growing bivalve
of the intertidal zone. Whereas in typical molluscs have large biomasses, but even when
European estuaries salt-marsh plants are lim- the heavy shell is discarded, the remaining flesh
ited to a region above the height of high water may have a low productivity as expressed by a
of neap tides and 9095% of the estuarine inter- low P : B ratio (or turnover rate). Other organ-
tidal areas are bare mud and sandflats. Thus in isms such as annelid worms or amphipod crus-
the case of Spartina-dominated estuaries the taceans are faster-growing and shorter-lived
surface-floating plant fragments will be carried and may often achieve production for the next
seaward, whereas in mudflat-dominated estu- trophic level that is over five times their bio-
aries the denser detrital fragments will be kept mass. The smallest organisms of all, within the
in suspension and carried into the estuary by small food web may have the least biomass, but
the bottom inflowing saline currents. the highest production. The importance of all
Thus it can be recognized that the high level the primary consumers as prey organisms will
of primary production within estuaries is due be discussed in the next chapter.
to a supply of dissolved nutrients from various
sources, especially by release from the sedi-
4.7 Role of primary consumers in the
ments and by currents carrying nutrients from
estuarine ecosystem
the sea and to a lesser extent rivers. The pri-
mary production of phytoplankton and phyto- Initially many studies of estuaries concentrated
benthos is greatly supplemented by the on single species, measuring their consump-
addition of POM either from salt marshes or tion, growth, and the production of energy.
sea grasses, or from adjacent seas. Due to the In time such studies have been combined in
P R I M A RY C O N S U M E R S 71
(g AFDW m2)
OS
Ythan estuary, Scotland they found that the
40
main flows of energy were through the sedi-
ment pools and that all living components in 30
B2
the estuary were involved in the cycling
process. For the much larger Ems estuary a 20 EWVM
B1 SFB + CB
pelagic (or water column) cycle structure was BF LIS B = 1.5 + 0.105P
10 WS LY r 2 = 0.77
separate from the benthic cycle, with the ben-
ED
thic suspension-feeders as a link between the COL
0
two cycles. This difference was due to an 0 100 200 300 400 500 600 700
apparent lack of resuspension of sediment par-
System primary production (g C m2 year1)
ticulate organic carbon in the Ems. Baird and
Ulanowicz (1993) also showed that the func- Figure 4.15 Relationship between system-averaged
tioning of a pristine estuary was very similar macrobenthic biomass and primary productivity of shallow
to more anthropogenically enriched estuaries. well-mixed systems. (After Herman et al. 1999.)
These studies do show the crucial role of the
benthos in estuaries as the principal energy
link between the primary food supply and the web, and most of the organic matter that enters
consumers such as birds and fish. They also the estuary is used there. Only the more refrac-
emphasize the need to analyze the role of ben- tory ($difficult to break down) part escapes to
thos suspension-feeders separately from the the sea.
deposit-feeders and the benthic carnivores. In a review of the ecology of estuarine macro-
Modeling of the Westerschelde estuary benthos, Herman et al. (1999) conclude that
shows that 80% of the carbon that is respired the coupling between benthic and pelagic sys-
is imported into the system (Soetaert and tems is intense. Average benthic biomass is
Herman 1995a), mostly from upstream limited by primary productivity of the system.
(inland), but also from the sea, including the In Fig. 4.15 data on the relationship between
importation and subsequent death of marine benthic biomass and system productivity is
zooplankton. Locally produced carbon is rela- assembled, where it can be seen that for these
tively unimportant. Due to the high het- shallow well-mixed estuarine systems there is
erotrophic activity (net heterotrophic means a clear and highly significant relationship
that respiration is greater than autotrophic pro- between primary productivity and macrofau-
duction; i.e. a food web based on detritus), nal biomass. The relationship suggests that
nearly all imported and locally produced car- between 5% and 25% of the annual primary
bon is lost in the estuary itself, and the estuary productivity is consumed by macrobenthos.
is an insignificant source of organic material to On an estuary-wide basis suspension-
the adjacent sea. The estuary thus acts as a trap feeders can be the dominant component (with
for reactive organic matter, both from the land, respect to biomass) of estuarine benthic assem-
from the sea or produced within the estuary. blages, with values of 4186% of biomass
Internal cycling, mainly in the water column, being due to suspension-feeders in several
results in the removal of most of the carbon. less-turbid estuaries. In more turbid estuaries,
All the imported, relatively reactive organic however, the deposit-feeders dominate the bio-
carbon is incorporated into the estuarine food mass. Suspension-feeders typically occur in
72 T H E E S T UA R I N E E C O S YS T E M
much higher local biomass (e.g. Mussels or the flux of carbon into the sediment, and the
Oysters in beds) compared to deposit-feeders, availability of particulate organic carbon in
which are more evenly spread throughout the the sediment, all of which are determined by
estuary at lower biomass levels. Thus at a large complex processes, including biotic and abi-
scale food may be the limiting factor for benthic otic factors. Hydrodynamic processes, such as
biomass. Depending on the characteristics of the currents discussed in the first chapter,
the system, grazing by benthic suspension- largely determine sediment texture or particle
feeders may be the most important factor deter- size. The organic content of the sediment
mining dynamics of the entire estuarine system. depends both on deposition from the water
The general conclusions are that a substantial column as well as the reworking of the sedi-
fraction of the carbon flow in estuarine systems ment by its faunal inhabitants.
passes through the macrobenthic populations, Deposit-feeders transport particles and
and that therefore macrobenthic populations fluid during feeding, burrowing, tube con-
at an estuarine system level may be limited by struction, and irrigation activity. By enhancing
food fluxes to the sediments. transport of particulate organic carbon to
The distribution of suspension feeders is deeper sediment layers, these organisms
critically controlled by the availability of stimulate anaerobic degradation processes
food, and the seston within the body water as and so affect the form and rate at which
well as water currents. Grazing by benthic metabolites are returned to the water column.
suspension-feeders has been shown effect- This bioturbation causes and upward move-
ively to control phytoplankton in estuaries ment of reduced components such as sulfide,
such as the Oosterschelde in the Netherlands, as a consequence of which sedimentary oxy-
and it has been shown that benthic grazing gen uptake can increase. The release of nutri-
can limit phytoplankton bloom development in ents such as nitrogen can be affected, and the
shallow water situations where vigorous verti- reduction in the release of nitrogen due to
cal mixing is occurring and where grazing is benthic denitrification can directly influence
sufficiently intense. But is has also been shown the availability of this nutrient for phytoplank-
that as soon as even a mild form of permanent ton production. The animals of the primary
vertical stratification affects the system, then consumer community thus both influence
benthic grazing is no longer a major factor. the productivity of the primary producers, as
The distribution of deposit-feeders is con- well as providing food for the secondary
trolled in large part by the nature of sediment, consumers.
CHAPTER 5
Adult
carcinus
Haematopus
Pleuronectes
Crangon
Nereis
Figure 5.1 Predator spectrum of
the cockle Cerastoderma edule
Carcinus
growing from spat to adult size
Length (mm) 1 10 20 30 on tidal flats at the Island of
Sylt, German Wadden Sea. (After
Age (years) Spat 1 2 3 Reise 1985.)
73
74 T H E E S T UA R I N E E C O S YS T E M
Spawning
feed on the animals of the primary consumer i g r
ts
trophic level, but it should be noted that a al m dul
Tid ub-a
S
few birds by-pass this trophic level and instead
feed directly on the primary producer plants.
Most notable of these are the widgeon (Anas Benthic
penelope) and Brent geese (Branta bernicla) feed- adults
ing on eelgrasses and algae. Other birds such
as the fish-eating birds (e.g. cormorant and
merganser), or mammals such as seals, feed Subtidal Tidal
on the members of the secondary consumer Figure 5.2 Common life cycle in demersal fish and mobile
trophic level, and should properly be called the crustaceans living in estuarine and coastal regions. The
tertiary consumers. pelagic larvae develop offshore, drifts inshore, and there
Migrations between habitats represent a fun- become benthic juveniles. Later stages migrate tidally. The
damental aspect of the ecology of populations estuarine intertidal areas function as nursery grounds.
(After Reise 1985.)
and individuals. The complex movements
between breeding, wintering, or feeding areas
during various life-history stages are often The fish fauna of estuaries may be classified
critical to the survival of species. Estuaries are into various categories (Box 5.1). The marine
viewed as critical nursery environments sup- components are generally the dominant con-
porting a rich variety of seasonal visitors. The tributors to the diversity of estuarine species,
best-known examples are probably waterfowl, here fish, and true estuarine, and other com-
but estuaries are used in a similar way by fish ponents contribute fewer species.
and shrimps for feeding and wintering. Estuaries perform a crucial role in the
population dynamics of many fish. They pro-
vide a migratory route for anadromous and
5.2 Fish
catadromous species and an environment for
Fish populations in estuaries can be abundant truly estuarine residents. Freshwater fish
with a wide diversity of species. Much of the colonize the head of estuaries while typical
abundance is seasonal as marine fish move into marine species use the mouth. Many marine
the estuary to breed and having used the estu- fish enter and remain in estuaries for a brief
ary as a nursery the young fish grow and move period of time, often in large numbers and
out to sea again (Fig. 5.2). Other fish such as particularly during the early part of their life
salmon (Salmo salar) and eels (Anguilla anguilla) cycle. A number of mechanisms have been
use the estuary as a migratory route to get from suggested to explain this feature, including the
rivers to the sea and vice versa, rarely feed- avoidance or attraction to abiotic factors or grad-
ing in the estuary. Only a few species live in the ients, the reduction intraspecific competition
estuary throughout the year. Notable amongst with adults of the same species, the reduction
these residents is the killifish (Fundulus in predation levels and the increase in food
heteroclitus) in American Spartina-dominated availability.
estuaries and various species of flatfish Young fish utilize estuaries and near-shore
such as flounders (P. flesus, Pseudopleuronectes marine areas as nursery areas in order to
americanus) in mudflat-dominated estuaries. benefit from the availability of food and
The flounders food consumption may often perhaps also to gain protection from predators.
rival or exceed that of the birds. On the large estuary of the Patuxent in
T H E S E C O N D A RY C O N S U M E R S 75
Box 5.1 Ecological guilds of estuarine fishes (from Elliott and Dewailly 1995)
Maryland, for example, a clear seasonal pattern estuarine fish populations can be gained
of abundance of fish has been noted with col- from an examination of the large numbers
lections of fish more abundant and diverse in of fish that become trapped on the cooling-
warmer water (spring and summer) and on the water intake screens at power stations. At
shores of the estuary. Collections in colder the Fawley power station in Southampton
water (autumn and winter) are less abundant Water, England, for example, fish catches of
and diverse and mainly in the main channel of up to 60,000 per week occur. Elsewhere in
the estuary. These differences reflect the large Britain a massive influx of sprats (Sprattus
number of young fish that use the shore areas sprattus) blocking the cooling-water intake
as nursery feeding areas in spring and summer. screens of a power station caused extensive
Estimates of the fish population in estuaries damage. Collections at three power stations
are more difficult to make than, for example, on the Scheldt estuary show both the num-
estimates of estuarine bird populations. bers and variety of fish that enter estuaries
Some idea of the diversity and abundance of (Fig. 5.3).
76 T H E E S T UA R I N E E C O S YS T E M
(a) Total species number plaice then consume as much as half of the
25 invertebrate food available for all secondary
consumers between March and July. It has
20 been found that much of the food of the young
Number of species
Table 5.1 Occurrence of seven macrobenthic species at three tidal levels on the tidal flats of the Oosterschelde estuary,
compared to the average number of prey specimens in the stomach and guts of plaice and flounder
diversity in the coldest month, February. It was 0.54 g dry wt m!2 year!1 are typical so it can
also found that areas with the maximal salinity be seen that these studies of Fundulus are
variation had the most diverse fish faunas. between 4 and 20 times higher. The killifish in
The killifish or mummichog, F. heteroclitus Massachusetts have a 3-year life span, during
is one of the most abundant resident fishes which time they are subject to intense preda-
within the tidal creeks of Spartina-dominated tion by birds and other larger fish. In contrast
salt marshes where they feed principally to many other estuarine fish species that are
on benthic invertebrates. Studies in the seasonally migratory, the Fundulus remain for
salt marshes of Delaware and Massachusetts their whole life confined to a single marsh sys-
have revealed annual productivity values of tem, apparently relying on the tides to bring
1016 g dry wt m!2 year!1. The productivity their rich food supply to them. However, their
values from Delaware were calculated on sedentary behavior has permitted a close
the basis of the populations of Fundulus within examination of their productivity to be made,
a 36-m long portion of a 3-km long creek. in contrast to the many estuarine fish popula-
Within the 36-m portion the mean biomass tions elsewhere about which little is often
was 32597298 g-wet wt, and the population known.
numbered 378814 individual fish. The On mudflat-dominated estuaries the most
Massachusetts study was based on the results common fish are often the flounder (P. flesus)
derived from eight tidal creeks, in which min- and the sand goby (Gobius minutes). On the
now traps were placed, and the fish caught Ythan estuary, Scotland, for example, the num-
were marked, released, and then a proportion ber of flounders varies from 0.05 m!2 in winter
caught again. Such estimates of production to 0.24 m!2 in July, representing biomasses
suffer from possible errors in the estimation of from 5 to 35 g wet wt m!2. The total food
of the size of mobile populations, never- consumption of flounders in the Ythan has
theless, the consistency between the two stud- been calculated as 58.1 kcal m!2 year!1, which
ies does point to the high productivity of these may be compared to the consumption of
fish populations. In non-estuarine studies gobies at 8 kcal m!2 year!1, and the combined
of fish populations, productivity values of consumption of birds (Oystercatcher, Dunlin,
78 T H E E S T UA R I N E E C O S YS T E M
Redshank, and Shelduck) in the same area at cycle, but most use it only during some
23.9 kcal m!2 year!1. Thus in this estuary at seasons or for different stages of their life cycle.
least the fish populations appear to consume Many of these species make considerable feed-
three times the amount of food consumed by ing and spawning migrations, such as Herring,
the birds. which drift from their spawning areas on the
The winter flounder (P. americanus) in East of Scotland and England into the local
Newfoundland feed mainly during the estuaries, as well as across to the German Bight
summer, consuming principally polychaetes, and NetherlandsGermanyDenmark coasts.
plant material, molluscs, capelin eggs, and For many, if not most, of the gadoids (Whiting,
fish remains. The winter flounder is the Saithe etc.) the older age groups occur in
dominant fish in the polluted Mystic River deeper waters, while the juveniles occur in
estuary in Massachusetts, comprising 89% of coastal waters and estuaries. The estuaries of
the total fish biomass. The winter flounder the North Sea are firmly recognized as import-
goes through its entire cycle in the estuary ant nursery areas for North Sea fish stocks.
feeding principally on the Capitella worms that The process of selective tidal stream transport
are the dominant members of the benthic by larvae of the Flounder (P. flesus) encourages
fauna. Only two other fish species spent the the retention of the larvae inside the estuary as
entire year in this estuary, the alewife (Alosa a nursery ground. This is shown by the vertical
pseudoharengus) and smelt (Osmerus mordax). movement of the larvae within the estuary
Another 18 species visited the estuary season- deviating from the movement of suspended
ally, mostly in summer. The biomass of fish for matter, so that the larvae could utilize tidal
this estuary averaged 2 g wet wt m!2, a low streams to remain within the estuary.
value that reflects the polluted conditions in Plaice, Sole, and Dab are classic examples of
this estuary. species with nursery grounds. Their bottom-
A study of the fish assemblages in estuarine living 0-group stages occur exclusively in very
waters around Sweden has shown that the shallow water (littoral or sub-littoral). These
number of species is limited by the salinity, three species are therefore relatively scarce in
with fewer species in less saline waters, the northern North Sea, where there is a
but the abundance of fish was determined dearth of suitable nursery grounds, while they
by temperature, with more fish in warmer are abundant round the perimeter of the
waters. The fish coexist with mobile inverte- southern North Sea, especially in the Wadden
brates, such as crabs (C. maenas) and shrimps Sea and other North Sea estuaries. Dabs spend
(Crangon, Palaemon), which feed on the same a comparatively short time in the nursery
bottom fauna. In summertime the small fish areas, but Plaice and Sole may spend 1 or
have to compete with shrimps as well as other 2 years. Cod too spend time inshore and
fish species for their food, and this may serve in estuaries, with the German Bight being
to limit the numbers of fish in an estuary. a major nursery. It is probable that a large
Apart from competing with the shrimps and proportion of the cod in the northern North
other mobile fauna for benthic food, fish such Sea spend their first 1 or 2 years inshore along
as cod and flounder also eat these animals and the UK coast, where they cannot be caught
in a study in one shallow bay it has been by conventional survey gear.
shown that the fish consumed about 7% of the Reviews of the possible physiological rea-
production of the mobile epifauna. sons for estuarine usage by fish conclude that
The estuarine fish communities of the North the most likely cause is the presence of high-
Sea comprise several components each with density prey and hence favorable feeding
differing requirements in the ecosystem. Some areas. For the Forth they show that the estuary
fish use the estuary during their whole life supports a population of about 0.5% of the
T H E S E C O N D A RY C O N S U M E R S 79
North Sea stocks of similar sized Plaice and most commonly between about 5 and 15 g m!2.
Cod. The figure of 0.5% may appear small, but These values are an order of magnitude higher
it must be remembered that the Forth estuary than those of tropical non-estuarine sea grass
is much smaller than 0.5% of the area of the areas in shallow waters are (Blaber 1997).
North Sea, so therefore it must be considered The number of fish species in subtropical
that its usage is much higher than expected on and tropical estuaries is much greater than in
a simple proportional basis. temperate regions. Where individual temperate
Three percent of North Sea Plaice utilize the estuaries may have about 20 species (Elliott
outer Humber estuary as a nursery feeding and Hemingway 2002) and warm temperate
ground. Studies into the year-class strength ones about 50 (Lenanton and Hodgkin 1985),
of Plaice in the North Sea have suggested most medium to large subtropical and tropical
that processes acting during their post-larval estuarine areas have at least 100, with some
demersal stage in the nursery areas such as reaching over 200 (Table 5.2). Additionally,
estuaries or the Wadden Sea may determine the number of species in adjacent estuarine
this. Year-class strength was in large part coastal areas often exceeds 300, but depends
controlled by the influence of the predator, upon the presence of specific habitats such
C. crangon, which feeds upon the Plaice when as sea grass beds. The composition of the
they are in these nurseries. For other fish, fish faunas results from the interplay of a
Crangon is essential as their principal dietary whole range of factors among which the most
item, as shown by stomach analyses of important are:
juvenile Whiting, Cod, and Haddock found in
(a) Estuary size, depth, and physical regimes,
estuaries.
particularly salinity and turbidity, as well as
The species richness of fish in tropical
the types of habitats, particularly the composi-
estuaries is greatest in the Indo-West Pacific,
tion and extent of mangroves.
but larger systems usually have more species
(b) The nature and depth of adjacent marine
than smaller ones. Deep open water channels in
waters and to a much lesser extent, freshwaters.
the larger systems favor more of the larger
(c) The geographical location of the estuary,
species, particularly carangids and sharks, in
both in terms of latitude and in relation to
addition to a higher number of occasional visi-
marine features such as ocean currents,
tors. In addition, larger systems usually have
canyons, and reefs.
a greater diversity of habitats. Hence compar-
isons of species richness between estuaries of The classification of estuarine fishes has been
the Indo-West Pacific and the East and West almost as numerous as the classification of
Tropical Atlantic should be viewed with cau- estuaries (Blaber 2002) with taxonomic, physio-
tion. However, taking into account the large logical, and ecological groupings based on
size of some Atlantic systems (e.g. Orinoco attributes such as salinity tolerance, breeding,
estuary, Termins and Lagos Lagoons), it is feeding, and migratory habits. Most research
apparent that even small estuaries in the on estuarine fishes has been in temperate and
Indo-West Pacific are usually more species rich warm temperate regions of Europe and North
(Table 5.2). Comparisons of overall fish America where salinity has long been regarded
biomasses and productivity (Tables 5.3 and 5.4) as a key factor regulating the composition of
in different tropical estuaries are difficult estuarine fish faunas. The lower salinity of
because there are few published figures, and estuarine waters is considered to be the out-
also because many of the results may not be standing difference between estuarine waters
comparable due to differences in methods. and seawater. While this phenomenon has
However, the scarce data suggest that fish bio- been amply demonstrated for warm temperate
mass does not usually exceed 30 g m!2 and is and temperate areas it is not at all clear in the
80 T H E E S T UA R I N E E C O S YS T E M
Table 5.2 Numbers of species recorded from different types of estuaries (O $ open, B $ blind, CL $ coastal lake) in the
four major zoogeographic regions
Indo-West Pacific
Alligator Creek Australia O small 150
Trinity Australia O medium 91
Embley Australia O large 197
Vellar Coleroon India O large 195
Chilka India CL large 152
Hooghly India O large 172
Chuwei Taiwan O small 30
Ponggol Singapore O medium 78
Matang W. Malaysia O medium 117
Kretam Kechil E. Malaysia O small 44
Purari New Guinea O large 140
Pagbilao Philippines O medium 128
Solomon Islands Solomon Islandsa O small 136a
Morrumbene Mocambique O medium 113
Tudor Creek Kenya O small 83
Gazi Bay Kenya CL small 128
Kosi South Africa CL large 163
St Lucia South Africa CL large 110
Mhlanga South Africa B small 47
East Atlantic
Lagos Nigeria CL large 79
Fatala Guinea O medium 102
Ebri Ivory Coast CL large 153
Niger Nigeria O large 52
West Atlantic
Guaratuba Brazil CL medium 61
Itamaraca Brazil O large 81
Orinoco Venezuela O large 87
Terminos Mexico CL large 122
Cienaga Grande Colombia CL large 114
Tortuguero Costa Rica O small 70
Sinnamary French Guyana O medium 83
East Pacific
Rio Claro Costa Rica O small 22
Guerrero Lakes Mexicob CL small 105
a
Incorporates 13 small estuariesno one system more than 50 species.
b
Sum of species for 10 small coastal lakes.
Source: Modified from Blaber (2000).
subtropics and tropics. Salinity is only one of a role in the distribution of fishes in tropical
an array of factors that determine which estuaries, but as these systems often undergo
species are found in any one subtropical or very great fluctuations in salinity (often from
tropical estuary. Salinity tolerance may play almost 0 to 35), both diurnally and seasonally.
T H E S E C O N D A RY C O N S U M E R S 81
Table 5.3 Biomasses of fishes from various subtropical and tropical estuaries
most species. Based on their occurrence in The fish communities of subtropical and trop-
low salinity waters almost all fit into a similar, ical estuaries of all four major tropical zoogeo-
very euryhaline category. This has led to graphic regions (Indo-West Pacific, East
a number of classification systems for sub- Atlantic, West Atlantic, and East Pacific) have
tropical and tropical estuarine fishes (Blaber many common characteristics. In almost all
1997; Whitfield 1998), based mainly on how cases, fishes of marine origin dominate them
they utilize estuaries and where they spawn. with more than half the number of species as
The following classification scheme is similar, well as the number of individuals, being con-
but gives increased importance to the estuar- tributed by either fully estuarine species or
ine category in the tropics due to recent marine migrants. Given the estuarization of
advances in knowledge of spawning behavior many tropical coastal areas, this is not surpris-
(Blaber 2000). ing and perhaps the term marine for many
of these species is a misnomer, for many of
1. Estuarine species that can complete their them do not occur outside of estuarine coastal
entire life cycle in the estuary. In the Subtropics areas. The dominant taxa in each region are
they represent a small but significant part of broadly similar, but there are some interesting
the estuarine fish fauna. In the Tropics they contrasts. In all regions except the Indo-West
represent a large part of the estuarine fish Pacific, Sciaenidae are one of the dominant
fauna, particularly in West Africa, South East families. In the Indo-West Pacific sciaenids are
Asia, and tropical South America. important in the equatorial regions of South
2. Marine migrants from the seausually East Asia, but much less so elsewhere. This
characterized as marine spawners. This is a pattern may be connected with the amount of
large group in subtropical and tropical estuar- rainfall and the degree of estuarization of
ies where they may occur both as adults and or coastal waters. In East Africa, for example,
only as adults. However, it is important to note there are few estuarine coastal waters and sci-
that the division of this category from that of (1) aenids, and although present they are not a
above is often difficult in the tropics, because dominant component of the fauna. Except for
many of the species that fall into the marine the sciaenids, the other dominant families are
migrant category in the subtropics, form part of broadly comparable across all regions. It is
the estuarine category (above) in the tropics. noteworthy, however, that within the Indo-
3. Anadromous species that breed in fresh- West Pacific, clupeids and engraulids are much
water and on their way to and from their more diverse and numerous in the estuarine
spawning grounds spend time in estuaries. coastal waters of equatorial South East Asia
This group is important in temperate estuar- than in other areas.
ies, but is a rare category in subtropical and
tropical estuaries.
5.3 Invertebrates
4. Freshwater migrants that move varying
distances down estuaries, sometimes to Invertebrate predators are principally crabs
spawn (e.g. eleotrids), but usually return to and shrimps, and carnivorous polychaetes,
freshwater to breed. In the tropics this group is the former mobile with the tides, and the latter
best represented in South and Central buried in the mud. Shore crabs (C. maenas) in
American estuaries where a number of the English estuaries show a seasonal migra-
pimelodid catfish, poeciliids, and characids tion. In the wintertime they are spread
penetrate estuarine waters. The freshwater throughout the estuary, but during the sum-
cichlid Oreochromis mossambicus and the archer mer the crabs move upstream and the lower
fishes (Toxotidae) are well known African and half of the estuary becomes devoid of crabs. In
Southeast Asian examples. November they again spread throughout the
T H E S E C O N D A RY C O N S U M E R S 83
estuary. This upstream estuarine migration is in population size as food becomes available.
apparently similar to the onshore summer In Narragansett Bay, United States, a biomass
migration that takes place in marine crabs. In peak of 60 g wet wt m!2 of Mnemiopsis has been
the Isefjord, Denmark, a negative estuary, the reported.
shore crab (C. maenas) move inshore in May as The mobile shrimps and prawns (e.g.
the temperature rises above 9 &C, with the C. crangon) and the opossum shrimps (e.g.
largest ones first, followed by the smaller indi- Praunus flexuosus and Neomysis integer)
viduals. They feed actively inshore during the are often important predators in estuaries.
summer and autumn and move offshore in The high densities of shrimps may often make
OctoberNovember and hide more or less them one of the main commercial catches in
inactively in deeper water during the winter. estuaries. Like crabs, shrimps tend to migrate
Such migrations seem to be a clear adaptation further into estuaries and into shallower water
to make use of the rich benthic production in summertime and retreat to deeper water
available during the summer months. in wintertime. When feeding in estuaries
Caging experiments designed to exclude they tend to be omnivorous, utilizing plant
crabs and gobies, and monitor the effects of fragments as well as smaller planktonic
these species on benthic infauna, have shown animals and members of the benthos. In
that predation by the shore crab C. maenas the Dutch Wadden Sea the juvenile Crangon
can significantly reduce the abundance of feed on planktonic copepods, but as they grow
small annelids on an estuarine mudflat, partic- the Crangon switch to feeding on benthic
ularly the dominant polychaete Manayunkia foraminifers and polychaetes. A study of the
aestuarina. Predation could be responsible predators feeding on Corophium in a Swedish
for year-to-year variations in abundance of estuary, has shown that 98% of the annual
this species, whereas in this experiment, the production of Corophium is consumed by
fish had little direct effect on the abundance shrimps, crabs, and fish, with 57% being
of prey. consumed by Crangon, 19% by Carcinus, and
In the Lynher estuary, Devon, the carnivo- the balance (22%) divided between three fish
rous benthic polychaete, N. hombergi occurs species (Fig. 5.4). Crangon also feeds on Nereis
abundantly with a mean annual biomass
of 3.947 g m!2 and an annual production of
7.335 g m!2 year!1. Since populations of other 6
macrofaunal invertebrates were inadequate
Production (g AFDW m2)
4
be meiofaunal. The carnivorous polychaete, Crangon
Nereis virens often forms substantial popula- 3
tions, feeding on smaller polychaetes. The rag-
2 19
worm Nereis diversicolor has been shown to be Carcinus
a predator on Corophium volutator, small 1
Fish 22
Macoma balthica and chironomid larvae, as
well as an omnivorous feeder on detritus. Apr Jun Aug Oct
Predation also occurs within the zooplankton,
for example the planktonic sea gooseberry, Figure 5.4 Cumulative production of C. volutator, and
the ctenophore, Mnemiopsis leidyi, which is cumulative consumption of Corophium by the dominant
epifaunal predators, in shallow areas of Gullmarsfjord on
a carnivore feeding on smaller members of the Swedish West Coast. Percentage consumption indicated
the estuarine zooplankton, can reach large on right. Production given as g ash-free dry weight m!2.
biomasses through its ability to rapidly increase (After Pihl 1985.)
84 T H E E S T UA R I N E E C O S YS T E M
and young bivalves, as well as juveniles of Some of the ducks such as Shelduck or eider
Carcinus and its own offspring. From several breed in the estuaries around the North Sea in
studies it has been concluded that Crangon is summer, but they are supplemented in winter
one of the major carnivores of shallow marine by others migrating from further north. Geese
and estuarine areas, and exerts a major impact may be present in large numbers in estuaries
on the infaunal community. as they pass through on migration while the
Gizzard contents of the predatory snail Brent geese (B. bernicla) may be resident in
Retusa obtusa show that the youngest and the winter. Living throughout the year in the
oldest feed on foraminifers, but the growing temperate estuaries of Europe are birds such
intermediate stages eat the snail Hydrobia as mallard, gulls, and cormorants (Fig. 5.5).
ulvae. There is a close match of predator and The bird predators of estuaries are highly
prey size, so that young Retusa feed on small mobile and typically exhibit clear tidal rhythms
Hydrobia, and the sizes increase correspond- of activity associated with water movements
ingly throughout the summer of growth. and the activity of the prey in relation to
The total impact of invertebrate and fish the tides. The Redshank (Tringa totanus) and
predation in estuaries is large, especially as the Shelduck (Tadorna tadorna), which feed on
many of the predatory species move into the Macoma, Hydrobia and other infaunal inverte-
estuary in summertime when the maximal brates feed mainly on the intertidal zone at
biomass of prey is available. low water, tending to be near the waters edge.
Eider ducks (Somateria m. mollissima) feed in
the shallow water at low tide, collecting in
5.4 Birds
particular the mussel, Mytilus edulis. Others,
The numbers and variety of birds of estuaries such as the diving ducks, (goldeneye, Bucephala
vary throughout the year. In Europe, many of clangula or scaup, Aythya marila), along with
the most abundant species of Waders arrive in cormorants (Phalacrocorax carbo) and mer-
the autumn on migration from summer breed- gansers (Mergus serrator) feed at high water by
ing habitats further north or east. Some of diving for their prey. In addition to the birds,
these stay in the estuaries of Britain and the which feed in an estuary others such as geese,
Netherlands throughout the winter, while various species of gulls, certain ducks, and
others move on south to overwinter in Africa. swans may use an estuary primarily as a place
100,000
80,000
60,000
Figure 5.5 Monthly mean
40,000 numbers of ducks, geese,
20,000 waders, and other water
birds counted in the
0 Scheldt estuary during
J A S O N D J F M A M J the period 19911997.
Month (After Ysebaert et al. 2000.)
T H E S E C O N D A RY C O N S U M E R S 85
to roost and obtain their food either at sea or The waders are one of the most important
from land sites. groups of estuarine birds, especially in
Many shore birds have adapted their food- autumn and winter. They feed on a variety of
searching behavior to suit the tidal and prey organisms depending on what is avail-
seasonal rhythms of their preferred prey. It able in their particular estuary. The principal
has been generally shown that the bill-length prey of Waders in the Wash, eastern England
of particular species of birds is suited to partic- is shown in Table 5.5, in which it can be seen
ular species of prey organism, but more how the prey resources have been divided
subtle behavioral adaptations also occur. The between the species of Waders with only
bar-tailed godwit (Limosa lapponica) feeds prin- limited overlap between species.
cipally on the lugworm Arenicola marina, and Oystercatchers (Haematopus ostralegus) feed on
waits and watches for new Arenicola casts so mussel beds when they are completely exposed
that it can catch the lugworms as they defecate. at low tide, by hammering on one of the bivalve
The Grey plover (Pluvialis squatarola) searches shells until it splits open, or inserting their
by sight for its worm prey, relying on the move- bill between the valves to cut the adductor
ment of the prey (the ragworm, N. diversicolor) muscle. Once the shell is open, the flesh is picked
at the surface of the substratum. Knot (Calidris out and the shell discarded. In the Wash the
canutus) on the Tees estuary feed on Hydrobia, Oystercatchers eat cockles and their daily con-
and like Shelduck on the Forth (see below), sumption of cockles has been estimated as 181
rely on the re-emergence of Hydrobia as the cockles for each bird each day ($57.5 g total wet
mudflats are reimmersed by the tide. The weight). Oystercatchers in North Wales feed on
ragworm N. diversicolor retreats into its burrow
as the tide ebbs off intertidal mudflats, but
is nevertheless caught by curlew (Numenius Table 5.5 Principal prey species of the main wading
birds in the Wash
arquata) relying on its sense of touch. Curlew
also feed on shore crabs (C. maenas), but as Bird Prey
these move offshore in winter, they are only
important in the summer diet. Oystercatcher C. edule
The invertebrate prey may adjust their (H. ostralegus) M. edulis
behavior in response to the activity of the Knot (C. canutus) M. balthica
predators. Macoma, for example, in the Wash C. edule
Dunlin (C. alpina) H. ulvae
apparently remain close to the mud surface
N. diversicolor
at low water whenever their predators are Redshank (T. totanus) C. maenas, Crangon sp.
absent, but bury deeper into the substrate at H. ulvae, Nereis sp.
low waters whenever predators are present. Bar-tailed godwit (L. lapponica) L. conchilega
The position of burrowing animals within the Nereis sp., M. balthica
substrate is most closely linked to the state of Turn stone (Arenaris interpres) C. edule
the tide. In experimental studies of Wadden Among mussel beds
Sea animals, it has been clearly shown that Grey plover (P. squatarola) L. conchilega and
H. ulvae and N. diversicolor migrate from being various
buried in the mud to the surface in direct Curlew (Numenius arquata) C. maenas,
response to the inundations of the tide over L. conchilega,
A. marina
the mudflats. There is no need to invoke any
Ringed plover (Charadrius Not recorded
inherent rhythm to the animals behavior, and hiaticula)
thus the animals only come to the surface to Sanderling (Crocethia alba) Not recorded
feed just prior to inundation and when they
are actually covered by the tide. Source: Goss-Custard et al. (1997).
86 T H E E S T UA R I N E E C O S YS T E M
the buried bivalve Scrobicularia plana by thrusting between feeding hours (i.e. number of birds
their bill deep into the mud, almost up to their and hours spent feeding per tide) and numbers
eyes, then jerking out the bivalve. The extracted per kilometer squared, and the density of at
bivalve is then wedged apart by the bill, and the least one of their main prey. The significant
soft parts removed. associations, which explain up to 96% of the
Within the Morecambe Bay area, at the observed variation in bird distribution
mouths of the Leven, Kent, and Lune estuaries, were found for Oystercatcher with mussels
northern England is a large overwintering (M. edulis), for curlew, Redshank and Dunlin
population of knot (C. canutus). The knot with N. diversicolor, and for knot with
breeds in the summer in the tundra of Cerastoderma (!Cardium) edule. Additionally, the
Greenland and NE Canada, where it feeds distribution of bar-tailed godwit, knot, and
almost entirely on insects. In winter it moves Dunlin was related to the area of the intertidal
south to the estuaries and coasts of Europe mudflat, and Redshank to the exposure
and North Africa and about 16% of the total sequence (length of time exposed by low tide).
population spends the winter in Morecambe The results of this study of bird distribution
Bay. The 70,000 birds at Morecambe Bay feed in relation to prey density are summarized in
predominantly on M. balthica, supplemented Table 5.6.
by M. edulis and H. ulvae. The choice of Macoma The steady fall in the numbers of Dunlin,
appears to be dictated by its availability, as the Calidris alpina spending the winter in Britain
knot feed predominantly on the lower half of may be caused by the spread of Spartina over
the intertidal areas and thereby feed mainly the mudflats where the birds feed. It has
on the Macoma, which are generally lower been shown that the decline in the number
down the beach than Hydrobia. of Dunlin varies from estuary to estuary. In
The preferred diet of Redshank (T. totanus) estuaries where Spartina has not spread over
at many estuarine sites in the British Isles is the mudflats, the numbers have not changed,
the amphipod C. volutator, and the feeding and the largest fall in numbers has been where
rate of Redshank depends mainly on the den- the grass has spread most rapidly.
sity of that prey in mud. Where the density of Studies on the Tees estuary, NE England sug-
Corophium is low, they take the polychaete gest that up to about 90% of the macrofaunal
worms N. diversicolor and N. hombergi instead. biomass is removed by seven species of wader
The densities of both Redshank and curlew and duck. Most of the bird populations feed
(N. arquata) on the Orwell, Stour, Colne, at the Tees during the winter time when the
Blackwater, Crouch, and Roach estuaries in invertebrate prey is not undergoing growth, so
SE England are closely correlated with densi- it is therefore reasonable to compare the birds
ties of their main prey (C. volutator and feeding requirements with the biomass of the
N. diversicolor). The distributions observed are invertebrates. The birds select different prey
due either to the birds responding behav- items (see Table 5.7) to some extent, which
iorally to the density of the prey in different enables them to live alongside each other. In the
estuaries and dispersing themselves in rela- Sheepscot estuary, Maine, gulls (three Larus
tion to it, or to the birds dispersing themselves species) have been estimated to remove 6.8% of
in different estuaries but subsequently dying the worm N. virens population biomass,
of starvation disproportionally in estuaries whereas in the Humber estuary, United
where food is scarce. Kingdom, birds consume only 23% of benthic
The relationship between bird usage of an production. Clearly estimates of bird predation
estuary and prey density has been developed vary, but a value of approximately 20% of ben-
for five species of Wader on the Forth estuary, thic production being consumed by birds can
Scotland. Here there is a significant association be considered as an average value.
Table 5.6 Correlations for wader feeding-hours and wader feeding densities in kilometer squared in relation to
invertebrate prey densities and site characteristics in the Forth estuary
Macoma X0 X X X0 X X
Cardium X0 X X X X
Mytilus X0 X
Hydrobia X X0 X0 X
Corophium X X
Nereis X0 X X0 X0
Nephtys X
Oligochaete X
Area 0 0 0
Inaccessibility Index 0
Coverage sequence 0 0
Note: Zero indicates a significant association in multiple regression analysis, and X indicates items found to occur frequently in the
diet in the analysis of guts.
Source: Bryant (1998).
Table 5.7 Importance of different invertebrates in the diet of shorebirds feeding on seal sands, Tees estuary, England
Shorebird Shelduck Grey plover Curlew Bar-tailed Redshank Knot Dunlin Energy content
godwit (invert. kcal x 106)
T. P. N. L. T. C. C.
tadorna squatarola arquata lapponica totanus canutus alpina
Invertebrate prey
H. ulvae
1 ) year class * ** * ** 9.0
01 year class * * * 3.0
N. diversicolor
1 ) year Class ** ** ** * * * 9.4
01 year class * 4.0
M. balthica
All classes * * * 0.6
Small polychaetes ** * ** 85.5
Small Oligochaete ** * * ** 222.0
Total energy content 333.5
of inverts.
Food requirements 73 2 23 9 17 93 68
of birds (kcal ' 106)
Total food 285.0
requirements of
birds (kcal ' 106)
Note: *Indicates regular food items; **Indicates the chief food providing daily biomass; indicates food item not taken at all or
only occasionally. The total energy content of the invertebrates and the total food requirements of the birds are expressed as millions
of kilocaloroie.
Source: Evans et al. (1979).
88 T H E E S T UA R I N E E C O S YS T E M
Birds, which feed intertidally in estuaries The gastropod snail, H. ulvae, is a major
during daylight hours, may find conditions food source for wintering Shelduck (T. tadorna),
particularly severe during the wintertime at and it is probable that most Shelduck
high latitudes. In a study of waders in the concentrations, at least in Britain, occur
Wash, England, it was shown that, during where Hydrobia is abundant and available. The
the autumn, feeding conditions for the birds feeding activity of Shelduck can be classified
were adequate and over 30% of the available into five methods, as shown in Fig. 5.6. These
daylight hours were spent in non-feeding methods are:
activities such as resting. During the short
1. Surface digging for Hydrobia buried in mud.
daylight days of the winter, however, they had
2. Scything action on exposed mud with a
to spend over 95% of the available hours
moist surface.
feeding and even that may have been insuffi-
3. Dabbling and scything in shallow water.
cient to sustain many individuals who appar-
4. Head dipping in 1025-cm deep water.
ently died due to food shortage. During the
5. Upending in 2540-cm deep water.
winter the Redshank on the Ythan estuary,
Scotland, are only able to obtain 50% of their Hydrobia displays a clear tidal rhythm, with
food from the estuary in daylight, and have the majority of individuals burrowing into the
to collect the balance at night or from the mud surface during low tide. As the tide reim-
surrounding land. merses the mud, the snails reemerge onto the
Eider ducks (S. m. mollissima) feed by mud surface to commence their grazing
dabbling in the shallow water at the edge of behavior. By adopting the head dipping and
the ebbing tide, or by diving into deeper upending modes of feeding, the Shelduck
water. The mussels they pick up are swal- are able to catch the emerging snails, and to
lowed whole and crushed by the birds large extend their time of feeding. Even at high tide
muscular gizzards. Such a feeding method many Shelduck manage to continue feeding,
precludes the intake of mussels larger than especially by upending. The scything action of
about 40 mm, and mussels less than 5 mm are the Shelduck is an effective means of sifting
rarely taken. out Hydrobia and other surface-dwelling
Scything Dabbling
and and Head
digging scything dipping Upending
100
% Hydrobia
80 Figure 5.6 Shelduck feeding methods,
on mud surface
showing the relationship between the
60 different methods used and the water
0 10 20 30 40 50
depth (cm), and the proportion of the
40 H. ulvae population at the mud surface.
Exposed mud Inundated muddepth of water (cm) (After Bryant and Leng 1976.)
T H E S E C O N D A RY C O N S U M E R S 89
animals such as Oligochaete from the mud Table 5.8 Productivity of selected secondary consumers
surface, but its value at particular locations
may depend on the moistness of the mud at Secondary Productivity Location
consumer (gC m!2 year!1)
low tide.
Estuarine birds 0.06 Dublin Bay, Ireland
Killifish 17.3 Tidal creek,
5.5 The impact of the secondary
(F. heteroclitus) United States
consumers Flounder 0.5 Ythan estuary,
The secondary consumers as a group often (P. flesus) Scotland
efficiently utilize the food that is available to Cape silverside 1.7 Bot estuary,
them, and also partition it between themselves (Atherina beviceps) South Africa
Clipfish (Clinus 0.05 Bot estuary,
in response to the pressures of ecological
spatulatus) South Africa
competition. As an example of this partitioning
of resources three bird species exploit the same Source: Wilson (2002).
mussel beds of the Ythan estuary with herring
gulls taking the 0-group mussels, eider the
I-group, and Oystercatchers the II)-group. and weather conditions may limit the time
Of the summer production of 1300 kcal m!2 that can be spent feeding. Third, much of the
by the mussels, 112 kcal are consumed by production of the primary consumers will be
the herring gulls, 275 kcal are consumed utilized after death by the decomposer organ-
by the eider ducks and 93 kcal are consumed isms, and through decomposition the organic
by the Oystercatchers. Man also removes matter becomes available again to the detritus
240-kcal m!2, and the remaining 580 kcal m!2 feeders.
is utilized by the mussels to support their The clear impact of predators of the inter-
metabolism in winter. Thus it may be seen in tidal macrofauna of muddy areas can be
this population that the requirements of the demonstrated by placing cages on the mud,
predators and the metabolic requirements of which exclude the predators and then observ-
the mussels are all provided by the production ing any changes within the cages. In predator
of the mussels. exclusion experiments in the German parts of
We have already seen, however, that in the Wadden Sea, off the Island of Sylt, Reise
the mudflats of the Ythan estuary as a whole (1985) has shown a remarkable (up to 23 times)
the flounders and gobiid fishes consume three increase of abundance of macrofaunal animals
times the total amount consumed by the four within exclusion cages as compared to control
major bird consumers. In various studies it sites without cages. The muddy sediments
appears that the fish and invertebrates are of this area are inhabited principally by the
more important predators in terms of total small annelid worms, Tubficoides benedeni,
consumption than the more conspicuous birds Heteromastus filiformis and Pygospio elegans,
(Table 5.8). In some estuaries not all of the with the main predators in the area the shore
total primary consumer production is utilized crab C. maenas, the shrimp C. crangon, and gob-
by the secondary consumers. This apparent iid fish Pomatoschistus microps. Within cages of
failure to consume all the available production 1-mm mesh only a slight increase in the abun-
may be due to several reasons. First, the dance of macrofauna was noted in the March
seasonal movements of the predators due to June period, but in the June to October
to migration in and out of the estuaries, or period of study the numbers within the cage
across continents, means that they are not increased dramatically, while the numbers
always present to utilize the food available. outside the cage decreased substantially. The
Second, the limitations imposed by tidal bivalve spat within the cage grew substantially
90 T H E E S T UA R I N E E C O S YS T E M
in this period, whereas extensive mortality Europe. Their areas of summer residence show
occurred outside the cage. At the end of the an even greater seasonal variation in food
study period the number of macrofauna availability with a rich abundance in the sum-
within the cage were 86,475 m!2, with the mer and virtual absence of food in the winter.
numbers outside 3750 m!2. From a comparison The birds are thus driven to the estuaries of
of different mesh sizes, Reise was further able milder regions, where even though food may
to show that in this part of the Wadden Sea the not be at its maximal biomass, it is at least
predation of shore crabs, shrimps, and gobies available. The availability of prey species
is responsible for major changes in the inter- throughout the year is ultimately due to the
tidal macrofauna, and that the impact of birds dependence of the primary consumer on detri-
and larger fish is negligible. The results of such tus as their main food source since this, almost
cage studies should always be treated with uniquely amongst primary food sources, is
some caution, as the presence of the cage may available throughout the year, albeit at a some-
alter the environment. what reduced level in the wintertime.
The numbers of birds that forage on the
Dutch part of the Wadden Sea are at a max-
5.6 Role of the secondary consumers
imum in autumn through to early spring and
in the estuarine ecosystem
then low in late spring and summer. The
numbers of birds clearly fluctuates independ- We have seen that the primary consumers of
ently of food supply, as the biomass of the estuarine intertidal areas are heavily exploited
prey species is at a peak in July and lowest by the secondary consumers that visit the
in February. The factors, which control the sea- area. These visitors are primarily young fish,
son of peak numbers of birds must be sought little crabs, and shrimps. The adults often stay
elsewhere and relate to breeding migrations in deeper water, while their larvae or juveniles
and the availability of food in other ecosys- come close inshore, feeding first on the meio-
tems. The numbers of fish and invertebrate fauna, and later on the macrofauna. Biomass is
predators, especially plaice (P. platessa) and exported from the tidal flats in terms of
shore crabs (C. maenas) in the Wadden Sea are
closely linked to the maximum availability of
their food species. As the biomass of prey Import Export
species reaches a peak in June through to
September so do the numbers of plaice and River
shore crabs. detritus Birds
In general, within the temperate mudflat- and plankton
dominated estuaries of Europe there is a sea-
sonal variation in the biomass of potential Land Intertidal
prey organisms with a maximum during the
production
summer months. The migration of fish and Sea
invertebrate predators into the estuaries is
often synchronized with this peak in the avail- Young fish,
Marine
ability of the prey, and when calculated on an detritus crabs,
annual basis the fish and invertebrates are and plankton and shrimp
often clearly the main secondary consumers.
Many estuaries particularly in Britain and The
Netherlands see peak numbers of birds in the
winter months, as the waders and wildfowl Figure 5.7 The tidal flat turntable for the flow of organic
migrate there from eastern and northern matter between the land and the sea. (After Reise 1985.)
T H E S E C O N D A RY C O N S U M E R S 91
Effect on abundance of prey
nurslings that have grown to adult size and healthy and sustainable an ecosystem must
return to the sea to grow and breed, and also maintain its metabolic activity level, must
to a generally lesser extent by birds, which maintain its internal structure and organiza-
feed on the same areas at low tide, and then fly tion, and must be resistant to external stress
off to the land at high tide. Reise (1985) in over time and space scales relevant to the
describing this situation uses the term an ecosystem. The ecosystem can thus be exam-
ecological turntable (see Fig. 5.7). Energy is ined on the basis of its productivity, its fish
received by intertidal flats from land and the and fisheries, its health and its socioeco-
sea as imported nutrients and detritus, and nomic values.
exported from the same area back to the land On an economic basis the estuarine eco-
and sea by migrating birds, juvenile fish, and system provides many valuable services,
as dissolved nutrients. The various predator which are often not traded in the stock mar-
effects on the prey populations are summa- kets of the world, but which are essential and
rized in a speculative manner in Fig. 5.8. The valuable services. These include the services
estuarine ecosystem is as we have already provided by estuaries and coastal wetlands as
seen heavily subsidized by the adjacent nurseries for fisheries, as natural pollution fil-
ecosystems, but through the activities of the ters, and as storm buffers. Even though the
secondary consumers at least some of this estuaries are the recipients, from rivers and
energy is returned to support the marine and the land, of most of the pollution that subse-
terrestrial ecosystems. quently enters the sea, by their ecosystem
Sherman and Duda (1999) have recently activity they manage to reduce the amount
reviewed the ecosystem approach to the entering the sea, while at the same time pro-
global assessment and management of coastal viding essential feeding grounds for juvenile
waters, and show how worldwide the ecosys- fish and migrating birds. Estuaries are a small
tem approach is developing as a means of proportion of the sea as measured by surface
linking sciencebased assessments to the area or water volume, but it is clear that the
management of these resources. They warn, ecosystem functioning of the estuaries that
however, that management cannot wait for they provide an essential service to the ecosys-
science to catch up with a full understanding tem of the sea, far out of all proportion to their
of ecosystem structure and function. To be apparent size.
CHAPTER 6
92
E S T UA R I N E U S E S A N D U S E R S 93
Driving forces: the Drivers, the human activities chemistry of the system, the biological responses
responsiblethis refers to the big aspects, for at any level of biological organization (cell,
example, the need for food such as fish and individual, population, community, ecosystem),
shellfish, the need for moving materials, the need at any spatial scale (i.e. extent) or temporal scale
for building materials, the use of space, the need (i.e. duration);
to dispose of waste; Response: the human response to the
Pressures: the causes of problems derived causes and effects, the derivation and
from the Drivers; each Driver will create many measurement of policy options, the need to
pressures, for example, emissions of waste, ensure that the six tenets (environmentally
seabed damage through fishing, oil spills from sustainable, economically viable,
tanker movements; technologically feasible, legislatively
State: the change in the environment, the permissible, administratively achievable, socially
background status, structure and functioning, desirable) are met for a sustainable management
environmental variables and physical, chemical, policy, the use of BPEO (Best Practicable
and biological features of an area or system; Environmental Option), EIA (Environmental
Impact: the changes to the human system, the Impact Assessment), BAT (Best Available
Man-induced modification to the physics and Technologies).
human activities in the estuarine ecosystem: mans actions as having a pathology and so we
can define and study that change as attributes
the presence of xenobiotics (i.e. materials
for the diagnosis of ecosystem pathology
foreign to the natural system) and toxins
(Harding 1992; Elliott and Hemingway
physical additions
2002). The predominant changes to the system
energy change
relate to:
physical structures
Pollutant A substance introduced into the natural environment as a The input and effects of micro- and macro-
result of mans activities and in quantities sufficient to organisms to produce adverse effects.
produce undesirable effects.
Pollution (i) is a change in the natural system as a result of mans The effects of introduced, invasive species sufficient
activities; (ii) has occurred if it reduces the to disturb an individual (e.g. internal biological
individuals/populations/species/communitys fitness to pollution by parasites or pathogens), a population
survive. The introduction by man, directly or indirectly, of (by genetical change) or a community (by increasing
substances or energy into the environment resulting in or decreasing the species complement); including
such deleterious effects as to harm living resources, the production of adverse economic consequences.
hazards to human health, hindrance to marine activities
including fishing, impairment of quality for use of
seawater, and reduction of amenities (GESAMP).
Contamination An increase in the level of a compound/element The introduction of species without noticeable
(pollutant) (as the result of Mans activities) in an effects (e.g. microbes killed in seawater; species
organism or system which not necessarily results in a occupying available and vacant niches).
change to the functioning of that system or organism.
Responses to (i) lethalorganisms are killed thus resulting in (i) the introduction of pests which increase
pollution community change; (ii) sub-lethaleffects which may predation/mortality; (ii) an increase in an
occur before the concentration of toxic substances immunological response in an individual due to
reaches lethal levels. exposure to microbes.
Stress The cumulative quantifiable result of adverse A reduction in health due to pathogens and
environmental conditions or factors as an alteration in parasites and the loss of genetic fitness due to
the state of an individual (or population or community) escapees. The alteration in community structure
which renders it less fit for survival. with single invasive species at low densities may be
difficult to detect.
Episodic pollution Major but often short-lived (temporary) discharges, e.g. Discharges of organisms via ballast-water may be
shipping accidents. regarded as episodic although there may be an
insufficient influx to establish a population.
Chronic pollution Diffuse, low-level inputs which cannot be traced to a Regular, continuous inputs of organisms as escapees
particular incident, e.g. from rivers, atmosphere; may or the dispersive, reproductive stages of non-native
contribute 90% of total inputs; often the cause of effects cultured organisms that have adapted successfully
which are sub-lethal and difficult to detect. and can reproduce; gradual movement of species
through climate change.
Acute pollution Concentrated pollution, with an identifiable source and Pathogenic micro-organisms discharged from
readily observed effects; often the cause of lethal effects. waste-water outfalls.
Aesthetic pollution Unpleasant material likely to cause visual or olfactory The aesthetic aspects of changes to natural faunal
(sic) offence but which (usually) causes little biological harm. and floral communities, i.e. a reduction in
naturalness.
Bioaccumulation An increase with time in the content (or body burden) The uptake and accumulation (culture) of
and/or concentration of a contaminant within an pathogens (and/or cysts) in filter-feeding bivalves
organism. may occur. The accumulation after successful
reproduction of invasive species in a community.
Biomagnification Increasing levels of a pollutant from the lower trophic The successful establishment (and displacement of
strata to the higher trophic strata. indigenous species) of an invasive population.
Direct input A point source discharge, often the cause of acute The release of pathogens from waste-water
pollution. discharges, of genetically-modified organisms from
culture, or parasites from transplanted shellfish.
Indirect (or diffuse) A widespread, low-level discharge often likely to result in The changes in distribution of species through
input chronic pollution. man-induced global climate change.
Conservative Materials with slow degradation rates (long half-life), The accumulation in the marine environment of
wastes/inputs more likely to bioaccumulate, e.g. trace metals, successfully-colonising organisms, to the extent that
halogenated hydrocarbons, and radioactivity. their population becomes established and increases.
Degradable Materials more liable to be assimilated after being The demise and degradation of micro-organisms
wastes/inputs degraded through physical, biological and chemical (e.g. faecal coliforms) in seawater after liberation
processes in the short to medium term, e.g. sewage, from waste-waters. The unsuccessful colonisation of
hydrocarbons, pulp/paper wastes. introduced species.
E S T UA R I N E U S E S A N D U S E R S 97
feature in the public domain, despite the fact City have been dumped near Ambrose Light.
that over long timescales, more oil could enter The bottom water oxygen concentrations are
the estuary from the industrial discharge than depressed at and around the dumpsite, and in
is spilt from a stricken tanker. spring and summer 1976 an area of 12,000 km2
experienced oxygen concentrations less than
2 mg l!1, with a smaller area becoming com-
6.3 Sewage pollution
pletely anoxic, resulting in the death of at least
Man has long used estuaries for the disposal 143,000 tonnes of the clam Spisula. Reporting
of waste material and sewage is discharged the situation in 19802, Steimle (1985) has
into many estuaries. In many cases the raw shown that there is not an azoic biological
untreated sewage is discharged, and in other desert in the dumping area, but the benthic
cases the sewage is treated on land in sewage fauna showed enhanced production of the
treatment works and only the liquid effluent stress-tolerant species, leading to an enhanced
produced is discharged into the estuary. benthic biomass (127344 g m!2 wet wt) with
The waste so discharged may, if there is only a productive demersal fisheries.
little, become incorporated into the estuarine Prior to the banning of seaborne sewage
ecosystem as another source of detritus. The sludge disposal in 1998, the dumpsite for the
quantities discharged may, however, be so sewage of the city of Glasgow was at Garroch
great as to cause major changes to the fauna Head, where 1.5 million tonnes of sewage
and flora. The solid waste from sewage works, sludge were dumped annually. This was a typi-
known as sewage sludge, has to be disposed cal accumulating site, and the biomass and
of, and several countries including the United abundance of the fauna was very high at the
States have allowed the disposal of such mate- center of the dump site (being 500 g m!2 and
rial to sea or estuary. Prior to 1998, the United about 500,000 individuals m!2) although the
Kingdom and Ireland dumped sewage sludge species diversity was low with the principal
into the sea but this was then banned in species being nematodes (Pontonema spp.),
Europe under the European Commissions the Oligochaete Tubificoides benedeni and the
Urban Waste Water Treatment Directive and polychaetes Capitella capitata and Scololepis
other international conventions. fuliginosa (Pearson et al. 1986). The species
Whether sewage works provide primary diversity increased away from the dumping
treatment (the removal of solids only) or sec- site, while the biomass and abundance declined
ondary treatment (full biological treatment of (Fig. 6.2). These changes in species diversity,
liquid waste as well), considerable volumes of abundance, and biomass conform well to the
solid waste, sewage sludge are produced. The general pattern described below, termed the
coastal area was long regarded as the most eco- Pearson Rosenberg model. It was of note that
nomical method of disposing of this sludge by this sludge, from a highly industrialized city,
transporting the waste by ship from the also contained high levels of persistent pollu-
sewage-works to a dumping site at the mouth tants such as heavy metals that then were
of the estuary, or further offshore. Two types of retained in the sediments and the fauna.
dumping site can be distinguished, either accu- Whereas many of these pollutants were retained
mulation sites where the material accumulates at the disposal site, some were exported in
on the bottom, or dispersal sites where currents migrating fish feeding on the high levels of
disperse the material over a wide area, hence invertebrates at the site. Despite all of this, the
using the ability of the sea to disperse, degrade, site was regarded as a marine land-fill site.
and assimilate the waste. For example, a major In contrast to the Garroch Head site, the
sludge-dumping site is in New York Bight, sewage from the city of Edinburgh (up to
where large quantities of waste from New York 500,000 tonnes annually) prior to 1998 was
98 T H E E S T UA R I N E E C O S YS T E M
estuaries bordering it. These nutrients are (4) oxygen depletion resulting from the die-
derived from inland sources, and carried from back of the increased plant growth, which kills
the rivers, through the estuaries into the sea. fish and benthos;
Without doubt the biggest source of such (5) organically enriched sediments following
nutrients is the river Rhine. the dieback of the plant material and changes
The addition of extra organic matter (carbon) to the benthic community.
to estuaries and the sea is generally known as
organic enrichment, and the increase of nutri- Rosenberg (1984, 1985) showed that eutro-
ents (nitrogen, phosphorus) from the same phication has been known to be occurring in
source is known as hypernutrification. Hyper- the Baltic Sea for some time, as a consequence
nutrification can lead to a set of adverse effects of the large nutrient inputs to the area coupled
under the right environmental conditions and with the long residence time of this brackish
those adverse effects are known as eutrophica- sea. During the 1980s, the symptoms of
tion. Eutrophication may be natural or from eutrophication were also found in the
mankind, that is, anthropogenic. In fresh- Kattegat, the shallow sea between Denmark
waters it is the supply of phosphates that may and Sweden, with marked changes in the
cause eutrophication, but in estuaries and the bottom fauna observed, and mass mortality of
sea it is usually the supply of nitrogen, which bottom-dwelling animals such as Nephrops
is the critical factor. De Jonge and Elliott (2001) norvegicus (scampi, or Norway lobster). The
and Elliott and de Jonge (2002) indicate the demersal fisheries (for cod, etc.) declined,
causes and consequences of nutrient enrich- but the pelagic fisheries (for herring, etc.)
ment as a set of bottom-up causes and top- increased.
down responses. High inputs of nutrients The phytoplanktonic blooms may be toxic
require optimal conditions of a long residence to other organisms. In 1987 such a red tide
time and good light conditions in order to bloom of toxic algae forced the closure
be used for plant growth. In usually turbid of 170 miles (250 km) of clam and oyster
estuaries, the nutrients may occur in elevated beds in North Carolina. Similarly, in the late
levels but do not generally lead to excessive 1990s, blooms of these organisms in west
algal growth because of light limitation. Scottish coast sea-lochs were thought to
If there are good conditions for the use of the be associated with increased nutrients
nutrients then the symptoms of eutrophication and organic matter from fish farming.
can occur. These are: Eutrophication has now been recorded in
many estuaries of the world; as an example,
(1) increased macroalgal growth as macro- the steady increase of phytoplankton in the
algal mats such as Enteromorpha, which may be EmsDollard estuary in response to eutrophi-
floating in shallow brackish waters, as along cation, measured here as total phosphates, is
the western Swedish coast, or on intertidal shown in Fig. 6.3.
flats such as the Tees estuary, NE England; On the intertidal mudflats of many estuaries,
(2) blooms of planktonic microalgae, which for example, the Ythan estuary in NE Scotland
may be toxic, such as the red-tide algae and the Tees estuary in NE England, the
producing Amnesic, Paralytic, or Diarrhoeic growth of extensive algal mats is regarded
Shellfish Poisoning (ASP, PSP, DSP) such as as a clear symptom of eutrophication. Algal
Alexandrium and Cingula, or just a nuisance by mats of Enteromorpha or Cladophora smother
producing large amounts of mucilage and the primary consumers in the benthos, and
foam such as Phaeocystis; prevent oxygen reaching the sediments as
(3) decreased water transparency as the result well as increasing oxygen uptake by the
of the blooms; degradation of the organic matter. This
100 T H E E S T UA R I N E E C O S YS T E M
6.5 Organic enrichment problem. Figure 6.4 shows the water column
with dissolved oxygen levels in the upper Forth
A small amount of organic matter, well dis-
estuary and in particular illustrates both a sea-
persed, can be readily utilized within the estu-
sonal (i.e. summer) and spatial (i.e. upper estu-
arine ecosystem to enhance the levels of
arine) dissolved oxygen sag or minimum.
biological production. Problems arise, however,
The development of low oxygen, or even
if a large volume of organic effluent is dis-
anoxic, conditions leads to the extinction of the
charged into an estuary with a low flushing
normal macrofauna of the estuary, and its
rate, or if too much effluent is discharged at a
replacement by annelid worms, especially
single point, that is, the assimilative capacity of
Oligochaetes. In areas of greatest organic
the receiving area is exceeded. When excess
enrichment even such worms may be excluded.
quantities of organic matter are present, then
The microbial degradation of organic waste
bacteria and other microorganisms, which util-
will, apart from consuming the available
ize the organic matter, will consume all the
oxygen, also lead to increased sulfide produc-
available oxygen in the water. Such problems
tion. The oxidation of sulfide leads to further
may be particularly severe in summertime
oxygen depletion. The effects of organic enrich-
because of reduced river flow, shallow water
ment are most clearly seen by monitoring the
conditions, coupled with temperature increases,
fauna and measuring the redox potential (Eh)
which lead to enhanced bacterial activity,
of the sediment at successive distances away
and thus accelerate the depletion of the oxygen
from a known organic discharge. The redox
content of the water. The shallow nature of the
potential may be determined with special plat-
upper estuarine areas then exacerbates this
inum electrodes inserted into the sediment,
13
Mar.
11
Jan.
9 Feb. Dec.
DO (mg l1)
Apr.
Nov. May
7 Oct. Aug.
UE 1 Sep.
Jul. Jun.
UE 2
UE 3
5 UE 4
UE 5 Figure 6.4 Dissolved oxygen and
UE 6 temperature profiles of the Upper Forth
Estuary, Scotland, showing monthly averages.
(Forth River Purification Board, Edinburgh,
4 6 8 10 12 14 16 18 20 198890 data, UE1 in mid -estuary and UE6
Temperature ( C) at the tidal limit.)
102 T H E E S T UA R I N E E C O S YS T E M
with the resultant negative Eh values indicating and Rosenberg 1978). Such S-A-B curves are
reducing (anoxic) conditions, while oxygen- now regarded as typical for the effects of
ated sediments have positive Eh values. organic enrichment, and can be interpreted as
Pearson and Stanley (1979) give full details of follows. Close to the effluent, conditions are
the measurement of Eh. anoxic, and no macrofauna can live. As one
The results of the measurement of fauna moves away from the effluent the sediment is
and Eh at distances of up to 800 m from the colonized by opportunistic animals, mostly
discharge of organic effluent from an alginate small annelids, which occur in enormous
(seaweed) factory are presented in Fig. 6.5. numbers. This zone, the peak of the oppor-
Close to the effluent were relatively few tunists has low species diversity and modest
species, with low numbers and biomass, cou- biomass. Beyond this is the ecotone, where
pled with negative Eh values. At 100 m the the more normal fauna replaces the oppor-
abundance of animals (mainly small poly- tunists, and as shown at 200 m in Fig. 6.6, a
chaetes) increased dramatically, but there are decrease in the biomass may occur. Beyond
still few species. The biomass and Eh increased the ecotone, is the transition zone where the
more steadily. Between 100 and 400 m the total fauna shows enrichment due to the organic
numbers (abundance) of animals decreased, source, with maximal biomass and species
while the species diversity and Eh steadily diversity. Beyond this enriched zone the fauna
increased. The biomass showed a dip at declines to its background levels.
200 m, but otherwise increased. Beyond 400 m The change in species composition is shown
the species diversity, abundance, and biomass pictorially in Fig. 6.6. Water movement and
all fell, while the Eh continued to rise. Such renewal is an essential factor affecting the
patterns of change in the species diversity (S), impact of organic material in a particular area.
total abundance of animals (A), and total bio- The importance of water renewal is shown in
mass (B), have been recorded in the vicinity of Fig. 6.7. The importance of water renewal is
many organic discharges, and the form of the such that a given organic discharge into the
curves have become known as S-A-B curves well-mixed mouth of an estuary may have lit-
or the PearsonRosenberg model (Pearson tle or no measurable effect, whereas the same
discharge into the restricted confines of the
head of the estuary may produce severe
symptoms of organic enrichment.
60 Total The nature of estuarine and marine commu-
abundance (100) +400 nities under both polluted/stressed and clean/
Eh
+300 unstressed conditions can be listed accord-
40 Species ing to the changes in the primary community
number +200
Eh (mV)
Aerobic
sediment 2
Anaerobic
sediment 3
4
Zone Normal Transitory Polluted Grossly polluted
cm Depth in sediment
Typical Nucula Labidoplax Chaetozone Capitella No Macrofauna
macro- Amphiura Corbula Anaitides Scolelepis
Terebellides Goniada Pectinaria Surface covered by
fauna
Rhodine Thyasira Myriochele fiber blanket
domi-
nants Echinocardium Pholoe Ophiodramus
Nephrops
Figure 6.6 Diagram of the changes in fauna and sediment structure along a gradient of organic Enrichment. The species
listed are typical macrofauna species for north European marine waters and equivalent species will be found elsewhere.
(From Pearson and Rosenberg 1978.)
Input
Organic matter
Sedimentation
High organic
content in sediment
Bacterial
decomposition
Well
flushed Stagnation
Moderate water
Oxidized sediment re- newal Anoxic sediment
with plentiful Reduced O2 Emerging Eh and overlaying
food supply in water in sediment water
Figure 6.7 Diagram to show some pathways of organic input to the marine and estuarine environment,
and its effects in relation to water renewal. (From Pearson and Rosenberg 1978.)
104 T H E E S T UA R I N E E C O S YS T E M
Table 6.2 Conceptual basis and assumptions inherent in macrobenthic impact studies. From Elliott (1994) modified and
expanded from McManus and Pauly (1990)
A. Natural state
A natural macrobenthic assemblage either tends toward or is in an equilibrium state;
Under non-impacted conditions, there are well-defined correlation and relationships (which therefore may be modeled)
between faunal and environmental (abiotic) variables;
In approaching the normal equilibrium state, the biomass becomes dominated by a few species characterized by low
abundance but large individual size and weight;
Numerical dominance is of species with moderately small individuals, this produces among the species a more even
distribution of abundance than biomass;
The species are predominantly K-selected strategists;
B. Moderate pollution
With moderate pollution (stress), the larger (biomass) dominants are eliminated, thus producing a greater similarity in
evenness in terms of abundance and biomass;
Also, with moderate pollution, diversity may increase temporarily through the influx of transition species;
C. Severe pollution
Under severe pollution or disturbance, communities become numerically dominated by a few species
with very small individuals;
Those small individuals are often of opportunist, pollution-tolerant species, which have r-selected strategies;
Under severe pollution, any large species that remain will contribute proportionally more to the total biomass relative to
their abundance than will the numerical dominants;
Thus under severe pollution, the biomass may be more evenly distributed among species than is abundance;
However, under severe pollution, species with large individuals may be so rare as to be not taken with normal sampling;
The change in assemblage structure with increasing disturbance is predictable, follows the conceptual models and is
amenable to modeling and significance testing;
D. Recovery
Opportunists are inherently poor competitors and may thus be out-competed by transition species and K-strategists if
conditions improve;
McManus and Pauly (1990) also consider that under normal conditions:
The biomass-dominants will approach a state of equilibrium with available resources;
The smaller species are out of equilibrium with available resources;
The abundances of the smaller species are subject to more stochastically controlled variation than the larger species.
fishes of the North Sea (Fig. 6.8). Organic pol- One estuary, which has recovered dramat-
lution of the Wadden Sea reflects the distribu- ically from decades of organic pollution, is
tion of populations and industry, with the Thames estuary, which flows through
untreated or minimally treated effluent from London, United Kingdom. Due mainly to mas-
dairies and slaughterhouses in the Danish sec- sive amounts of domestic sewage the state of
tor, effluent from the estuaries of the Elbe and the Thames declined steadily during the first
Weser in the German sector, and from major half of the twentieth century, until by the
Dutch cities. Viewing the Wadden Sea as a 1950s it was completely anoxic over the mid-
whole the organic effluents cause severe local dle reaches, inhabited only by Oligochaete
problems of oxygen deficiency, but on a gross worms, and mallard and swans, which fed on
scale the area remains as probably the most grain spillages. Conditions only began to
important and productive estuarine ecosys- improve in 1964 with the completion of a new
tem in Europe. sewage works at Crossness, and improved
E S T UA R I N E U S E S A N D U S E R S 105
Denmark
Esbjerg
Skallingen
N Fan
Mand
Rm
Sylt
0 km 100 Fhr
Amrum
SchleswigHolstein
Pellworm
Wangerooge
Spiekeroog
Schiermonnikog
Norderney
Langeoog
Elb
Brokum
Treschelling
e
Ameland
Just
Weser
Cuxhaven
Jade
Vlieland
Ost Bremer-
Ems
Friesland haven
e n
Texel
Emden v
Delfzijl a
sh Figure 6.8 Map of the Wadden Sea.
Harlingen m Bremen
el Stretching from Texel in The Netherlands
li h
W Niedersachsen to Esbjerg in Denmark this is the largest
Den Helder
The Netherlands Federal Republic of Germany estuarine area in Europe. (After Essink
1978.)
further in 1974 due to another sewage works numbers of wading birds and other ducks
at Beckton, coupled with the closure of several have increased dramatically in response to the
old discharges. As a result of these improve- return of a wide diversity of prey organisms.
ments the condition of the estuary water has
steadily improved, and by 1975 the lowest
oxygen content was 25% of saturation. Since 6.7 Fisheries and aquaculture
then the water quality has steadily improved,
6.7.1 Wild fisheries
and as it has done so, fish and birds have
returned to the estuary. Figure 6.9 records the Worldwide, the main estuarine fisheries are
number of fish species caught during the for shellfish, with the collection of some of the
recovery period. In the period 192060 no fish abundant natural populations of inverte-
were caught. The eel was the first to return, brates, such as shrimps, crabs, oysters, cockles
and by 1975, 36 species were found, including and mussels. Increasingly the catching of nat-
most dramatically the return of salmon, Salmo ural stocks is supplemented by mariculture,
salar. The number of fish species has increased most notably for oysters and mussels. Bait
gradually with, by 2002, over 120 species digging by sport fishermen occurs widely in
being found. Similarly, a wide diversity of estuaries, and may involve considerable dis-
invertebrate fauna now occurs, although there ruption of the intertidal fauna.
are fewer Oligochaetes. As the Oligochaetes In addition to the shellfish industries are the
have reduced so have the numbers of tufted vertebrate fisheries. While many of the fish,
duck and pochard, which fed upon them. The which enter estuaries are not commercially
106 T H E E S T UA R I N E E C O S YS T E M
100
Cumulative species nos.
80
60
40
20
0
1964
1966
1968
1970
1972
1974
1976
1978
1980
1982
1984
1986
1988
1990
1992
1994
1996
1998
2000
2002
Year
Figure 6.9 Number of fish species recorded in the Thames estuary, England.
Drawn from data supplied by Dr. S. Colclough, Environment Agency, London.
exploited because they are the nursery stocks, the activities, these have been summarized in
or unwanted species, other species are heavily Fig. 6.10. For example, as well as the removal
exploited. Salmon, sea trout, and eels all pass of target and nontarget fish and shellfish, fish-
through estuaries on route from the sea to ing activity in estuaries can disturb or destroy
rivers, and many commercial fisheries exploit benthos and thus affect their dependent fish
them. The Gulf of Mexico, stretching from and bird populations (Jennings and Kaiser
Florida to Texas and Mexico, is a key fishery 1998; Hall 1999; Blaber et al. 2000). The main
area for the United States, with 28% of their effects of fishing are:
total fish landings coming from the brackish
bays and lagoons of this area. The catch is (1) to cause local extinctions, where there is a
mainly menhaden (Brevoortia spp.), striped large-scale removal of a particular species,
mullet (Mugil cephalus), croaker (Micropogon often as the target species, for example, the
undulatus) plus Penaeid shrimps, blue crab, smelt, Osmerus eperlanus, which disappeared
and oysters. All except the croaker are estuar- from the Forth Estuary, Scotland in the middle
ine species, which in general spawn at sea. of the twentieth century mostly as the result of
The larvae enter estuaries, grow there, and over-fishing;
when fully mature return to the sea. Other rich (2) to affect the population viability, genetics,
estuarine fisheries are at the mouth of the and maturity of the target organisms, for
Amazon, in Nigerian estuaries, and in Indian example, the removal of spawning stocks
estuaries, such as the Ganges, where estuarine through over-fishing of pelagic and demersal
conditions extend 160 km upstream from the populations;
mouth. (3) to affect nontarget organisms, both other
As with all types of fishing, there are now fishes, such as the taking of juvenile herring
major concerns on the ecosystem effects of during estuarine fishing for sprat, and other
Fisheries
effects on:
Figure 6.10 The effects of fisheries on estuarine ecosystems. (From Elliott and Hemingway 2002.)
108 T H E E S T UA R I N E E C O S YS T E M
and dab during shrimp fishing, and by remov- population changes at the ecological and
ing the mudflat habitat used by juveniles; genetic levels, and the effects on competing
(5) to affect trophic interactions such as the predators)
removal of prey fishes, such as sandeels taken the physical integrity of the system (through
by seabirds and crustaceans, or the shrimps, bed disturbance and damage to the benthos)
which are central to the functioning of many contamination of the estuarine system
estuarine food webs, and by changing food (through discharges of soluble pollutants and
webs by increasing scavengers, especially ben- large and small particulate materials, includ-
thic megafauna, following seabed damage by ing litter and gear loss)
fishing gear and the production of detrital the creation of infrastructure (habitat loss,
Community
Increase in structure Pollution Damage and
Discards
scavengers Selective change ghost fishing
mortality
(diversity)
Regrowth Lethal and
Increase in
Benthic Habitat integrity sublethal
scavengers (birds
anoxia Density effects Habitat
and fish)
reduction loss
Juvenile flatfish
(e.g. shrimp catch) Opportunistic Seabird and
Energetics
species seal prey Biomass
change
loss export
Figure 6.11 The effects of trawl fishing on the estuarine ecosystem. (From Elliott and Hemingway 2002.)
110 T H E E S T UA R I N E E C O S YS T E M
Figure 6.12 A floating sea-cage fish-farm for salmonid fish (salmon or trout). Such farms have become common sights
on the fjordic estuaries of Norway, Ireland, Canada, New Zealand, Scotland, and elsewhere. (See also Fig. 6.13.)
organic matter (DOM), excess food, feces, lead to the development of opportunist
mucus, and detritus). The wastes will add populations and thus benthic prey quality
nutrients thus having the potential to change palatability has been changed for fish and
oligotrophic (nutrient-limited) systems to megafaunal predators. Brown et al. (1987)
more nutrient rich ones. Naturally nutrient- have shown that underneath the floating
rich systems (hypernutrified) can become salmon cages the sediment is highly reducing
eutrophic. There may be a reduction of (negative Eh) and azoic. A highly enriched
Dissolved Oxygen (by fin/shell-fish demand zone, dominated by the opportunistic and
and the Biochemical Oxygen Demand of the pollution tolerant polychaetes C. capitata and
wastes) and the potential for red-tides (nox- S. fuliginosa, occurred from the edge of the
ious microalgae blooms) has been suggested cages out to approximately 8 m. A slightly
for the Scottish West Coast. There will be a enriched zone occurred at up to 25 m, beyond
hydrographic disruption by the structures which the fauna was unaffected by the cages.
(they will impede water flow, especially where Thus the ecological effects of this form of
there is excessive weed attachment) and the organic enrichment are severe but limited.
inputs from the caged organisms will lead to In the case of shellfish, farming of mussels
microbial change through and input of fecal has expanded in Spain, New Zealand, and
microorganism input. Finally, there may be Ireland. The mussels are grown on suspended
an increase in suspended solids hence an ropes, feeding on natural planktonic material.
increased turbidity, leading to potential for a In studies of mussel farming in the estuarine
reduction in primary productivity. Many of Ria de Arosa of northwest Spain, Tenore et al.
these effects fall within the generally acknowl- (1982, 1985) have shown that mussels are
edged symptoms of eutrophication. a key species in determining ecosystem
Arguably the greatest impact will be in structure and dynamics of the whole area. By
changes to the sediment and benthos. There intensive mariculture, man has replaced the
will be an accumulation of solid wastes on zooplankton with mussels as the dominant
the bed given that 510% of food is wastage herbivores in the area. The major changes are:
and 2530% of the food weight will be excreted
1. The surface area of, and detritus from, the
as feces. In the case of bivalve shellfish there
mussels support a dense epifaunal commun-
will be the biodeposition of sediments through
ity, which utilizes 90% of the mussel feces, and
pseudofeces production, hence changing the
supplies food to demersal fish and crabs.
nature of underlying sediments. The hydro-
2. Epifaunal larvae, rather than copepods,
dynamic change because of impeded water
dominate the zooplankton community.
flow can lead to sediment structure change
3. Nutrient cycling by mussels dampens phyto-
through the production of a low energy area.
plankton oscillations and contributes to high
The organic input and enrichment of sediment
seaweed production on ropes.
will produce anaerobic sediments, which may
4. Heavy sedimentation of mussel deposits
ultimately affect the water quality through the
changes the sediment regime and lowers
production of hydrogen sulfide (H2S) and
infauna production.
methane (CH4), which in turn are toxic. Anoxic
5. Transport of particulate organics derived
sediments will affect the water column if the
from mussel deposits from the farming areas
Redox Potential Discontinuity, as the boundary
enhances benthic biomass outside this area,
between the upper oxygenated levels and the
and might support near-shore fisheries, espe-
lower anaerobic levels, migrates out of the
cially for lake.
sediments. This is then reflected by the devel-
opment of Beggiatoa mats utilizing the hydro- Tenore concluded that the raft culture of
gen sulfide. In turn, the anaerobic conditions mussels affected food-chain patterns and
112 T H E E S T UA R I N E E C O S YS T E M
production in generally positive ways, although and the introduction of exotic species. In
admitting that the infauna benthos near the cases where fishes such as salmonids have
farms was typical of polluted conditions. been reared especially for culture, escapees
Considering this study along with several from the genetic monoculture will produce
others, we can summarize in Fig. 6.14 the pos- an alteration in the genetic makeup and a
sible effects of mussel farming on the estuarine possible reduction in genetic fitness of the
ecosystem. While mussel rafts or ropes reroute natural populations. There may be new and
the flow of energy or materials, they do not add increased disease introduction as well as para-
any extra nutrients to the ecosystem, unlike site introduction/concentration. Finally, the
caged finfish fed on prepared food, and any removal of locally or distant caught marine
changes involve processes different from fishes for fish meal for feeding the farmed
eutrophication caused by an enhanced nutrient fishes will impact on those stocks.
supply. Given the importance of these areas for
Both finfish and shellfish culture will pro- birds, aquaculture has the potential for
duce impacts on the indigenous fishes impacts on this use. There may be acute and
through the alteration of normal habitats, chronic disturbance of breeding, feeding, and
impeding of water flow, and disruption of overwintering areas and the disruption of
migration routes. There is the introduction the site (through the loss of land and water
and/or displacement of nonnative stocks usage) as well as an influx of opportunist birds
Release of nutrients,
particularly NH3. Provides attachment +
Increased cycling food for large epifaunal
Decreases phytoplankton community
biomass + changes
composition by filter feeding
Mussel
Increase in epifaunal
raft
larvae
Dampens phytoplankton
oscillations
Competition with other
natural filter feeders.
Increased organic
Reduction in copepod
loading to sediment
zooplankton community
Reduces Decreased
phytoplankton community Increased bacterial
productivity diversity and activity Bedload transport
biomass enriches other areas
Possibility of anaerobic
conditions. Increased benthos and
Reduces Reduced
demersal fish in other
release of working of
areas
nutrients sediment
Production of H2S + CH4.
Figure 6.14 Summary of the possible effects of mussel farming. Note that some of the effects are contradictory,
and not all effects will be seen at one site. (From Gowen et al. 1988.)
E S T UA R I N E U S E S A N D U S E R S 113
feeding. The activities can lead to mortalities using low-wastage feed and hand-feeding,
through the control by shooting (which coupled with the type of food (slow sinking,
requires an exemption to bird protection Acts and with a balance controlled to reduce NH3
in certain countries) and there may be accid- input) will reduce the effects. The use of good
ental or deliberate trapping of birds (through practice such as the removal and destruction of
the use of anti-predator nets etc.). These mortalities, rather than into the water column,
adverse effects may be minimized by site con- and the use of biological control for pests
siderations, gear improvement, and the use of (e.g. lice control using goldsinny wrasse) will
scaring devices or sacrificial food, and the minimize biological effects. In addition, fish-
adoption of a policy on shooting. farmers and environmental protection agencies
Aquaculture requires the addition of chem- determine the potential effects using numerical
icals and these may be given as enteric modeling (based on volume of stock, water
treatments (by mouth, e.g., with the fishfood) flow, depth), coupled with self-monitoring for
or by immersion treatments. These include the health of sediments and benthos. Sampling
food additives such as vitamins, mineral and/or photography then achieve the assess-
mixes, and pigments, the latter are required in ment. Finally, the use of polyculture, as a mix-
the case of salmonids in order to produce ture of fin- and shell-fish farming, or the use of
pink flesh, which is acquired in the wild by ranching as compared to cages can be a wiser
eating a crustacean diet not used in farming. use of the environment.
Anaesthetics and narcotizing agents are
required to minimize stress of the fish during
6.8 Industrial contamination
handling and vaccines and other therapeutants
are administered for health. Disinfectants and Waste from industrial sources enters estuaries
antibiotics will be used to minimize microbial from industries located on the banks of the
effects and, given the increased possibility of estuary as well as that from inland sites may
external parasite transfer when large numbers be discharged into rivers or public sewers,
of fishes are confined, pesticides (such as which subsequently arrive in estuaries. The
Nuvan/Aquaguard (dichlorvos based) and waste from industrial sources may be treated
ivermectin) are used to combat infestation before discharge, so that the effluent arriving
of fish lice. Finally, antifouling agents/ in the estuary has a considerably reduced
treatments (such as Cu, TBT, bitumen, and slip- concentration of waste materials, or the waste
paints) may be used to control macrofouling of may be discharged in an untreated form. All
the gear. industries produce some amount of waste,
Aquaculture thus has the potential for ranging from large volumes of water, which
impacts at different biological levels. Manage- have been used for cooling purposes, through
ment strategies may be employed to reduce to chemical waste products which may be
or remove the effects of aquaculture. The deve- extremely toxic even in small quantities.
loper may collect the excess solids by funnel
or suction system although decomposition,
6.8.1 Petrochemical discharges and inputs
a release of nutrients, and poor water quality
can still occur. Pumping or bed trawling to aid Crude oil is a complex natural mixture dom-
breakdown could disperse the wastes. Good inated by hydrocarbons but also containing
siting principles will minimize the effects or the non-hydrocarbons and metals, each fraction
cages may be moved at intervals, with those have a varying toxicity, behavior, and persist-
intervals being judged by the degree of biotur- ence in the environment. It has been estimated
bation under the cages, hence using the assim- that over 2 million tonnes of oil per annum is
ilative capacity of the area. Feeding techniques discharged into estuaries, being approximately
114 T H E E S T UA R I N E E C O S YS T E M
one-third of all the oil discharged into the could affect mudflats, salt marshes, and
worlds oceans. Thus estuaries do indeed reedbeds. Once on the sandy beaches, the oil
receive a disproportionately large burden. can be pushed into the coarser sediment by
Whereas oil spillages and tanker accidents tidal pumping and once in the sediment, espe-
receive a large amount of press coverage, by cially if there is no oxygen penetration then
far the greatest proportion of oil enters the aerobic degradation will be reduced. Hence
estuaries and coastal seas from diffuse sources the oil can remain for many months. On the
such as urban and river runoff or domestic or shore the oil acts mechanically, smothering
industrial waste. The effects of the oil industry animals in burrows or on rocks, and excluding
on estuaries may be divided into (1) the impact the light from plants. Given that mudflats have
of gross spillages, due to shipping accidents or larger infauna and predator populations than
human error at a loading terminal, (2) the sandy beaches then oil deposited on the for-
impact of effluents produced by refinery and mer has a greater smothering effect and affects
petrochemical industries, and (3) the effects of predatorprey relationships. Oil deposited on
oil extraction. higher energy sandy beach surfaces and on
rocky shores will be subjected to greater phys-
Oil spillages ical degradation and thus may breakdown
Oil spillages from harbors or collisions have more easily. In contrast, oil deposited in low
many and varied potential effects, which are energy estuarine environments such as
the result of the nature of the oil spilled, the mudflats, seagrass beds, salt marshes, and
characteristics of the estuary and the climatic reedbeds will persist for longer and is also
conditions. Some of the different fractions more difficult to remove by mechanical meth-
may be toxic while others are relatively inert ods. It will infiltrate creeks and between the
and all the fractions will be subject to physical, plants and thus be protected from mechanical
chemical, and biological degradation. When breakdown. On a salt marsh the soiled parts of
oil is spilt from a harbor or a collision it the plant die, but if it can survive this, the plant
reaches water as a separate phase, which can regrow.
is largely immiscible with the water, and gen- The oil will also be mixed with the water
erally forms a surface slick (Fig. 6.15). The column forming first water-in-oil emulsions
slick will spread and be subject to physical and then, with increasing mixing, oil-in-water
and chemical breakdown through photo- emulsions (often called chocolate mousse due
oxidation, evaporation and, during high to the color and consistency). Zooplankton
winds, aerosol formation. Hence the import- will ingest the oil droplets and produce fecal
ance of the weather conditions affecting the pellets, which can aid breakdown by bacterial
fate and effects of the spilled oil. Many of the biodegradation on the large surfaces. These
lighter and most toxic elements can be lost by droplets and pellets may then settle out of the
weathering, but unfortunately, in the confined water column. The water-soluble or more
space of an estuary there may not be time for volatile compounds are most toxic, and
this to occur before the spilt oil is deposited on spillages in American estuaries of fuel-oil
the shore. containing 45% low aromatic oils have devas-
The surface slick and the sinking oil will tated commercial shell fisheries. The heavier
be transported to areas of the estuary depend- compounds within the oil, which remain after
ing on the wind and tidal conditions and evaporation, will tend to sink, where they
the nature of the estuary. For example, a undergo microbial decomposition by many
spillage at the mouth of the estuary will microorganisms, which are capable of degrad-
impact on local rocky shores and sandy ing petroleum hydrocarbons as well as
beaches whereas that pushed up the estuary naturally occurring biogenic hydrocarbons.
E S T UA R I N E U S E S A N D U S E R S 115
Atmosphere
Solution (water
Absorption soluble fraction)
and Convection
Adsorption
Dispersion Up-welling
Solution, etc. Micro-particulate tar
Emulsification
Chemical
degradation
In suspension
Absorption and Dispersal CO2
Adsorption, etc.
Biodegradation
Food chain
Oil Resuspension
(subsurface spill) Sea bottom
Biodegradation
and fauna
Sediment Sediment
Reworking benthos
Long term burial
Figure 6.15 The fate of oil spilled or discharged into surface waters
(modified from CONCAWE 1981, www.concawe.org).
116 T H E E S T UA R I N E E C O S YS T E M
The most resistant oil fractions, the heavier from chemical works. A refinery effluent may
and less volatile components, however, contain only 1020 ppm of oil, which is dif-
remain as tar-balls for many weeks if not ficult to remove and seen perhaps only as
months. cloudy water with an oily surface sheen, but
Given the serious consequences of oil pene- as a flow of 7.58 cumecs this produces more
trating the more delicate habitats in estuaries, 6.55 tonnes of oil per day. The impact of efflu-
especially those where oil is difficult to ent from petrochemical complexes in the
remove once it settles, then the most common Medway and Forth estuaries was examined
advice on how to deal with oil spills in estuar- before major clean-up operations. In the
ies is to contain the oil by a boom if possible, vicinity of the outfall an abiotic zone was
and then to use mechanical removal with found, with no life at all. Beyond this was
pumps for the trapped oil. If it comes ashore a grossly polluted zone in which only
on sandy beaches then mechanical removal Oligochaetes were found, in small numbers.
with shovels or excavators is the preferred Then followed a polluted zone with abun-
mode of treatment, rather than emulsifiers dant Oligochaetes and Manayunkia (a small
if at all possible. The earlier emulsifiers, or polychaete), and occasional specimens of
detergents, used in clean-up campaigns to other species such as Hydrobia. Finally came
disperse oil were often more toxic to estu- the largely unpolluted zone at over 1-km
arine life than the original oil. Newer emulsi- from the effluent, with Macoma, Cerastoderma,
fiers are less toxic. Oil washed off rocky Nereis, Nephtys, and Hydrobia often abundant,
shores with hot and/or freshwater can cause and fewer Oligochaetes. It may be seen that
greater community effects than the oil if the zonation of species and their abundance is
left alone. Oil settling on to mudflats, salt remarkably similar to that noted above for the
marshes, sea grass beds and reedbeds in effects of organic enrichment. Since oil is a
estuaries cannot be removed without con- biological product derived geochemically
siderable damage to those habitats and so if from organic material, it is perhaps not so sur-
these areas are oiled then the decision is very prising that the effects of oil on estuarine
often to leave the oil to natural clean-up. ecosystems are rather similar to the effects of
Consequently, an estuarine oil-spill contin- other excess organic materials. Throughout
gency plan may involve recovering the oil the world oil refineries have made strenuous
from sandy beaches rather than other habitats, efforts to reduce their waste effluent, and
especially as the beaches may have good modern oil refineries have considerably less
access and will support heavy machinery. impact on the estuarine ecosystem than their
Figure 6.16 shows the recommended decision older counterparts. When an oil refinery dis-
processes involved in dealing with an oil spill charge is reduced, or terminated, then the
within an estuary. resilient estuarine fauna can generally recolo-
nize the area rapidly.
Petrochemical discharges
Less dramatic, but potentially more dangerous Oil and gas extraction
to estuaries than oil-spills, are the insid- Oil is extracted from several estuarine loca-
ious effects of the continual discharge of tions, most notably the Nigerian oilfields are
industrial effluents from petrochemical com- located within Nigerias estuarine delta
plexes. The effluent from such industries is region, and blow-outs from the wells have
mostly hot freshwater, which may contain occasionally led to environmental problems.
certain amounts of chemicals, including The shallow areas of eastern England and the
hydrocarbons (oil) from refineries, or chemical Dutch coast border on the southern North
waste products such as phenols or ammonia Sea gas fields and the large estuarine area
E S T UA R I N E U S E S A N D U S E R S 117
Oil spilled
Observe or predict
Implement No
Yes Yes
Is a dispersion
Continue Clean up
operation No
actions on-shore
possible? (a)
Yes
Figure 6.16 Decision tree for the response to an oil spill in an estuary, considering in particular whether a chemical
dispersant is acceptable. (From Concawe, 1981.)
Morecambe Bay in NW England also sits on trapped with the oil and gas, which may
a gas field and thus these areas support gas contain high levels of minerals and be at
rigs as well as seabed collecting systems. Gas a high salinity whereas production water is
and oil come ashore by pipeline into several that pushed down into ageing wells to
estuaries, and so formation water and produc- increase production. Each of these will contain
tion water has to be treated and disposed. oily residues and thus require treatment
Formation water is the fossilized water before disposal into the estuary.
118 T H E E S T UA R I N E E C O S YS T E M
60 20
enclose the metal within a membrane, has
lin
15
Sa
40
20 10
been found. These vesicles prevent contact of
5 excess metal with vital constituents and effec-
5 10 15 tively detoxify the metal. Estuarine annelids,
Temperature ( C)
such as Nereis have been shown to be able to
Figure 6.17 The effect of temperature and salinity on median detoxify metals by accumulating them within
survival time (h) of Corophium volutator at a chromium their jaws. Although detoxification may pre-
concentration of 16 mg l!1. (After Bryant et al. 1984.) vent a mollusc or an annelid from being
E S T UA R I N E U S E S A N D U S E R S 119
48 h 96 h 192 h
15 C
5 C
10 C
1000
10 C
5 C
15 C
LC50 mg l1 10 g scale
10 C
5 C
15 C
100
10
15 20 25 30 35 15 20 25 30 35 15 20 25 30 35
Salinity
Figure 6.18 The effect of temperature and salinity on the median lethal concentration (LC50) of nickel to
Macoma balthica. (From McLusky et al. 1986.)
adversely affected by excess metals, a preda- estuarine sediments, and only a lesser amount
tor eating it may not possess the same ability is eventually discharged to the sea. If estuaries
and may be affected by the total concentration are to serve as such vital traps for pollutants, it
of the metal. Unlike some other pollutants, is essential that the natural intertidal mudflat
heavy metals may show biomagnification. That areas remain intact and undisturbed. In sev-
is, they show an increase in concentration as eral cases it has been shown that mans activ-
one proceeds up the trophic levels of the estu- ities may disturb such metal burial, for
arine ecosystem. For that reason, humans example, dredging will often liberate heavy
should always avoid eating any shellfish metals from sediments, and even bait-digging
exposed to heavy metals. for worms has been shown to lead to an
The processes of sedimentation within estu- increase in the total concentration and
aries, discussed in the first chapter, may serve bioavailability of metals.
to relieve the estuarine and marine ecosystem
of some of the worst effects of heavy metal
6.8.3 Organohalogens
pollution. As metals are discharged into the
turbid waters of estuaries they may become The addition of pesticides and organohalogen
rapidly bound onto the surface of the fine sed- compounds to estuaries poses considerable
imentary particles. As the sedimentary parti- threats to the estuarine organisms. The
cles settle on to the intertidal mudflats, the organohalogen compounds are the poly-
metals are gradually buried. In many of the chlorinated biphenyls (PCBs), including
estuaries bordering the North Sea, it has been hexacholorobenzene (HCB), lindane, hexa-
estimated that about half of the metals enter- chlorocyclohexane (HCH), dieldrin, and
ing the estuary become trapped within the dichloro-diphenyltrichloroethane (DDT) used
120 T H E E S T UA R I N E E C O S YS T E M
both as pesticides, and for industrial purposes unique detectability of radioactive isotopes
(especially in electrical equipment). Unlike enables the anthropogenic additions to the
petroleum hydrocarbons chemical oxidation or environment to be identified clearly and quan-
microorganisms cannot degrade these materi- tified. It is, however, often forgotten that the
als, generally known as the halogenated hydro- man-made additions may be very small in
carbons (hydrocarbons containing chlorine, comparison to the amount of radioactivity that
bromine, iodine, or fluorine). Thus, like metals, naturally occurs in seawater. Over 90% of the
these substances are permanent additions to natural radioactivity of seawater arises from
the estuarine ecosystem, but, compared to met- potassium-40, almost 1% from rubidium-87,
als, even minute quantities of these man-made and the remainder is virtually all from the
substances can pose a hazard to estuarine life. isotopes of uranium. The concentrations of
Any animals, which ingest them, cannot these elements vary proportionately with
excrete these substances, and they thus accu- salinity, so the natural radioactivity of estuar-
mulate within their bodies, principally in fatty ine water is governed by salinity, with greater
tissue although detoxification can occur. Food- concentrations in saline waters and lesser con-
chain biomagnification can occur so that top centrations in brackish waters. At a typical
predators have the highest concentrations, and salinity of 33 the natural radioactivity has been
may suffer most. Until the effect of these measured as 12 Bq L!1 (1 Bq (Bequerel) is one
substances was realized, and their production disintegration per second). In the Forth estuary
curtailed, about 1 million tonnes of PCBs were and the Firth, the volume of water is 20 km3 or
produced around the world. Much of this 2 ' 1013 l, so Leatherland (1987) calculated the
material has ended up in estuarine and marine total natural radioactivity of these waters as
organisms, and will continue to circulate 240,000 GBq, indicating a total of 60 tonnes of
through estuarine ecosystems. dissolved uranium in these waters. In this area
Pesticide or industrial PCBs in very low con- a naval dockyard adds 7 GBq per annum, and
centrations, for example, 1 %g l!1 ($1 ppb) low levels of isotopes will be added in sewage
have been shown to be lethal to estuarine life systems derived from hospital or laboratory
both in laboratory tests, and in field mortalities sources. A nuclear power station has added a
around industrial and agricultural discharges further 200 GBq per annum, but it can be seen
containing pesticides. Due to the biomagnifica- that these sources are minute compared to the
tion effects of PCBs, top predators such as natural levels of radioactivity present.
birds and seals are particularly at risk. Within In Britain and France the major anthro-
the Baltic Sea, for example, high levels of PCBs pogenic source of radioactivity is from nuclear
within seals have been held responsible for the reprocessing plants at Sellafield and La Hague.
failure of reproduction in many female seals. These plants recover uranium for re-use from
the fuel rods of nuclear power stations, and
discharge large quantities of waste water with
6.8.4 Radioactive discharges
a low radioactive content, but because of
The subject of the addition of radioactive the volumes involved they generate a consider-
substances to estuarine environments arouses ably greater input of radioactive material than
considerable public concern. Man has, during nuclear power stations or other man-made
the latter part of the twentieth century, added sources. Some of the waste products decay
radioactive material to seawaters from the rapidly or are adsorbed onto fine sediments.
fallout of atmospheric bomb testing, from The fine sediments can be carried into local
waste discharged from nuclear power stations estuaries, and in the case of Sellafield these
and reprocessing plants, and from the use need to be regularly monitored. The isotope
of radioactive material in submarines. The caesium-137 remains in solution, and since it
E S T UA R I N E U S E S A N D U S E R S 121
does not occur naturally it serves as an effective 6.8.5 Threats from industrial pollutants
marker for the Sellafield waste. Regular
Of the various pollutants discussed, three
annual surveys, of British coastal waters,
groups of substances pose particular threats to
show that waste from Sellafield is predom-
the estuarine ecosystem. These are heavy
inantly carried north and west away from the
metals, PCBs, and radioactivity. All these
discharge, and after passing around the north of
substances are persistent, and cannot be
Scotland enters the North Sea and its estuaries.
degraded, although they show some slow
Close to the discharge point the concentration
reversibility due to burial in sediments. All
of caesium-137 is 0.5 Bq kg!1, and declines to
show bioaccumulation, and in the case of
0.05 Bq kg!1 in the North Sea (Fig. 6.19). These
PCBs also show clear biomagnification. All
concentrations are considerably less than the
show effects on the fauna at low concentra-
natural radioactivity levels of seawater, but are
tions, often at concentrations well below those
greater than any other anthropogenic sources of
shown to be toxic in standard 96-h toxicity
radioactivity. In recent years the quantities of
tests. The threats posed by metals and PCBs
waste discharged from reprocessing plants has
can be demonstrated in present estuarine
diminished considerably, but they remain the
ecosystems, but radioactivity effects have not
main source of man-made radioactivity in estu-
been shown to disrupt ecosystems so far. By
aries. Laboratory studies have shown acute
contrast, organic materials and oil may show
lethal effects of high radiation doses on many
severe effects in the vicinity of discharges, but
organisms, but it must be said that existing lev-
these materials can be biologically degraded,
els of natural and man-made radiation in estu-
and their effects can be completely reversible
aries and the sea have so far produced no
at the estuarine ecosystem level. So, when con-
measurable environmental impact on these
sidering pollutants in estuaries one must con-
ecosystems.
sider not only the quantity and quality of the
waste, but also whether it can be degraded
12 8 4 0 4 and whether its effects are reversible.
sea walls converted them into productive over 90% of the Tees estuary, leaving less than
agricultural land. In Britain such land-claim 10% of the original intertidal area (Fig. 6.1),
has been predominantly on the east coast, and and in the Forth estuary, approximately 50%
in Denmark such land-claim has occurred in of the intertidal area has been lost through
West Jutland. In India and Bangladesh such land-claim in the past 200 years (Fig. 6.20). At
claimed land has proved to be particularly first such land-claims were for agricultural
suitable for rice cultivation. Such processes of purposes, followed by harbor works, and
agricultural land-claim are, however, gradual more recently for industrial purposes, includ-
in comparison to the large-scale land-claim ing sites for power stations, refuse disposal
schemes undertaken to meet the needs of and fly-ash dumping. Within Southampton
industry. Modern industrial developments Water land-claim has taken place over many
seek areas of flat land close to available trans- years to meet the needs of new docks, oil
portation, and as other land may not be read- refineries, power stations, and other industries
ily available, large-scale estuarine land-claim (Fig. 6.21). While the salinity regime has been
is often suggested. For such a land develop- hardly affected by the land-claim schemes, the
ment scheme, the first stage is usually the con- removal of extensive salt marshes has reduced
struction of a seawall or bund across the substantially the input of organic detritus to
intertidal mudflats, followed by the infilling of the ecosystem, and made the whole ecosystem
the bounded area with dredged estuarine more dependent on phytoplankton produc-
mud, or hard-fill material derived from quar- tion. As intertidal areas have been obliterated,
ries, mine waste, ash waste from coal-fired the commercial clam harvest has fallen from
power stations, or domestic refuse. In other 100 to 60 tonnes per annum, and the bird
schemes, the impounded area may be filled population feeding intertidally has declined
with freshwater to form reservoirs, and espe- substantially.
cially in the Netherlands the impounded area San Francisco Bay is the largest estuary on
may be drained and converted into agricul- the Pacific coast of the United States, with a
tural land or polders. water surface area of 1240 km2, and probably
Whatever the use of the claimed land, it the most modified estuary in America. Large
represents a total loss to the estuarine eco- areas of wetland, or salt marsh, have been
system. In the estuaries of eastern Britain, for destroyed by land-claim, so that only 6% of
example, industrial land-claim has obliterated the original 2200 km2 of wetland remain.
Stirling Alloa km
0 1 2 3 4 5
N
New docks
River Itchen
Old docks
0 1000 m River test
Due to inland water engineering schemes less estuaries of Rhine, Meuse, and Scheldt have
than 40% of the original river flow now been dammed and converted from tidal estu-
reaches the estuary, a factor that has consider- aries into freshwater lakes, brackish lakes,
ably reduced the flushing of pollutants from or sea areas with restricted tides (Fig. 6.22). The
the Bay area. On the east coast of the United southernmost estuary, the Westerscheldt,
States more than 25% of the estuarine wet- remains open, but the Oosterscheldt estuary
lands have already been lost through land- has been cut off from its freshwater supply (the
claim. Wherever, and for whatever purposes, Rhine Meuse), large areas of shallow wetland
land-claim occurs, the birds and fish, which have been claimed for land, and an 8-km wide
feed on estuarine mudflats, simply find that a tidal storm surge barrier restricts the inflow
large proportion of their food supply has dis- of seawater. As a consequence it is now a
appeared. American legislation now requires sheltered marine bay, not an estuary. The
that new land-claim schemes must compen- Grevelingen estuary has been closed at both
sate for the loss of habitat by the provision of its river and seaward ends to form a semi-
alternative localities; however, it is very stagnant nontidal brackish-water lake, with a
doubtful if such compensation can ever fully stable salinity of about 18. The small Veere
replace a habitat which has taken centuries to estuary has also become a brackish-water lake
develop. with a salinity of 7 in the winter, rising to 12 in
the summer. The Haringvliet estuary has been
closed at its mouth, preventing the entry of salt
6.9.1 Dutch delta works
water, but still permitting the exit of freshwater.
Faced with the problem of the threat of large- Consequently it has become a freshwater lake,
scale flooding from the North Sea, the fed by the Rhine. The necessity for the Delta
Netherlands has claimed extensive estuarine Plan to the safety of the Netherlands is clear,
areas. In the past three decades major changes but the consequence has often been the obliter-
have occurred in the estuaries of the southern ation of mussel beds and the elimination of
Netherlands, in the Delta Plan, whereby the feeding sites for birds and fish.
124 T H E E S T UA R I N E E C O S YS T E M
N
North Sea Dam Haringvliet
Canal
0 10
km
Westerschelde
Figure 6.22 The Delta scheme area of southwest Netherlands. The map shows that four previously open estuaries have
been partially or completely closed off from the sea. In addition inland compartment dams have created freshwater lakes
from previous estuaries. Only the Westerschelde remains as an estuary. (After Nienhuis and Smaal 1994.)
some of the areas that previous generations and often repeated removal of material,
have removed by land-claim. This reoccupa- typically sand, silt, and gravel deposited in
tion, known as managed retreat is being low energy, accreting areas by river discharge,
trialled at several locations and may represent tidal currents, or wave action in areas pre-
the future for many estuarine areas. viously dredged. In contrast, capital dredg-
ing is the first time removal of sediments,
which may be necessary to create a new
6.9.3 Ports, navigation, and dredging
channel, build a new bridge or lay a new
The use of estuaries for transportation requires pipeline of telecommunications cable. Hence
harbor works, which can range in size from a it is likely that sediments for maintenance
single pier to a vast container or bulk-cargo dredging are recently deposited and relatively
terminal. A single pier may cause localized un-compacted although they may be accumu-
change in current pattern and sedimentation, lating organic matter and pollutants, which
whereas a large terminal will have a similar easily settle, flocculate, or precipitate from
impact to other major land-claim schemes. the water column.
The major ports of Southampton, Rotterdam, The nature of the material, the area, and the
Antwerp, and Hamburg are all situated on type of equipment used dictate the environ-
major estuaries and all have developed by mental impacts of dredging and dredged
building on land-claim, building wharves into material disposal. The different types of
the deep channel. Estuaries have port develop- dredger include those that remove material
ments, which depend on the position of the mechanically by means of a grab or bucket,
channel, for example, the four ports of Hull, whereas suction dredgers transport the mate-
Grimsby, Immingham, and Goole on the rial as a suspension via a pipeline. Silts and
Humber estuary, which are the largest UK fine sands can also be cleared from harbors
ports complex, each developed where the deep and dock entrances by agitation dredging,
water channel touched either the north or the that is, resuspending the material and then
south banks in an otherwise shallow estuary. relying on water currents to carry it away.
In order to maintain access for shipping, Usually the dredged material is loaded into a
many estuaries are dredged. At the dredge hopper, either in the dredging vessel or in a
site, the substratum is removed along with separate vessel, for transport to the disposal
any fauna it contains, and the dredge spoil is sites where the material is discharged by pipe
then transported to a disposal ground, usually or bottom-opening doors. The environmental
at the mouth of the estuary, or outside it. The effects thus result from the way the sediment
fauna of the dredged and disposal grounds is obtained at the dredge site and liberated at
may be affected by being moved or smoth- the disposal site.
ered, and pollutants accumulated in estuarine Disposal sites were first selected for conveni-
sediments, which act as a sink, may be ence in being close to the dredged area but
released. Harbor sites may also be a cause of sufficiently far away so as not to affect naviga-
pollution, especially from oil, through accid- tion. The disposal site for a small fishing harbor
ents as ships are loaded and unloaded, or due or marina may receive maintenance dredging
to the risk of collision, or from leaks from on an annual or bi-annual basis, consisting of a
ships engines or bilges. Ship repair and demo- few thousand tonnes of marine sand moved
lition sites also present potential hazards for into the harbor area from the adjoining seabed
considerable further accidental pollution. by winter storms. In contrast, the disposal
Dredging operations in estuaries can be ground for large estuarine ports may receive
divided into either maintenance or capital many millions of tonnes of low density silt
dredging. Maintenance dredging is the periodic several times each day.
126 T H E E S T UA R I N E E C O S YS T E M
In the United Kingdom in 2000, 10 sea objectives (Elliott et al. 1998). As with many of
disposal sites received #1 million wet the activities considered here, we can use our
tonnes of dredged material, 24 sites received knowledge of these effects and the linkages
1 million100,000 wet tonnes, 40 sites received between the different responses to create a
100,00010,000 wet tonnes, and 21 sites conceptual model which, by the nature of the
received "10,000 wet tonnes. While a rela- system and the potential changes to dredging
tively small number of sites accounted for a and marine disposal, is naturally complex
large proportion of the United Kingdoms (Figs 6.23 and 6.24, adapted from Elliott and
2000 dredged material disposal, at least 95 sites Hemingway 2002).
had some material deposited in them. A num- Dredging will alter the bed topography and
ber of the sites received material from several bathymetry, which in turn will remove the bed
different dredging areas. Despite this, not all organisms and it substratum, as well as
dredged material is disposed of at sea as sig- changing the overall hydrodynamic regime of
nificant tonnages are placed ashore in lagoons, the area. The structure and functioning of
land-claim areas, and landfill sites. On-shore those bed sediments and the overlying hydro-
disposal may be used for infilling develop- graphic regime (water currents, tidal circula-
ment land and as beach nourishment for coast tion, etc.) will be intimately linked to the
protection. In cases where the material being structure and functioning of the bed biological
dredged is deemed to be too contaminated for community, principally the invertebrates
sea disposal then it may be placed in contain- (Elliott et al. 1998). In turn this will influence
ment sites such as the Slufter system created the fishes and, in estuarine nearshore areas,
by the Port of Rotterdam in the 1990s. the birds feeding on those invertebrates. The
Both dredging and disposal of estuarine second major effect of dredging will be the
sediment can affect the physical, chemical, and resuspension of the bottom sediments and
biological features of the water column and the its effects on the water turbidity, and the
seabed and the impacts could be short-term or liberation of any materials contained and
long-term. The potential impacts could include sequestered within the sediments. Dredging
a deterioration in the overall health quality of will result in the sediments being exposed to
the marine ecosystem; a reduction in the aerated waters and this change in chemical
socioeconomic aspects of the sea, including nature can re-release the contaminants. In
fishery and amenity interests; an interference addition, anoxic and organic rich estuarine
with the legitimate uses (i.e. those legally per- sediments may generate methane and hydro-
mitted) of the sea recreation and navigational gen sulfide, which is liberated on disturbance
aspects, and a reduction in the aesthetic qual- by dredging. The release of those materials
ities associated with the area. into the water column will then have the
The potential effects of dredging and potential for a biological effect. All of these
dredged-material disposal can be regarded effects have the potential to influence the fish-
as a set of bottom-up causes, in which the eries and nature conservation value of the area
physical system (both in the water column (Fig. 6.24).
and on the bed) is altered and in turn affect The disposal of dredged material will sim-
the health of the biological system. The even- ilarly have the potential to affect the water
tual effects on the biological system and its column, the bed conditions and their biota
uses by Man can be regarded as a set of top- (Fig. 6.23). Reductions in water clarity through
down responses, for example, the effects on an increased turbidity will in turn affect the
the higher levels of the ecological system primary production by the phytoplankton.
(such as fishes, seabirds, and marine mam- The release of any materials contained within
mals) as well as on fisheries and conservation the dredged material, either as the water
E S T UA R I N E U S E S A N D U S E R S 127
Figure 6.23 Potential environmental impacts of estuarine dredged material disposalconceptual model.
(Modified from Elliott and Hemingway 2002.)
soluble fraction or the release of particulate Whereas most of the above impacts of
materials may be the result of the changed dredging and dredged-material disposal
chemical environment, that is, anoxic fine relate to the ecological system, the resultant
sediments liberated into the oxygenated water impacts on the uses and users of the marine
column will cause the release of pollutants pre- environment are often of greater public con-
viously sequestered due to the anoxic chemical cern. These include the actual or perceived
conditions. Similarly, any organic matter in the effects on socioeconomic aspects such as
sediment will create a water column oxygen fisheries, and aesthetic aspects, including
demand. The deposited sediment will change recreation and tourism. Similarly, the per-
the nature of the bed sediment, if it is of a ceived or actual effects on the conservation
different particle size and it can have a smoth- importance of an area will be of concern, espe-
ering effect on the bed community as well as cially where the habitats and species within
bringing new organisms to an area. Both of and adjacent to the dredging and disposal
these features will affect the structure of the areas are of importance.
bed community and in turn the demersal and In addition to capital and maintenance
benthic fishes feeding on that bed community. dredging, estuaries are often the sites of the
128 T H E E S T UA R I N E E C O S YS T E M
Figure 6.24 Potential environmental impacts of estuarine dredgingconceptual model. (Modified from Elliott and
Hemingway 2002.)
winning of aggregates to supply the needs of now supporting large power plants and having
construction industries. Sand and gravel are barrages placed across the estuary for large-
often dredged from the bed of the lower scale tidal power. Power generation can be sep-
reaches of an estuary. However, given that arated into conventional plants and alternative
bed sediments are rarely a single type of mat- sources. The former use fossil fuels, such as oil,
erial, the desired grade of sediment requires to gas or coal, or nuclear energy to heat water to
be separated from other, often finer material create steam, that turns turbines and need large
and the latter is then discharged over the volumes of water to condense the steam after
side of the dredging vessel. This can lead to use. In contrast, alternative sources, often called
smothering of the bed sediments and fauna renewable generation, uses the tides, waves,
and changes to the water quality, especially water currents, winds, the sun, or freshwater
the turbidity levels. However, given that impoundment (hydroelectric).
estuaries are often highly turbid and their Many of these generating sources occur on
biota are adapted to sedimenting condi- estuaries or at least impinge on estuarine
tions then the effects may be less than on the processes, especially conventional plants and
open coast. tidal power systems. Wave power is not suc-
cessful in sheltered estuarine areas and as yet
there are no water current systems in which an
6.10 Power generation
open propeller is placed within the water
Estuaries have long been the sites of power body. Hydroelectric schemes are not found in
generation, from having small-scale water estuaries, despite this these schemes may
wheels operated by tidal flow to grind grain, to affect the inhabitants of estuaries such as
E S T UA R I N E U S E S A N D U S E R S 129
impeding the migration of diadromous fishes, other cases the material has in the past been
such as the shad (Alosa spp.) or the salmon dumped at sea. Such a fine material input
(Salmo salar). Conventional power generating changed the nature of the bed sediments by
plants are often sited in estuaries because of blocking pore spaces, smothering the bed
their associated infrastructure and the availabil- fauna, and affecting colonization. Trace metals
ity of a large supply of cooling water. As with all in the ash can also leach into the surrounding
major structures, the power plants will have an waters. This material has been used in pro-
impact on the estuarine environment but also ducing breeze blocks for construction and
the estuarine environment has an effect on the sometimes for creating artificial reefs, such as
power plant. the experimental site near Southampton,
The initial and often long-term effects are southern England.
from the construction and site development The greatest environmental impacts are,
and maintenance. Land-claim land loss, espe- however, due to the main reason for siting
cially of intertidal area, may occur to provide the power plant on the estuarythe intake
space for construction or for the waste from and hence discharge of large volumes of
the power plant. Any change to the shape of water for cooling the condensers. This is
the estuary will produce hydrophysical dis- referred to as direct cooling and thus removes
ruption, with changes to the hydrography, the need for cooling towers common to inland
sediment regime, and bathymetry. As with power plants. These impacts can be separated
any major construction, there can be construc- into those related to the intake and those for
tion discharges and runoff of oils, solids, and the discharge. A large power plant may
litter. In the case of large power plants there require 3050 m3 s!1 (cumecs), the equivalent
will be dredging, seabed aggregate extraction, of a medium sized river, and so the power
aggregate movement, and storage as the plant has to be positioned where there is an
seaborne delivery of building materials is adequate water source, hence an estuary. In
often the most efficient. The creation of nuis- taking in this volume, large and small material
ance, often termed aesthetic pollution, from is sucked into the plant, termed impingement.
light, noise, and vehicle and ship movement, Material greater than 1 cm2 will be retained on
will affect the large overwintering wading the initial rotating screens inside the power
bird populations. In addition there will be hin- station. This material includes fishes, large
drance to resource exploitation by boat and and mobile invertebrates such as lobsters,
fixed net fishing and to recreation and crabs and, in the outer parts of estuaries, squid
tourism. Finally, the visual impact of the plant and octopus, seaweed, jellyfishes, and litter.
may affect the amenity value of the area. The amount and nature of this material
The operation of power stations and pro- depends on the type of biological commun-
duction of power causes environmental ities adjacent to the plant and the position of
impacts both from the waste, especially from the intake pipes.
coal-fired power plants, and because of the Up to several tonnes per day of fish, other
intake and discharge of cooling water. Coal- organisms and debris can be impinged, espe-
fired power plants use pulverized coal as cially if large inshore and estuarine fish migra-
fuel and thus they produce pulverized fuel tions occur. For example, a power station in
ash or fly ash (PFA), which may be up to a the Forth estuary, Scotland, is situated in an
million tonnes per year for a large power area supporting large overwintering estuarine
plant. In some cases this material has been fish populations (Elliott et al. 1990), which
used to reclaim estuarine intertidal areas, such become impinged. Hence power plants have
as at Culross Bay, Scotland, thus removing been described as stationary trawlers. The
productive wader feeding areas, whereas in impinged fish suffer injury and death through
130 T H E E S T UA R I N E E C O S YS T E M
the pressure effects, which cause swim-bladder heat produced in the system leads to the
damage in round fish, external damage to production and liberation of chlorine residuals
scales and fins, for example, in delicate fish and organohalogens (bromoform, bromoamine,
such as clupeoids, and the hemorrhaging of chloroform, chloroamine, etc.), themselves
eyes. If there is a high mortality then this polluting materials when discharged from the
might have an influence on the inshore and plant into receiving waters.
estuarine nursery function and on migration The cooling water discharged into the estu-
routes, hence disrupting fisheries. Such an arine receiving waters can also lead to environ-
influence is greater on round-fish than flatfish. mental changes given that it is warmer than
Large amounts of impingement, such as of ambient and may produce an obvious thermal
jellyfish in the Emshaven power plant on plume covering many square metres. There
the Ems Estuary, the Netherlands, can also can be an influx of non-native species and the
severely affect the operation of the plant and physiological disruption and acclimation of
even lead to the plant being shut down. The native species. The temperature changes can
impinged material, termed trash, can be disrupt reproductive cycles, for example,
returned to the estuary or disposed to sea or summer spawning invertebrates will have
landfill under licence although power plants their spawning period extended whereas
would generally prefer to reduce the impinged winter spawners will have theirs curtailed.
levels by the use of intake design or acoustic, These could lead to population and commun-
light, or bubble screens. ity changes. The high flows of discharged
Materials smaller than 1 cm2 will pass cooling waters can lead to bed erosion and
through the rotating screens and thus pass substratum disruption as well as the disper-
into the main cooling system of the plant. sion of chlorine residuals and organohalogens
This small material includes larval forms as into receiving waters. In turn, there will be
well as the silt found in high concentrations potential bioaccumulation and cellular and
in the estuarine waters. This leads to the physiological effects of the latter.
fouling of the pipes inside the plant by ses- Coastal and estuarine power plants have
sile fauna (spat, barnacles, saddle oysters, other emitted and produced materials such as
bryozoans, hydroids, and tubeworms) and aerial emissions, leading to atmospheric
bacterial film. If left unchecked this leads to inputs, of CO2, NOx, SOx, etc. (greenhouse
blockages and a reduction in the cooling effi- gases and nutrient inputs). Radioactive mate-
ciency, especially condensers and areas within rials may be discharged in water emissions as
the cooling water stream, any auxiliary stream well as fuel waste storage and reprocessing
and, in the case of nuclear stations, the reactor from nuclear stations.
cooling water. As such there is the need for bio- Given the potential effects then it is neces-
fouling prevention measures involving biocide, sary to produce mitigation measures for the
antifouling, and anticorrosion treatment. The reduction of the effects. There should be
latter relies on the transport, production, stor- plant design and operation to environmental
age and use of biocides from oxidizing com- standards (e.g. ISO 14001) and the rehabil-
pounds such as liquid chlorine, powdered itation of areas after use. Trash could be
sodium hypochlorite, electrochlorination, and returned or impingement minimized through
bromine-enhancers. There will be large mortal- intake design and deterrent methods or the
ities of planktonic organisms and larval stages, plant operated on a regime related to fish
although there are vast quantities of these in migration patterns. The discharge apparatus
the water column. The introduction of halo- could be designed to reduce outflow scouring,
genated antifouling agents coupled with high for example, by using plastic seaweed on
levels of organic matter in the water and the the bed, and to disperse heated water. Heat
E S T UA R I N E U S E S A N D U S E R S 131
treatment could be used instead of chemical a storm surge to develop in the northern
antifouling and fuel gas desulfurization (FGD) North Sea and then progress down the eastern
used to control atmospheric inputs. UK coast and then move around to the eastern
North Sea coasts in Belgium and the
Netherlands. This produced high tidal levels
6.11 Estuarine barriers and barrages
in the adjacent estuaries and led to over-
Many estuaries have long had small physical topping of seawalls. As a result of this, many
barriers at their upper regions, for example, lives were lost in the United Kingdom and the
weirs that mark the head of the estuary. In the Low Countries thus leading to the creation of
last few decades, however, larger permanent the Dutch Delta Plan and the need for higher
barrages and temporary barriers have been seawalls and estuarine barrages in Eastern
placed across several estuaries for various England.
reasonsto protect low lying areas from The estuary of the River Hull, leading into
flooding due to storm surges and high tidal the Humber estuary, Eastern England, has
conditions, in order to increase the amenity a movable barrier, which is lowered into
value of adjoining areas, and for tidal power place during high tides in order to protect the
generation. The interference of hydrographic city of Kingston Upon Hull (Fig. 6.25).
patterns within the estuary by the presence of Similarly, the Thames Barrier (Fig. 6.26) is
such structures will in turn change the phys- raised following the threat of high tides in
ical nature of the estuary and thus its biological order to protect low lying areas in London. As
structure and functioning. a much larger structure, the Oosterschelde
Low-lying areas are exposed to high tides (Eastern Scheldt) estuary in the southern
and especially storm surges, particularly in Netherlands has a storm surge barrier to pro-
cases where the tidal wave may get amplified tect the inner, low lying areas against flooding.
by a constricting topography such as within a Although this is a semi-open structure in
funnel-shaped estuary or a funnel-shaped which gates are lowered between piers follow-
semi-enclosed sea. For example, climatic con- ing the threat of high tides, this has changed
ditions, wind direction, and atmospheric the hydrographic nature and salinity intrusion
pressure differences, in January 1953 caused of the estuary.
Permanent barriers have been built across generation. A similar and larger structure was
Cardiff Bay, South Wales, and the Tees Estuary also built at Annapolis Royale in the Bay of
and Wansbeck estuary, North-eastern England, Fundy, Canada, thus using high tidal ranges in
in order to increase the amenity value of those both areas. There are proposals for more
areas. The closure of Cardiff Bay has raised major stations in the future. China and the
water levels, covered extensive mudflats per- USSR also have small stations in operation,
manently, and created waterfront areas more while they prepare for larger schemes and in
desirable for housing. The Wansbeck barrage Britain many estuaries have been assessed with
was designed to create a lake, in which the sea- regard to their energy generating potential.
water exchange is only at highest tides or, dur- In the 1980s and 1990s, many UK estuaries
ing the winter, when the barrage is partially with a tidal range in excess of 5 m were consid-
opened. The lake was designed to increase a ered and there were detailed proposals and
water sports area although siltation since the Environmental Impact Assessments (EIA) car-
barrier was built has reduced water depth suf- ried out for tidal energy barrages across the
ficiently to restrict such an amenity use. The Mersey, Duddon, and Severn estuaries. It was
Tees barrage was opened in 1995 in order to suggested that the Severn estuary, with a 16-m
create a high amenity urban area. tidal range, would be able to supply up to 20%
The use of tidal power in macro-tidal estu- of Britains electricity requirements.
aries has long been cited as an example of An energy-generating barrier would be
renewable, environment friendly, and altern- placed in the lower reaches of the estuary and
ative energy to conventional sources, especially then it works by allowing water to flow into
now with concerns about global warming and the estuary and thus creating a head-pond
the build up of greenhouse gases. Electricite De upstream. The barrier can then be closed as
France built the permanent barrier across the the tide ebbs seawards thus creating a head
Rance Estuary, Brittany, in 1972 for tidal power pressure, which can be released as water flows
(a) Estuarine barriers/barrages Storm surge/safety (e.g. Thames, Oosterschelde, Hull)
Amenity (Cardiff Bay, Wansbeck)
Tidal power (Bay of Fundy, Rance)
Changed shape/
restriction/constriction Changed hydrodynamic Riverine inputs
regime
Changed Changed Overall Increased
wave climate tidal regime Scouring at reduction residence Freshwater
Nutrients
inlet/outlet (move to time/ inputs
lower decreased
From From Outside energy) flushing rate
Inside Changed Sediments
outside inside sediment
Tidal bounce/ Persistent
Changed
reflection Reduction in pollutants Potential
benthic
Energy Increased community turbidity flooding
Reduced Reduce
reduction tidal range saline erosion
downstream Removal of Increased
penetration Reduction light limitation siltation
as fish refuge on primary
Biological Increased
community productivity capacity as a Water
Higher
Lower change Reduced O2 pollutant sink diversion
LW
HW demand in Bathymetry mechanisms
Reduced Tidal water change (M)
delivery of squeeze Drying of
marine salt marsh Increased primary Pollution
sediments production uptake,
into estuary storage and Estuarine
Removal Loss of effects basin
Wave of bird wetland Potential filling
erosion of feeding area productivity eutrophication
shores (macroalgal mats,
microalgal blooms, Waste reduction
Increase of fish Increased benthic or diversion Reduced
feeding area agricultural enrichment) techniques (M) estuary life-
land span
(Mmitigation or
compensation measures)
Overall reduction of Nutrient controls (M) (NBnot all features result from
intertidal productivity Habitat all types of barrage).
+ carrying capacity creation (M)
(b)
Interference with Remaining socio-
biological migrations Interference with economic repercussions
estuarine/marine uses/users
Low in
Turbine Increased operation
Barrier employment
damage Shellfish
Coastal defence
Fishing and High in
fisheries construction
Fish Obstruction to Different demands Estuary
seasonal and with area
spawning Infrastructure crossing
migration
Industry
Housing
Marine Egg
juveniles Shipping/navigation
Swimbladder + stripping
physical Recreation/amenity
(external)
Loss of feeding/ damage Pollution
Passage and port dispersal /
nursery areas interference assimilative
capacity Water sports
Diadromous Dredging
fishes impacts
Population effects in
estuary/freshwater/ Need for locking
coastal areas system
Change to natal
river populations
Amenity/
Requirement for recreation
fish pass (M)
Figure 6.27 Estuarine barrages and barriersconceptual model of impacts and consequences: (a) changed shape and riverine inputs, (b) interface with migrations,
uses and users and socioeconomic repercussions.
E S T UA R I N E U S E S A N D U S E R S 135
back through the barrage. The water movement Most notably, the estuary will become lower
can turn turbines either just on the ebb or both energy in nature and thus suspended material
on the ebb and flood. The presence of a barrier will have greater opportunity to settle out
across the mouth of an estuary has the potential and thus improve the water transparency.
to create many changes and adverse impacts in This could remove the light limiting control on
the estuary, the adjacent coastline, and the areas primary production within the hypernutrified
upstream and inland. The main impact is the areas, which in turn could increase primary
change to the shape of the estuary and thus production. Unless nutrient inputs into the
the distortion of the hydrographic regime. The estuary are reduced this would increase the
changed wave climate will prevent sediment possibility of symptoms of eutrophication
movement from the seaward side although the occurring. The increased residence time and
barrage will retain river-borne sediments that decreased flushing rate will affect water qual-
enter the estuary. It may increase wave-induced ity through the greater retention of any persist-
erosion inside the estuary where the water ent pollutants from the catchment, which
levels are kept higher than previously. Some of previously would be eventually exported from
the tidal energy does not enter and so results in the estuary.
tidal bounce which can increase erosion along In addition to the effects on the natural struc-
the coast (Fig. 6.27). ture and functioning of the estuary, there is
As shown in earlier chapters, fish and shell- interference with estuarine uses and users, for
fish migrations into estuaries are an important fishing both inside and outside the estuary, and
and central biological feature. Any barrier will for navigation and shipping. Industry, agricul-
interfere with fish and shellfish migration, tural, and urban areas upstream will have to
both by preventing migration into the estuary improve their pollution control mechanisms
and also by causing damage. Pressure changes and either prevent pollutants from entering
as organisms move through the barrage open- the system or divert any discharges around
ings and turbines will cause external damage the barrage. Despite this, there may be added
such as scale stripping, eye hemorrhaging, benefits for recreation and amenity, with the
and internal damage to swimbladders in creation of water sports areas as shown on
round fish, especially pelagic species such as the Rance estuary, and for infrastructure and
clupeids. Eggs carried on the underside of communications when a bridge crossing is
shrimps and crabs will be stripped by the incorporated into the barrage.
pressure changes. Several of the above adverse effects can be
The restriction of the tidal wave both enter- mitigated against although the concern about
ing and leaving the estuary will change tidal remaining impacts may prevent other bar-
regime by reducing the tidal amplitude in that rages being built. Despite this, the Rance tidal
high water is not as high, whereas low water power generating barrage has proved success-
is not as low as previously. This will keep low ful and the estuary, while changing from its
intertidal areas covered longer, thus restricting previous state, is considered to have reached
feeding by wading birds (but allowing fish a new equilibrium.
longer to feed), and cause upper areas to be
uncovered for longer perhaps causing drying
6.12 Other uses
out of salt marshes and reedbeds. Hence it is
likely that the carrying capacity of the inter- Salt extraction has long been practised on estu-
tidal areas will be adversely affected although arine areas, especially lagoons in warmer coun-
this has not been fully quantified. tries. For example, the Ria de Aveiro, the major
The barrage is likely to change the sediment- northern Portuguese lagoon, has long sup-
ary and water quality regime in the estuary. ported salt extraction in pans thus providing
136 T H E E S T UA R I N E E C O S YS T E M
the salt to produce salted cod, the classic of birds within the estuary may be the result
Portuguese dish bacalau. Compartments built of anthropogenic organic matter, which has
in shallow areas of the lagoon are allowed to displaced natural organic inputs. In Chapter 8
fill with water, which then is allowed to evapo- we will discuss the ways in which estuarine
rate, and the salt collected. Hence hypersaline conservation may be managed.
waters are produced, which changes the estu-
arine fauna from brackish tolerant to high
6.13 Conflicting uses
salinity tolerance.
Mention should be made of other positive We can thus see that the uses of estuaries by
uses of estuaries, such as for recreation or mankind are many and varied. In almost
conservation. However, where these require every estuary a variety of uses may be occurr-
human intervention to improve areas for ing, and many of the uses described are not
these then there may be adverse effects. mutually exclusive. Obviously some conflicts
Recreational interests in estuaries range do arise between the different users of the
widely; the sheltered waters are used for estuary. The requirements of fishermen or
sailing, swimming, or wind-surfing, often conservationists for clean undisturbed waters
accompanied by the construction of specialist may conflict with an industry or city council
harbors or marinas. Again the construction looking for the cheapest disposal site for waste
of any structure will change the hydrographic material. Despite the many uses of an estuary,
patterns, the sedimentary regime, and thus for most people living in cities their local estu-
the biota. An excessive number of boats will ary is the closest natural ecosystem that they
add sewage and antifouling paint residues to can visit or observe, since the resilience of
the waters, although the most polluting estuarine fauna has meant that it has survived,
antifouling paint has been banned on small whereas other natural habitats, such as the
("25 m) boats. countryside in all developed countries, have
Many people are attracted to the tranquillity been irrevocably altered.
of estuaries for bird-watching or simply escap- In this chapter we have seen the many ways
ing from their fellow men. Ornithologists have in which mankind uses and abuses the estuar-
in particular become a powerful pressure ies of the world. In the next chapter we will
group, campaigning for the conservation of consider how we can measure the effects of
the remaining parts of estuaries. In many those activities and then in Chapter 8 we can
cases, however, the presence of large numbers explore the management of the estuaries.
CHAPTER 7
137
138 T H E E S T UA R I N E E C O S YS T E M
discharging a certain type of waste into an from the affected area, animals within the
estuary? or what will be the effect of building sediments may burrow deeper, and stationary
a barrage across an estuary and how big will animals such as mussels may simply shut
that effect be? their valves to wait for improved conditions.
All organisms respond to environmental Depending on the animal and the cause of the
change from whatever cause by a series of stress, the animal may recover its metabolic
responses ranging from subtle metabolic functions, showing acclimation, or individual
adjustments to dramatic changes such as animals may be genetically selected to survive
escape or death. The latter may be the con- the stress, showing adaptation. Alternatively
sequence of the stressor not being removed or the stress may cause death or serious impair-
the organism does not have an ability to cope ment of normal functions, such as growth
with it. Figure 7.1 indicates the time-related or reproduction, leading to changes in popula-
sequence of the effects of environmental stress tions and communities. Such structural and
on marine and estuarine fauna. The first event functional responses can be detected by
of importance to the organism is the detection biological sampling of the components of the
of change by the sensory receptors, followed estuarine ecosystem.
by metabolic adjustments and or behavioral Because of this sequence and set of
reactions. Mobile animals can swim away responses, we can regard any effects of human
activities as acting at one or more levels of bio-
logical organization (Table 7.1) in which a
Stress
change at one level may, if the stressor is not
stopped or the biological system cannot cope
with the effect, then go on to affect a higher
Nervous system
detection
Escape level. Proceeding through these levels, the
time of response to environmental change
decreases and the inherent variability in the
Changes in system increases. For example, an organism
enzyme activities will respond rapidly to a change in water
quality whereas the community will take
longer to show changes; a cell is much less
Metabolic and variable in its functioning than a community.
physiological function These features have to be considered when
designing any sampling and survey program
Recovery Impaired
Table 7.1 Effects at differing levels of biological
organization
Acclimatization Death or
1. Biochemical/subcellular level
changes in
populations and 2. Physiological effect
communities 3. Change in the individual
Genetic selection 4. Changes in the population
5. Community structural change
6. Community functional change
Adaptation
7. Whole ecosystem effects
Figure 7.1 Schematic sequence of the effects of There is an increase from 1 to 7 in the complexity (and thus
environmental stress on estuarine animals. (After variability) of the system, the time for response and thus the
Blackstock 1984.) time required for the detection of effect.
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 139
and when interpreting the data. An effect of method should be precise in indicating change
a pollutant in a cell will eventually affect and have a large response compared with the
the whole organism if the cell cannot deal variability (i.e. a high signal to noise ratio)
successfully with the pollutant. Hence a suite but in order to produce a cost-effective tech-
of techniques are required to produce data to nique, which can be employed in manage-
indicate and measure change at each biolog- ment, there should be a rapid response after
ical level although those data are only useful contact with the stressor. In the case of pollu-
once converted into information. tion, the test should be applicable to a wide
Information regarding changed estuaries is range of phyla and their life histories should
required to answer focused questions: demonstrate a doseresponse relationship.
Some techniques are suitable for use for a gen-
Do the indigenous populations reflect the
eral survey, whereas others are a diagnostic
hydrophysical characteristics of the estuary?
test to give a detailed investigation in a hot-
Do the populations and growth rates differ
spot. Finally, the method should be relatively
from similar areas?
straightforward and at a reasonable cost.
What is the trophic system and how do any
Despite the many techniques being developed
organisms link with the general food web of
for assessing change, it is, however, necessary
estuaries and the adjacent coastal areas?
to be cautious as all methods have problems,
What is the structure of the communities
many are in early stages of validation and are
and how do their populations function?
not necessarily suitable as working tech-
How have the health of the individuals or the
niques. In many cases, the sources of variabil-
populations been affected by mans activities?
ity in the method have not been determined or
Have any environmental or ecological qual-
quantified accurately.
ity standards and objectives set for the estuary
The science of impact assessment has been
been met?
complicated further by the use of interchange-
A wide variety of techniques has been deve- able terms: the studies can be separated into
loped to study the changes, which occur in surveillance and condition monitoring, com-
estuarine organisms in response to human pliance monitoring, and diagnostic testing,
activities, including pollution, ranging from although different management protocols
laboratory-based toxicity testing procedures, may use different terms for these (see also
through to field-based population sampling Chapter 8). Surveillance, sometimes called con-
programs, and including physiological, cyto- dition monitoring, is the assessment of a small
logical, biochemical, genetic, behavioral, or large area, at one time or over time. In con-
pathological methods as well as bioassays and trast, compliance monitoring assesses those
ecologically based methods. characteristics but with a given end point in
There are many techniques suitable for mindthat is, what is the expected or
detecting change, each of which is more suit- required status of an area, its populations or
able than others for particular stressors, for communitiesin essence whether the area
example, some will detect the effect of heavy complies with what has been decided as the
metals from a polluting discharge, whereas preferred status. Condition and compliance
others may detect community change through monitoring may also be termed operational
sediment changes caused by creating an monitoring and, if the results are used to check
obstruction in an estuary. The techniques whether any legal licence has been met, also
should show a consistent relation of the Statutory monitoring. Diagnostic testing, also
response to individual or species pathology termed applied research, a trigger-exceedence
and it should be known whether the response study or investigative monitoring, can then be
is to either specific or general stressors. The used to determine the cause of change.
140 T H E E S T UA R I N E E C O S YS T E M
An essential
chemical testing of pollutants as an alternative. Chronic
testing may, for example, look at behavioral
Harmful changes, reproductive success, or metabolic
changes responses of animals subjected to low doses of
a pollutant. The use of low-level, long-term
testing is essential if we are to fully understand
the problems caused by dilute concentrations
Acceptable Safe limit of pollutants, however, they are more difficult
changes to administer, control, and reproduce than
A non-essential chemical
acute tests. Thus, despite the drawbacks, acute
Dose
testing remains the principal method for the
Figure 7.2 The response of a typical population to laboratory assessment of pollutants, which lead
changes in the dose of essential and nonessential chemicals. to guidelines for the disposal of toxic materials.
The position of the dotted line on the arbitrary vertical axis Although estuarine animals are often used,
varies with the toxicity of the chemical, and for some poisons, for example Corophium, Nereis, and Macoma
e.g. carcinogens, the dotted line may not exist, there being
a continuous gradient of response with concentration. The have all been greatly used in toxicity testing,
intercept of the dotted line with the response curve indicates there is an increasing use of macroalgae in
the safe dose limit. (After Pascoe 1983.) toxicity tests, both in the laboratory and in the
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 141
field. For example, mesh bags containing in power station and near chemical industry
excised discs of the green alga Enteromorpha or outfalls. Finally, regulators, especially in
the growing tips of the brown alga Fucus North America, have made large use of a
vesiculosus can be located in effluent concen- bioassay Microtox using luminescent bacte-
trations and then the viability of the cells or ria in which a measure of the decrease in their
the growth of the tissue can be measured as light output gives an indication of the toxic
a response. nature of any wastewater.
Lawrence and Hemingway 2003 for further environmental and management relevance,
details). The release of stress proteins may repair especially where food species are used, as
some of the damage caused by pollutants. well as the relative ease of measuring the
Binding (sequestration) to another molecule often higher levels of pollutants in the tissue
also reduces the bioavailability of pollutants, than in the water column. The degree of
which can lead to either excretion or storage. bioaccumulation has repercussions for the
For example, Heavy Metal Binding Proteins, a organism itself, its predators (including man
form of metallothionein proteins, which can in the case of commercial fishes), and its prog-
bind metal ions, become induced when there eny. Because of this, bioaccumulation studies
is exposure to high metal concentrations. The are widely used by statutory bodies charged
presence of these detoxification mechanisms with assessing and protecting the estuarine
or the bioaccumulation of the pollutant will environment.
indicate recent exposure. Accumulated metals The assessment of stress and the degree
may be stored in specific organs and slowly of contamination of estuarine fauna at the
excreted. In many vertebrates, including fish, subpopulation level require the use of common,
the main storage organ is the liver, which can
therefore be expected to give a better indica-
tion of recent contamination, whereas muscle
tissue may give long-term uptake and storage
Pathways within organism
(Fig. 7.3). (invertebrate or vertebrate)
At the whole-organism level, pollution may Food, water, air
act directly by increasing mortality, or indi-
rectly by causing changes in behavior such as
impairing reducing an organisms efficiency
of foraging ability thus reducing food uptake Transport molecule
Cell membrane
and thus somatic production. Hence, in turn, (gill and gut tissue,
lung, integument)
these effects on individuals can reduce popu-
lation growth, or produce population decline. Haemocyanin, red blood Transport by circulating
All of these mechanisms at cellular and indi- cells, plasma proteins, body fluids (haemolymph,
metallothionein blood or pallial fluid)
vidual levels use energy and resources, which
are therefore not available for production thus
creating a population effect. In turn popula- Enzymes, metalloproteins, Target organ
tion responses will produce effects at the com- respiratory proteins, (liver, hepatopancreas,
munity and ecosystem levels, hence the need metallothioneins and digestive gland,
polypeptides kidney, etc.)
for monitoring exercises to link the levels of
biological organization. This whole field of
biomarkers, used to indicate responses to Metallothionein, protein Storage organ
complexes, elemental metal, (kidney, liver,
pollutants, is rapidly expanding and gives inorganic concretions hepatopancreas,
great scope for detecting an early-warning of green gland)
pollution effects.
Into urine, faeces, Excretion
Bioaccumulation studies can additionally { concretions, shell (kidney, gut, gill)
be conducted to indicate both sublethal deposition
stresses within individuals as well as contam-
ination in estuaries (Phillips and Rainbow
1994). There are many benefits in the use Figure 7.3 Fate of trace metals taken up from the
of organisms over other methods for monitor- environment and diet within marine organisms. (After Engel
ing aquatic contamination, for example, the and Brouwer 1984.)
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 143
representative indicator (or sentinel) organisms of pollutants, hence a larva is more susceptible
that can be used for bioassay or bioaccumu- than an adult and an embryo is more suscep-
lation studies. Bioaccumulation monitoring tible than a larva. Their small size and
programs require species to be used, which often unprotected surface, which thus gives
fulfill several criteria: them a large surface area to volume ratio and
thus a large surface area in contact with any
being sessile or sedentary and thus repres-
entative of the study area
abundant in study areas, easy to identify 7
and sample and providing sufficient tissue for
analysis 4
able to tolerate wide ranges of contaminant
0
Solution
concentration and of physico-chemical vari- 0.3
ables such as salinity, thus even allowing an 0
Suspended
experimental approach to measure uptake 0.3 material
provide a simple correlation between the
0
Microplankton
tissue concentration and the ambient levels. 1.5
0
Most importantly, the accumulation strategy Meioplankton
1
of a potential biomonitor for a particular 0
contaminant should be known, for example, Sediment
5
whether it will take up pollutants in the partic- 0
ulate form common to highly turbid estuaries, Macroalgae
1 (seaweeds)
or from solution because of their solubility and 0
lipophyllic nature. Species such as estuarine Suspensionfeeders
(mussels etc.) 1.5
marine macroalgae such as Fucus vesiculosus 0
and Enteromorpha, the suspension-feeding Depositfeeders
marine mussels Mytilus, and the estuarine (polychaetes etc.)
0.2 0
fishes: the flounder, Platichthys flesus, and Epibenthic scavengers
the eelpout, Zoarces viviparous have many of ? and predators (crabs,
3 shrimps, etc.)
the characteristics listed above necessary for 0
sentinel organisms. Monitoring of the health Pelagic fish
1
(sprat, herring, etc.)
of estuaries will require the use of a sentinel 0
Estuarine demersal
organism or at least estuarine residents with a
6 fish (flounder,
small home range, hence reducing the uncer- eelpout)
0
Marine demersal fish
tainty in the interpretation of the data and ?
(cod, whiting)
increasing the link between the health of the 0
biota and the ambient stressors. For example, Wading birds
as sediments receive and store many pollu- (knot etc.)
tants then organisms feeding on these or on
Longannet area, contaminated mid estuary
their infauna are vulnerable, for example, the
Port Edgar area, cleaner lower estuary
flounder P. flesus, or wading birds (Fig. 7.4).
All values are milligram per kilogram dry weight
except solution where values are nanogram
7.2.4 Individual health and external and per liter; ? = no data.
internal body condition
Figure 7.4 The concentrations of mercury in components of
It has long been acknowledged that younger the Forth Estuary, Scotland. (Data from Elliott and Griffiths
organisms are more susceptible to the effects 1986, figure from Elliott and Hemingway 2002.)
144 T H E E S T UA R I N E E C O S YS T E M
waterborne pollutants, their place in the effect relationships with respect to fish health.
upper water layers while planktonic and their In addition, it may be difficult to determine
high metabolic rate, which requires the rapid whether an observed abnormality is due to
movement of materials into the organism, all a stressor acting on an individual or on its pre-
increase the uptake of pollutants. This has led vious generations and, especially in estuaries
to the production of developmental bioassays and for mobile species such as fish, whether it
such as the oyster embryo bioassays, which is operating in the area where the animal was
measures the rate of development and the caught or elsewhere. This is particularly
success of development in reaching the important with migratory fishes that use the
D-shape characteristic of most bivalve larvae. sea and or freshwaters as well as the estuary.
The health of individuals and the effects of The flesh condition of an estuarine organ-
pollutants discharged into estuaries may be ism, an index of fatness, and the relationship
shown by the presence of morphological between weight and size or volume, and its
changes. These have been used widely and are overall individual health may also provide
potentially valuable in pollution monitoring valuable information on the health of the
programs and especially with high-profile system. The condition factor varies within a
estuarine organisms such as fishes (Elliott and population, especially between life stages,
Hemingway 2002). Within fishes, for example, but also within feeding status and thus
morphological disorders range from mild skin it reflects emaciation created by spawning,
discolorations to major skeletal deformities poor food conditions, or overwintering. Thus
such as blemishes, lesions (raised areas of skin, any natural or anthropogenic stressors which
which may or may not be broken or infected), affect the energy balance will affect condi-
lymphocystis-type nodules (generally found tion, for example, the technique of assessing
on the dorsal, ventral, and anal fins and the scope-for-growth.
caudal peduncle), fin rot, eye-deformity,
mouth ulceration, and skeletal deformities
7.2.5 Molecular and genetic techniques
such as scoliosis (spine curvature). Because of
this, it is valuable to record routinely morpho- Molecular techniques are also included in the
logical anomalies during surveys of fish com- suite of approaches available for determining
munities. These give a simple qualitative change in the estuarine systemfor use for
analysis of the response to pollutants and other stock discrimination, that is, differentiating
stressors within the area, and they may alter between populations, and for assessing stress
the structure and functioning of the popula- damage. The chronic or occasional presence
tions although the effects are difficult to quan- of any substance in the environment can
tify. There may be changes in the reproductive exert a selective pressure upon organisms
capacity, or success, of the organisms, that is, and the pollutants may directly damage the
fewer number of eggs may be produced, or genetic material thus producing mutations
fatal deformities may arise within the larvae as (Beardmore et al. 1980) (Fig. 7.5). In addition,
a result of contaminants within the environ- within estuarine environments, genetic
ment. There will, however, be variations in changes to populations due to escapees from
their incidence due to season, age, migratory aquaculture are also possible. While there is the
patterns, and species as well as that due to need to determine the responses (as effects) of
external stressors. The latter may be anthro- populations in adapting (or attempting to
pogenic, for example, pollution and contam- adapt) genetically to stresses caused by human
ination, but also abnormal levels of natural activities, this has not been studied in estuaries.
features such as salinity and temperature. As Similarly, mutational changes have not been
such, it may be difficult to determine cause and studied as the result of estuarine stress.
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 145
Possible concentration of
Mosaics Neoplasia Gametes pollutant through food chain
Cell death
Some inviability
Some inviability
The speed of response of the population is edulis and the harpacticoid copepod Tisbe
related to the intensity of the selection pressure have been found to have a complex genetic
applied and by the ability of the population to population structure, which may result in their
respond to it. This in turn is influenced by inter- tolerance to environmental stressors found in
nal factors such as genome size, number of estuaries. Other genetic assessments are now
chromosomes, amount of recombination, popu- being carried out, for example, to determine
lation size, and generation time. Beardmore the effects of escapees from the culture of
et al. (1980) suggested that the following meth- salmonids in fjordic systems and imported
ods of response to polluted conditions are oysters in estuaries, particularly as these
possible: will have been reared especially for culture
(Gosling 2003). In order to produce a reliable
an organism may react entirely phenotyp- monitoring technique, the main needs are:
ically, usually in its growth rate or by modifi-
to investigate populations exposed to pollut-
cation of its reproductive output
by an increase of one or more genotypes or
ants to establish the effects on fitness, con-
sibling species at the expense of others. genital anomalies, resistance to disease, life-
span, fertility, etc.
These aspects have not been specifically stud- to determine the variation, in time and
ied in estuaries, although estuarine resident space, of the gene pools of populations living
species such as the common mussel Mytilus in clean and in polluted environments, and
146 T H E E S T UA R I N E E C O S YS T E M
under laboratory conditions; to analyze the of the cohorts. Growth can be determined by
genetic basis of any observed adaptive shifts several methodsby marking organisms
in phenotype across a range of species and releasing them before re-capture and
to assess the occurrence and distribution re-measurement, such as used extensively with
of mutagens in the marine and estuarine fishes, especially in freshwaters, but also with
environment sedentary invertebrates such as estuarine and
to test both singly and in combinations the marine bivalves. The use of growth ring discon-
effects of these mutagens on a representative tinuities in scales or otoliths of fishes and shells
range of species of bivalve molluscs can be used to determine age
to establish a monitoring scheme to screen at each size. Similarly, growth can be estimated
appropriate species for mutation and allele either by analysis of lengthfrequency his-
frequency changes. tograms or from the mean size of an organism of
a known age at the time of sampling. If one or
more particular size groups (cohorts) can be dis-
7.3 Population responses: Population
tinguished in a lengthfrequency histogram of
dynamics, growth, mortality rates,
a sample and followed through successive sam-
and population models
ples, the modal sizes plotted against time will
If there are sufficient changes to individuals of produce a good estimate of growth (for further
a given species and these are not compensated details see Crisp 1984). Alternatively, if all indi-
for by the organisms then they will lead to viduals of a species in a sample can be aged, the
population changes. An analysis of the popu- mean length of each age group can be used to
lation dynamics, that is, the relative sizes of estimate annual growth or growth between suc-
age stages in the population, will indicate if cessive samples over a shorter time span.
younger and more vulnerable stages have Once growth as size or weight at each given
been adversely affected. There is the need to age has been determined then it can be
measure the effects of activities on predator summarized in numerical models, which may
prey relationships and oscillations, especially be theoretical or empiricalthe former are
in relation to birds and fishes using the estu- designed to describe the sigmoid growth
ary. For example, if pollution or other stress curve followed by most organisms whereas
within an estuary removes a preferred prey the empirical models merely give a best fit line
size then this will affect a predator. to the observed growth at any time or place
Growth and mortality of individuals and their (see Bagenal 1978; Elliott and Hemingway
effects on population dynamics are an important 2002 for further details). While growth may be
measure of determining whether an estuarine reduced in estuaries as the result of pollution,
area is functioning as is expected given its envi- it may also be reduced in the case of primarily
ronmental characteristics. The data can then be marine species such as mussels due to the
used within population models aimed at pre- reduced salinity. The reduction in population
dicting the yield. These techniques have been size, that is total mortality, is necessary to
most widely used on fish populations. The determine the loss of organisms through
population structure of a species can be natural or anthropogenic stressors. It can be
described after measuring a representative set of expressed numerically in two different ways
organisms and then plotting a sizefrequency the actual mortality rate (A), as the fraction
histogram. The presence of the modes within of organisms present at the start of a period
that histogram will indicate the average sizes of of time which die during that period and where
cohorts and then changes in the frequency and the survival rate s is (1 ! A), and as the instan-
average sizes on successive sampling occasions taneous mortality rate (Z), an exponential rate,
will indicate the mortality (survival) and growth of reduction in numbers (see Gee 1983).
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 147
_
7.3.1 Biological production productivity (e.g. P : B $ 2.5) derived from
other studies can often be used to calculate the
Biological production is a valuable parameter
production from the mean biomass data. This
and will indicate the nature of the area, the
may only be an approximation as this ratio
success of an area supporting a population,
depends on the species, the age of individuals,
the success of the populations themselves, and
and the environmental conditions; it is often
the material available for higher and detrital
underestimated when juveniles are concerned,
foodwebs. For example, where there is the
although Elliott and Taylor (1989) found this
need to determine the carrying capacity of an
value to be suitable for estuarine invertebrate
intertidal estuarine mudflat for wading birds
species.
then it is important to know the amount of
Although there are difficulties in estimating
food produced. The production may be deter-
production, its values may be used with biomass
mined by species (individual, cohort, and
data, to assess the functioning, as the transfer
population) or by communities (by adding the
of material during feeding, of an area. Most
production by all the age groups of each
methods for estimating the production of estu-
species in the estuarine assemblage) (see Crisp
arine populations and communities require
1984 for further details). For an individual, the
input data on growth rate, mortality rate,
physiological production (P) depends on
and abundance at least at one time, although
the food consumption and the metabolism of
production may be extrapolated for groups of
the species and equals the sum of Pg (tissue
species (trophic levels or communities) from
growth), Pr (production of genital products),
production estimates of other parts of the food
Ps (secretions such as mucus), and Pe (dead
web (notably the primary producers).
tissues).
The population production available for
predators (including Man) is termed yield and 7.4 Community level assessments
can be determined using the cohort produc-
As shown in the previous chapter, any change
tion based on the growth as size or weight of
to the overall nature of the estuary such as
individuals belonging to the same ageclass
hydrographic and sedimentary features, will
(the latter is termed the Allen curve). This
change communities, especially the bed com-
indicates the relative decrease in abundance of
munity. Changes to sediment type or quality
a cohort, through natural mortality together
and water quality will sooner or later affect
with the increase in average size, and thus
bed community structure, hence these remain
weight, of the cohort members. Production
the mainstays of assessments of estuarine
may also be estimated using growth and
environmental change. All laboratory-based
average biomass data where the latter is the
assessments of pollutants should be supported
organic matter contained in the tissues, often
by comprehensive field-based studies of the
measured as ash-free dry weight or measured
actual effects of the pollutants on estuarine
as wet weight and then calculated using
organisms. For a field assessment of the effects
conversion factors.
of a pollutant discharged into an estuary,
Production values are most accurate when
or of the change to the community through
based on a large sample size which, given the
engineering works, the principal method is to
nature of estuarine populations, will be of
collect samples of animals or plants from areas
a few dominant species. _ Production can
or times believed to be affected, and to com-
be estimated using P : B ratios, derived else-
pare them with samples from areas or times
where or from the literature, together with an
believed to be unaffected. This has given rise
average biomass from the area under study
to the BACIPS approachthe Before After
(Elliott and Taylor 1989). A theoretical value of
148 T H E E S T UA R I N E E C O S YS T E M
Design Examples
Transect Down the shore (high, middle, low; tidally standardized profile or fixed points)
Transect Based on the direction of a hydrographic feature, for example, tidal movement
Skilled eye For biotope assessment and quantification
Ellipse Aligned according to the dominant current direction but with latitude for movement
perpendicular to that direction
Circular grid Based on a null hypothesis of no change in any direction around an interest point
Circular grid Hexagonal variant of central treatment area surrounded by a control area
Box grid Null hypothesis of no spatial effect but with regular spaced samples (e.g. based on
lat./long. points, intersections)
Point sampling Random sampling points across an area
Point sampling Stratified random in which a given number of sites are arranged in each habitat type
(stratum)
Treatment area vs control With replication, based on well-defined locations of both areas
Increasing spatial separation On the basis that the effect with decrease in intensity away from the source, hence a grid
or transect with station separation increasing (log, semi-log interval)
Temporal surveys To cover change with time as well as different impacts over time, for example, for a pipeline to
cover preconstruction, postconstruction/preoperation, postoperation,
postdecommissioning
Tow For nekton or benthos (using a dredge, trawl, plankton net) to give geographic coverage
(e.g. axial in estuary) over a known distance
Tow For nekton or demersal fauna (using a dredge, trawl, and plankton net) using tidally
standardized tows
7.4.1 Numerical analysis of estuarine data diversity of the assemblage, and the distribu-
tion of individuals and biomass among those
There are many numerical methods relevant
species. Estimates of abundance indicate
to the sampling and interpretation of estuar-
the size of the populations of each species
ine communities, especially invertebrate com-
although their accuracy is dependent on the
munities (e.g. see Krebs 1989; Table 7.3) and in
methods used and on samples being represen-
particular the estuarine benthos in relation to
tative of the whole population.
pollution or other stressors (Elliott 1993, 1994).
Many features can be used to describe an
Univariate methods are used for the interpre-
estuarine community and interpret changes
tation of the primary and derived community
in relation to human impacts, environmental
parameters such as species richness, abund-
variability and biological processes (see
ance, biomass, and diversity of the commun-
Table 7.3). Community or assemblage analysis
ity. Increasingly multivariate methods are
can be carried out using the primary commun-
used any data set in which a set of attributes
ity variables of taxon (usually species) rich-
(e.g. species presence or abundance) has been
ness (S), abundance (A), or biomass (B) data.
measured across a set of samples, which may
The different measures can be used at the
be sites, stations, or areas.
binary (presence absence) level (taxon rich-
The community structure refers to the
ness) or as quantitative data (abundance and
species composition of an area, which can
biomass), depending on the reasons for the
be described according to the number of
study and the methods used. Quantitative
species and the amount of biomass present, the
150 T H E E S T UA R I N E E C O S YS T E M
Table 7.3 Concepts and techniques for detecting and indicating change in estuarine and marine communities
Source: Elliott and Hemingway 2002, references for the topics therein.
data allow a higher level of analysis, with the techniques such as cluster analysis can be used
determination of the rarity of a species. to define communities, that is, as a set of attrib-
In addition to this taxonomic approach to utes such as species which are occurring
community structure description, analyses of together, whereas other methods define con-
biomass size spectra, and functional guilds tinua of both taxa and environmental features,
can be used and are an increasingly valuable for example ordination analysis. Any area
means of interpreting community structure. sampled will easily produce a matrix of attrib-
The functional groupings or guilds can be utes (e.g. species presence, abundance, and/or
based on the ecological preferences of species, biomass) against sample (e.g. site, salinity
their reproductive strategies, and their feeding regime, and area). There are a variety of meth-
modes (Elliott and Hemingway 2002 for ods available for multivariate community
details). Other functional aspects can be deter- analysis such as ordination and cluster analy-
mined in communities, such as using ratios to ses (Gauch 1982; Begon et al. 1990; Jongmann
compare taxonomic types or organisms with et al. 1987; Elliott 1994) and gradient analysis.
different habits. These allow arranging the different species
The primarycommunity variables can be abundance along a selected gradient or allow
used to derive other secondary variables such communities, or sites, to be ordered along con-
as diversity (e.g. H ShannonWeiner index) tinua according to similarities in their species
and Biomass (B/A) and Abundance ratios composition and abundance. These enable
(A/B). These univariate data can then be inter- the analysis of community attributes, taxon,
preted and interrogated using conventional abundance, or biomass, at a series of stations
hypothesis-testing techniques (e.g. ANOVA) or sites, and can be performed as either Q- or
although complex community data, such as R-mode. The former describes similarities
the abundance of each species in a sample, are between different stations according to the
often more suited to analysis by multivariate taxa present, its abundance or biomass, while
statistical techniques. Multivariate numerical R-mode classifications group taxa according
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 151
100 s
Cumulative % dominance
ma
ss
m ber
Bio Nu
rs
be
rs um ass s
be N as
50 um om om
N Bi Bi
Moderately Grossly
Unpolluted polluted polluted
0
1 5 10 50 1 5 10 50 1 5 10 50
Species rank
Figure 7.6 Hypothetical k-dominance curves for species biomass and number, showing
unpolluted, moderately polluted, and grossly polluted conditions. On the horizontal (x) axis,
the species are ranked in order of importance, using a logarithmic scale, and on the vertical
(y) axis, the percentage dominance is plotted, using a cumulative percentage scale. (After
Warwick 1986.)
to affinities in their distribution. For further numerical treatments all, however, require
information see Elliott and Hemingway further effort and testing in estuarine habitats.
(2002). Trends in all of these features can be deter-
There are several graphical techniques to mined and used to indicate the functioning of
indicate change such as the S-A-B curves the community in a defined area at a specified
(PearsonRosenberg model) described in time; these can then be compared to data from
Chapter 6. As a further example, Warwick other times and places (Moss et al. 1982).
(1986) assessed the pollution status of a ben- Thus it is possible to determine the normal
thic community by plotting the distribution of situation from a variety of sites and popula-
the numbers of individuals among species tions and then show if the study area is differ-
the abundancebiomass-comparison (ABC) ent, and if so, whether this is due to natural or
(Fig. 7.6). In unpolluted conditions the benthic anthropogenic factors. Many techniques have
community will be dominated by one or a few been used to separate the effects of natural
large species, each represented by rather few disturbance stress (such as found in estuaries
individuals. In polluted situations, benthic and sedimentary unstable areas) from anthro-
communities become dominated numerically pogenic stress and this has resulted in the
by one or a few small species; each repre- widespread use of indices of environmental
sented by many individuals. However, cau- quality. The methods are required to indicate
tion has to be exercised as an influx of small both the human and natural importance of the
organisms during recruitment can also create areas, habitats, and populations.
similar patterns in the community. Community studies, and especially those
There is an increasing amount of software, involving the macrobenthos have long been the
especially as integrated ecological and biolog- mainstay of estuarine impact assessments,
ical packages, used to calculate this wide often because its features and taxonomy are
range of numerical techniques and indices. well known and its response to pollution and
The increase in computing power has pro- stress are well described. The model for change
duced a greater and easier potential for ana- (see Table 6.2) indicates the nature of the
lyzing large data sets. The development of change and thus the characteristics of the
sampling, the treatment of replicates, the use fauna, which should be studied within an
of power analyses in determining the level of impact assessment. These changes underpin
sampling required and the cross-calibration of especially the ABC and SAB techniques. As
152 T H E E S T UA R I N E E C O S YS T E M
mentioned throughout this book, the presence to detect severe sources of pollution, or hot-
of large numbers of small organisms within a spots, but more subtle change requires a
few species, which is characteristic of stressed wide variety of approaches, and the skill and
and polluted conditions, is also typical of ingenuity of the scientist may be severely
natural estuarine populations and so this has to tested. Such skill is especially needed for estu-
be borne in mind during impact assessments. arine studies, since a wide range of natural
In order to assess the effects of disturbance environmental variables, such as salinity, tem-
from pollution, dredging or other activities, perature, or sediment type, occur alongside
field experiments can be established in estuar- any perturbations induced by mankinds use
ies, whereby an area is deliberately disturbed and abuse of the estuary.
or defaunated and the recovery of the In managing estuaries, we are often inter-
fauna observed. In such a study in Alewife ested in their ability to maintain the high pred-
Cove, Connecticut, USA, Zajac and Whitlach ator populations, especially the fishes and
(1982a,b) found that the response to distur- birds, and so we are interested in knowing the
bance was very variable, and that no set pat- trophic structure of the area. For example, we
tern for estuarine re-colonization succession need to know if land-claim removes an area
could be detected. Thrush and Roper (1988) supporting large populations of primary con-
have also shown that re-colonization studies sumers or if sediments change sufficiently
are difficult to assess for pollution monitoring. such that an area no longer supports deposit-
feeding infauna. Hence the loss of an estuarine
area through land-claim or a change in its tidal
7.5 Integrated estuarine assessments
height or sedimentary nature through accre-
and ecosystem changes
tion, will influence its foodweb. Because of
While some techniques are used in isolation, this, analyses of stomach contents and feed-
it is now becoming more common to carry ing behavior will determine the use of areas
out integrated environmental assessments in and the repercussions of loss of areas. Those
which a combination of complementary tech- analyses can give prey preferences, prey
niques is used. For example, the sediment importance, and the amount of prey required
quality triad approach (Long and Chapman to support the population. They can also give
1985) combines three measures: of community niche breadth, for example, using diversity
health, as seabed quality as shown by benthic indices to show what number of prey items
community analyses, of individual health as are taken and the opportunistic or specialized
shown by an invertebrate bioassay such as the nature of predator feeding, and niche overlap,
suitability of sediments for burrowing by that is, the similarity of feeding between two
infaunal species such as Macoma, and lastly an predators either within or between species to
environmental (abiotic component) such as the indicate possible competition.
degree of contamination (e.g. the concentration We can define the ability of an area, both
of persistent chemicals in sediments). The according to the amount of space and food
latter gives a cause component to help to available, to support those higher predators as
explain the former two effect components. its carrying capacity. Hence any change in
There are many methods at our disposal for that carrying capacity, especially because of a
detecting change in estuaries. All methods for loss of an estuarine area, a habitat or a biotope,
detecting the effects of pollutants and other is of major concern to the estuarine managers.
causes of change in estuaries are a compro- A unit of the ecosystem, which is increasingly
mise between the desirable, the practicable, and recently used in assessing the quality and
the affordable, and the reliable. It is generally managing ecosystems is the biotope. This term
true that almost any method will be able may be regarded as describing the habitat
H U M A N - I N D U C E D C H A N G E S I N E S T UA R I E S 153
and its component and dominant organisms. important to realize that a measurement is
For example, middle estuarine salt marsh, mus- never exact but it can be precise and accurate
sel bed, or mudflat biotopes include the main within certain limits (types of equipment used,
physical features of the habitat and the domi- skill of the scientist, etc.). Precision is the same
nant organisms (possible Spartina, Nereis, and measurement or class achieved on each occa-
Mytilus respectively). The presence or number sion that an observation is made, for example,
of biotopes in an estuary, their size, and degree the weight of an object on a particular balance
of functioning can then be used as a measure of or the same identification of a taxon by one
the health of an estuary. As such, biotope taxonomist. In contrast, Accuracy is the correct
mapping is increasingly used to determine measurement or class that is obtained on every
these features and may be carried out at either occasion that an observation is made, for
a lower level (a Phase 1 survey) in which all example, the weight of a standard calibration
assessments are done in the field and only the material or the correct identification of a taxon
obvious organisms are noted. Alternatively, if by any taxonomist. Hence, measurements can
greater data are required then a Phase 2 survey be precise without being accurate, for example,
will be carried out whereby samples are taken a badly maintained balance could be precise
for the full identification and enumeration. without being accurate. AQC/QA is designed
to ensure that all the errors in measurement
are known, accounted for, or minimized, for
7.6 Monitoring, surveillance, and
example, a laboratory has well-trained staff,
survey design
well-documented methods, and good equip-
In physiological terms the term homeostasis ment. In this way, data obtained from one
is used as the ability to cope with or absorb estuary can be combined with those from any
change by modifying the organisms func- other area thus allowing changes in one area to
tioning but without resulting in a long term, be put into a wider context.
deleterious change. We may also think of As the widest example of an integrated
estuarine populations, communities, and the study focused on a defined estuarine activity,
ecosystem as exhibiting homeostasisthe Environmental Impact Assessments (EIA) are
ability to absorb and adjust to change without now widely required to be performed before
the effects being observed widely. As shown permission is given for each estuarine activity.
throughout this book, estuaries are highly Those EIA should have a structured approach
variable systems and thus they may have (see Glasson et al. 1999) and thus consider
the ability to absorb anthropogenic change as each component of the activity and each part
well as natural change and still function as of the natural system likely to be impacted. It
expected. The aim of the estuarine scientist should determine the impact or an activity at
and manager is to detect whether those char- a place at a certain time and as performed in a
acteristics have been changed, thus a rigorous certain way, that is, a site and time-specific set
approach to monitoring is required. This is of effects. For example, the construction and
particularly important, as in some cases data operation of a pipeline into an estuary will
produced by the monitoring may be used in involve clearing a channel, bringing materials
legal discussions or proceedings. to the site, perhaps dredging an area, building
In any discussion of monitoring, it is the pipeline, and discharging the waste from
necessary to mention the quality of the data it. These aspects may affect different parts of
produced especially where those data are the water column, estuarine bed topography
to be used widely. Because of this, there is and sediment, and faunal and floral popula-
an increasing need for analytical quality con- tions. Hence the EIA should quantify the
trol (AQC) and quality assurance (QA). It is spatial and temporal impact of each part of the
154 T H E E S T UA R I N E E C O S YS T E M
sequence on each natural component and then whether a component affected by an activity
determine the size of that natural component within an estuary is important in an interna-
before determining whether the activity will tional context, such as a population of overwin-
have a long- or short-term effect or will affect tering wading birds, or only in a local context,
a small area or large area. Hence monitoring is such as an estuarine intertidal mussel bed. The
required to quantify each of these aspects. size of the affected component needs to be deter-
The monitoring is required to determine the mined, for example, the number of hectares
spatial degree of effect, that is, the extent, and covered by the mussel bed or whether the wad-
the temporal severity of effect, that is, the dura- ing birds are #1% of the European population
tion. The spatial degree can be the amount of (i.e. internationally significant). The final conclu-
area affected and the spatial importance of a sions from the matrix of impact (Table 7.5), after
component affected (Table 7.4). For example, it has been applied to the activity, produce as an
I N R D L Nil
Negative
Irreversible in 0
long term
Reversible in 0
long term
Reversible in 0 0
short term
No known impact
Positive
Short-term !!! !!! !! ! 0 0
benefit !
Long-term !!! !!! !!! !! ! 0
benefit !! !
Scales: I, International; N, National; R, Regional; D, District; L, Local and Nil, no known value.
Table 7.5 Example of use of impact scales for an estuarine pipeline construction
Element Activity Aspect Component Potential Intrinsic !ve !ve !ve #ve #ve
size of ecological irrev. rev. rev. short long
target value long short term term
term term
end point, a graphical and simple indication of partitioning by fish and bird consumers of
degree of effect, whether positive or negative. available production. Studies are likely to be
This then needs to be communicated to inter- applied, for example, attempting to determine
ested bodies. For example, if it is decided that whether the indigenous fish populations in
constructing a barrier will have a negative coastal areas are as they should be given the
effect, which is irreversible in the long term on hydrophysical characteristics of the area and
a population of international significance, such whether the benthic populations differ from
as overwintering wading birds, then a table similar areas. There may be the need to know
of impact will indicate this by giving six filled the trophic structure, the estuarine foodweb,
circles. and its functioning. Finally, there is often the
Following this process, it is then necessary need to know how the populations have
to indicate, in the EIA, whether any mitigation been affected by mans activities and whether
measures can be employed, for example, can a Ecological and Environmental Quality Object-
pipeline be constructed in such a way as to ives (EQO) and Standards (EQS) are being met.
minimize an effect on the bed of the estuary.
If mitigation is not possible, then managers Decision level 2: Monitoring definition
will have to determine whether compensa- In order to create a focused approach, the type
tion measures can be produced, such as by of monitoring needs to be defined. For exam-
creating or restoring a habitat elsewhere as ple, whether Statutory Monitoring is required
compensation for a estuarine habitat, which and, if so, to determine who is responsible for
had been destroyed by an activity. carrying it out. This will follow identifying,
for the human impact, the Uses and users of the
estuary or coast at risk and those who are interested
7.6.1 Decision Tree
in the results of the monitoring. This determines
The nature of monitoring and surveillance where the demand is for the monitoring and
required to answer the questions posed why it should be carried out. Within estuarine
throughout this chapter may be defined using a and coastal management, this will produce a
Decision Tree. Surveillance is regarded as matrix of effects on the different users and uses.
gathering data and then indicating change a The next decision is the Detail of monitoring
posteriori whereas monitoring should relate to required, which covers from the subjective
a pre-defined end-point, for example, the dis- and qualitative, through skilled-eye tech-
solved oxygen levels of waters required to niques to the fully quantitative and statist-
protect estuarine fish migrations. Once a pre- ically rigorous surveys. A definition of the
defined limit of 5 mg l!1, for example, is set then Components at risk will then indicate what
monitoring can be directed toward detecting aspects should be monitored, which in turn
the time and place where this is not reached. dictate the methods to be used. The scale of
the monitoring then requires to be considered,
Decision level 1: Definition of main first the Spatial extent, as the area to be mon-
questions and hypotheses itored, which involves defining station posi-
This first step will decide whether studies are tions, the strata to be sampled and often the
carried out to provide answers to either general station positions with regard to hydrographic
or precise questions, to test hypotheses, and characteristics. Second, the Duration and
to generate hypotheses. For example, many Frequency of sampling, which illustrates the
studies merely determine the usage of estuarine temporal component and the length of cam-
or marine intertidal and subtidal areas, the paign. Finally, the sampling may be required
respective usage of estuarine and coastal areas to detect changes in the biological and
and their associated freshwater areas, and the environmental components, hence the Degree
156 T H E E S T UA R I N E E C O S YS T E M
tion on natural features and their inherent nents, either as a single component (hard
spatial and temporal variability benthos, soft benthos, hyperbenthos, nekton,
condition monitoring, to determine if the plankton, etc.), or a combination (two or more
status of the estuarine individuals, popula- components)
tions, and communities are in a pre-defined ecotoxicological individual health, such as
158
T H E M A N A G E M E N T O F E S T UA R I E S 159
goes ahead and to decide what is the desired scientists had to be aware of a large number of
condition for the estuary. Because of this, there aspects. It was recognized that although indi-
is a need for indicators to describe the status of vidual activities should be controlled, the main
the ecological ecosystem elements, as in the aim is to manage the whole environment, and
DPSIR approach (Chapter 6), which summar- to protect habitats and species and the func-
izes the source of problems, the causes and tioning of estuaries. This created an overall
effects against the background characteristics, philosophy that incorporated sustainability,
and the responses required by Man in order precautionary action, the integration of all
to manage the estuarine area. The options for environmental aspects, and democratization
sustainable and cost-effective management (wide consultation, open, fully accountable)
then have to fulfil six tenets: being environ- (Table 8.1).
mentally sustainable, economically viable, In managing estuaries, we can take what
technologically feasible, legally permissible, may be called bottom-up or top-down
socially desirable, and administratively achiev- approaches. For estuaries, this is from the
able. For example, dredging of an estuarine (bottom-up) management of species and habi-
waterway will be required by society in order tats, through activities, and whole estuaries to
to make the area suitable for navigation but it (top-down) management of catchments and
has to be carried out using appropriate and coastal sea areas. For legislation and adminis-
cost-effective methods, in accordance with local tration, this is from (bottom-up) local and
laws and international obligations but without regional initiatives, national laws and bodies,
having an adverse and unacceptable environ- supranational ones such as the European
mental effect. This chapter aims to illustrate Union (EU), to (top-down) conventions and
many of these features. agreements covering regional seas and global
Over the past half century, there has been an aspects.
evolution of the thinking regarding environ-
mental protection relating to pollution and dis-
charges into estuaries. In the 1960s there was 8.3 Planning and designations
recognition of the problem, following public
8.3.1 Management and protection of
awareness as indicated by Rachel Carsons
species and habitats
book Silent Spring. The 1970s saw an increase
in ecological monitoring and the creation of There is an increasing awareness of biodiver-
end-of-pipe controls with the use of long-sea sity as the protection of all members of the
outfalls and preliminary treatment. The 1980s biological community although often these are
had some better treatment and tighter end- the high profile species, the rare, fragile, and
of-pipe controls whereas the 1990s led to the endangered species or the species important
holistic approach. The latter brought in in the overall functioning of the system.
Integrated Pollution Control (IPC) and Management initiatives are then required to
Environmental Management Systems (EMS), protect each of these species.
and clean technologies for industries. It The International Union for Nature Con-
acknowledged that pollution moves across servation (IUCN) has created lists (published
land, air, and water and the need to stop pollu- as the Red Data Books) of endangered species,
tion at source rather than merely treat it after it which may then form part of the species con-
had been produced. The 2000s completed the servation measures created by countries. In
cycle by showing a more complete recognition the case of European estuaries, these aspects
of the problemthe acknowledgement that an have been incorporated into the European
ecosystem approach was required in which all Commissions Habitats Directive, which in
features were considered and managers and Annex I give habitats for designation and
T H E M A N A G E M E N T O F E S T UA R I E S 161
Sources of legislationnational, To include specific laws that relate to estuaries or have been implemented to
supranational, etc. include estuaries; to determine whether estuaries are controlled waters and
identify the competent authorities
Types of agreement, protocols To include agreements, from inside and outside states, which may or may
national, supranational, global not lead to estuarine legislation
Identification of priorities for To determine which are the greatest potential or actual problems within an
action estuary, such as which pollutants are important, what habitats and species
should be protected, what water characteristics need to be assessed
Authorization for potentially To decide which of those priorities for action require a licence, for example, do
harmful actionlicensing, etc. estuarine industries require integrated pollution control authorizations, or which
dredging schemes require permits?
Precautionary principle, action To decide which of the activities should not be permitted on the basis that they
are likely to cause harm to the estuary
Impact assessment and action For each activity which has the potential to harm the estuary, to be subject to an
EIA, that assessment must include all anthropogenic and natural characteristics
outside of the estuary
Roles of physics, biology, and The need to ensure that all of the interlinked estuarine processes are included
chemistry in assessment and and to use information from one discipline (e.g. hydrography) to inform
control and the integration of decisions about another (e.g. biology)
environmental aspects
Role of monitoring and To ensure that the surveillance of the estuarine features is carried out
surveillance (i.e. determining spatial and temporal trends), but also that monitoring per se,
in which the assessment is carried out against a preconceived threshold/level of
acceptable change, is performed in a cost-effective manner
Sustainability for use vs non-use To realize that within a working estuary that supports many economic activities
such as navigation and industry, it is necessary to ensure that they are carried
out in a sustainable manner (i.e. one activity does not jeopardize another either
now or in the future). Similarly, non-(economic) use such as maintenance of
wildlife also needs to be sustainable
Carrying capacity of natural and It is argued that an estuary has a carrying capacity for each aspect, for example,
anthropogenic features the number of wading birds that it supports, or the amount of waste discharged
into it without creating harm, and that management has to be geared to ensuring
that these occur
Democratization, accountability, If there are changes to an estuary, management of its features, or the awarding of
openness permits for activities, then all stakeholders have to be involved
Polluter pays principle Estuarine users, such as dischargers putting waste into an estuary or navigation
monitoring, clean up, prevention authorities carrying out dredging, are responsible for the effects to be
monitored, clean up to be effected, new technology to be used
Classification schemes, The estuarine areas have to be classified according to their nature and their
performance indicators quality, and the amount of estuarine area within each quality class
can be used as a performance indicator to determine whether management
is being effective
Environmental management Within a costbeneficial and environmentally sustainable framework, industries,
systems, clean technologies, waste and developers are made to be responsible for ensuring that environmental
minimization. problems either do not occur, are mitigated (made less severe) or compensated
(that the loss of one habitat is compensated for by creation elsewhere)
162 T H E E S T UA R I N E E C O S YS T E M
protection. Its Annex II lists species important taken to protect the species. Wider measures
in a European context and which may be may be required for species subject to com-
requiring protection because they or their habi- mercial fishing; for example, No-Take Zones
tat are endangered. While there are few estuar- may be designated in certain areas.
ine (and nearshore marine) species on the list, Estuaries are not highly diverse areas in
there are 12 species of fishes (Table 8.2). Most terms of numbers of species but they have a
of these are diadromous species, that is, they unique and important functioning and they
migrate between freshwaters and the sea, and have habitats not found elsewheresuch as
so any barriers, whether physical in the case of mudflats, mangroves, salt marshes. There
weirs and barrages, or chemical, such as dis- is thus the need to protect the estuary as a
solved oxygen sags, will endanger the popula- habitat or the habitats within estuaries and to
tion. Because of this, European Member States protect habitats supporting large numbers of a
have to produce management plans to protect few species (or even a single species). Because
these species. Hence there is the increasing of this, habitat-specific Biodiversity Action
view that in estuaries and other marine envi- Plans (hab-BAP) are created to indicate how
ronments, in order to protect the species it is much of a habitat is desirable and how much
preferable to look after the habitat and ensure of that habitat should be protected on a
that a species is not taken excessively, as in national scale. Hence, the European Habitats
fishing. Following this, the species will look Directive Special Areas of Conservation (SAC)
after itself. Once the species requiring protec- have designated a type of habitat (e.g. sand
tion have been designated then species- banks) or a group of habitats (estuary, coastal
specific Biodiversity Action Plans (sp-BAP) embayment) thus providing protection for
can be created detailing the measures to be those habitats. Of particular relevance to
Table 8.2 The species of European estuarine and migratory fishes deemed to be fragile, threatened, or endangered
estuaries is the European designated Special Any noncompliance with the permit or licence
Protected Areas (SPA) for the protection of the will lead to legal proceedings. The main fea-
habitat for supporting birds. Similarly, sites tures given within a licence are indicated in
designated worldwide under the Ramsar Boxes 8.2 and 8.3.
Convention are aimed at protecting wetlands Different approaches can be adopted to
as bird habitats. bring about pollution control. Either, an
Although the protection of species and Environmental Quality Objective (EQO)
habitats is the main aim in management, it is policy, usually linked to an Environmental
likely that human activities will affect those Quality Standard (EQS) policy, or a Uniform
and so mitigation measures have to be Emission Standard (UES) policy. The essence
employed to minimize effects. Where this is of the EQO approach is that an estuary (or any
not possible, for example, if a new dock water body) can disperse, degrade, and assim-
removes mudflats, then compensation will be ilate pollutants that enter it. It is based on the
required such as the creation or restoration concept of assimilation capacity, which is
of habitat elsewhere in order to replace any the capacity of a body of water to cope with the
habitat lost. This feature has now led to the effluents discharged into it. Apart from any
process of habitat-banking in which a deve- chemicals, which bioaccumulate, it assumes
loper may buy land elsewhere in the estuary that the solution to pollution is dilution! The
prior to development at one site taking place. EQO approach is concerned to define the over-
The management of habitats on a larger scale all conditions of the habitat and then to select
in the United Kingdom is now being proposed for each estuary a set of permissible opera-
through the creation of Coastal Habitat tions, or levels of pollutants, discharged in
Management Plans (CHaMPs), which may order to achieve that condition. The EQSs are
indicate the overall management of habitats in
relation to those affected by the estuarine
activities.
Box 8.2 Conditions imposed within a
8.3.2 Management and licensing of activities Consent (Licence/authorization/permit)
In most areas, permission from a relevant for a discharge into estuarine waters
statutory authority is required in order to per-
(1) types of outlet and number of diffuser
form an activity. That permission may be in ports;
the form of a legally binding document, which (2) source of material discharged (type of
may be called a permit, a licence, an author- industry);
ization, or consent. For example, in the United (3) pH, BOD5, suspended solids, temperature;
States, a NPDES (National Pollutant Discharge (4) emission levels (as concentrations
Elimination Scheme) permit is required before amounts per liter) (based on toxicity
discharge of a land based effluent, whereas in information ) ability of receiving areas to
the United Kingdom an authorization is degrade/disperse/assimilate);
required for a complex industrial plant to (5) maximum loadings of key parameters
allow the integrated management of dis- (amount per day, month);
(6) volume of discharge per day and per hour;
charges to the air, into the estuarine waters, or
(7) position of sampling points within plant;
onto land. A consent to discharge is required (8) information on record keeping and good
by a sewage treatment plant in order to dis- practice;
charge its effluent into nearby water course, (9) toxicity-based element (Direct Toxicity
whereas a licence is required to dump dredged Assessment).
material taken from a navigation channel.
164 T H E E S T UA R I N E E C O S YS T E M
disposal of effluent must be controlled, either movements, and substratum type. Thus if a
by the reduction in volume, or by an improve- low energy area is protected and maintained
ment in quality of the waste so that it does not then it will support a mudflat containing large
impede the other uses. At the mouth of the numbers of prey for overwintering birds and
estuary, where there is plenty of water, this resident fishes.
objective might be attained by better disper- The last objective, of those given here, is
sion of existing waste through the construc- designed to eliminate contamination of, for
tion of a long sea outfall waste pipe. In the example, shellfish by microbes, such as fecal
inner estuary where there is less dilution coliform bacteria, which may be injurious to
water available, the attainment of the same any person eating the shellfish. It also aims to
objective might involve the construction of eliminate contamination of fauna by any per-
new sewage-works or treatment plants. The sistent accumulative pollutant, such as heavy
standard of effluent permitted may thus be metals (especially mercury or cadmium) or
different in different parts of the estuary. In Organohalogens that could taint them, affect
order to achieve the defined EQOs, a higher predation, or harm anyone eating them.
EQS would generally have to be applied to a In certain cases, the meeting of these objec-
sewage or industrial discharge at the head of tives is determined by testing for specific and
an estuary, than to one at its mouth. numerical standards, as EQSs, which may be
The second objective requires that the estu- created for licences or protocols. The EQS
ary be of sufficient quality to sustain the values may be defined as the maximum allow-
diverse components of the estuarine ecosys- able concentration, and or the 95%ile value
tem that we have described in earlier chapters. (95%ile $ the value that is not exceeded for
In particular, no discharge to the estuary 95% of the values taken), and or the annual
should remove the fauna in the vicinity of the average. These values may be included within
discharge, and reclamation and other engi- the licence or the discharge conditions may be
neering schemes that destroy the benthic habi- set allowing for the dilution, dispersion, and
tat should be avoided. assimilation within the receiving waters to
The third objective, to allow the passage of ensure that the EQS are not exceeded.
migratory fish, such as salmon or eels, at all Toxicity testing (see Chapter 7) has a role for
states of the tide, is designed to ensure that estuarine management in helping to deter-
the oxygen concentration, as a measure of mine the permitted levels of toxicant in the
water quality, is greater than 50% saturation discharge. For example, 1/10 or 1/100 of the
at both high and low tide, and in both sum- 96 h LC50 may be used as the safe level for
mer and winter. In many estuaries, this object- the discharge of a toxic effluent, these factors
ive is readily attainable in winter at high being termed application factors. Such
tides, when there is a large volume of water, application factors have been criticized,
but more difficult to attain in summer at low because exposure to sublethal concentrations
tide, when there is a smaller volume of water of a pollutant may influence physiological
and there will be less oxygen present due to processes such as reproduction or develop-
the water being warmer or more saline (see, ment, but not cause death. Despite such valid
for example, Fig. 6.4). This objective must criticism, the acute toxicity test and applica-
therefore be defined in terms of the most tion factors remain in use, since such tests are
problematic season or condition, not the most easy and reliable to perform, and provided
lenient. that the most sensitive life stage of an animal
The fourth objective is to ensure that the ele- is used, do give valuable guidelines for
ments of the biota are as expected given the determining priorities in the management of
areas hydrophysical characteristics, its water industrial effluents discharged to estuarine
166 T H E E S T UA R I N E E C O S YS T E M
ecosystems. Toxicity testing has long been Pollution discharged into an estuary is man-
used in determining the levels of single com- aged by determining the state of the estuary,
pounds for discharges. All discharges, how- the quality of its waters, and the condition of
ever, convey a mixture of materials, so it is its inhabitants at all times of the year. Dis-
necessary to licence the complex effluent and, charges entering the estuary from industries
because of synergistic and antagonistic or domestic sewage-works or the diffuse
effects, there is an increasing use of a direct sources from the river input should be identi-
toxicity elements in the licence (called toxicity fied and analyzed and, in the case of the
based consents or Direct Toxicity Asses- anthropogenic discharges, licensed by the rele-
sments). The toxicity element would therefore vant authority. The characteristics of the
be a clause in the licence whereby the whole licence (Boxes 8.2 and 8.3) control the levels
effluent would be tested (Whole Effluent entering the estuary and ensure compliance
Testing) and deemed to fail if its LC50 was with international and national standards and
likely to be breached after dilution in the agreements, such as whether the discharge
receiving waters. contains any of the priority substances of
When the policies of management in estuar- concern as well as the local situation of the
ies are implemented as planning objectives, discharge. The licences can then be reviewed
the concepts of EQO, EQS, and UES are per- at intervals as technology improves and
haps not as different as they first appear. In knowledge about the fate and effects of pollut-
practice they usually work together, in that the ants becomes available.
estuarine manager needs to specify the per- The US state and federal policy has gener-
mitted standards in any effluent entering the ally followed the EQO EQS approach. In the
estuary, in order to achieve the objectives, 1970s and 1980s, the UK government also
which have been set for that estuary. Whether followed an EQO EQS approach, but with the
the standards set are uniform in all countries, adoption of the North Sea declaration follow-
or parts of that country, is essentially a polit- ing the North Sea Ministerial Conference in
ical decision. The role of the estuarine scientist 1987, and the adoption of the precautionary
is to advise the politicians as to what stand- principle, it has moved to a UES policy for
ards are required, either in the effluent, or the those substances regarded as causing the
receiving waters, so that the estuary is fit for greatest problems, that is, being the most
its uses. In the context of estuarine water qual- polluting. These substances, given on the
ity, the concentration of a particular con- so-called Red List, will be controlled by cut-
stituent in a discharge may not be the most backs in all emissions irrespective of the body
important factor; it is the rate of entry to the of water to which they are discharged. The
estuary that matters. This is obtained by multi- precautionary principle could lead to zero
plying the concentration by the rate of flow to emissions given that all substances are likely
give the load. The assessment of loads is, to have an impact sooner or later, at some
however, often difficult since many of the place and in some concentrations. The precau-
sources of pollution are subject to wide varia- tionary principle may be defined as a philoso-
tions in both flow and composition (Gameson phy whereby preventative action should not
1982). As the effluent load enters the estuary it be postponed even if the causal link between
is usual to define a zone of acceptable eco- an activity and the impact has not yet been
logical impact, or a mixing zone, which is fully confirmed.
an area where dilution and dispersion occurs. Some EQSs for estuaries and seas in Europe
The zone should be as small as practicable, are listed in Table 8.3. Such lists are ultimately
and some latitude in the EQS for this area may based on the evidence of the toxicity of the
often be deemed acceptable. different substances with mercury clearly
T H E M A N A G E M E N T O F E S T UA R I E S 167
Table 8.3 EQS for Europe. All standards expressed as Table 8.4 The Black Listsubstances which should be
microgram per liter ($parts per billion), in receiving waters banned or substantially eliminated from discharges. The list is
(dissolved), except where indicated of the Paris Convention and List I of the European
Dangerous Substances Directive
Parameter Estuary Coastal sea
Priority substances for pollution control
Dissolved oxygen #55%
pH 6.08.5 List Icertain individual substances (see below) selected
Ammonia "0.025 mg l!1 according to their toxicity, persistence, and bioaccumulation
Cadmium "5.0 "2.5 potential, with the exception of those which are biologically
Copper "5.0 "5.0 harmless or which are rapidly converted into substances
Mercury "0.5 "0.3 which are biologically harmless
Arsenic "25 "25 1. Organohalogen compounds and substances which may
Nickel "30 "30 form such compounds in the aquatic environment
Lead "25 "25 2. Organophosphorus compounds
Zinc "40 "40 3. Organotin compounds
Chromium "15 "15 4. Substances in respect of which it has been proved that
Iron "l.0 mg l!1 "1.0 mg l!1 they possess carcinogenic properties in or via the aquatic
environment a
Source: After FRPB (personal communication) and Sayers
5. Mercury and its compounds
(1986).
6. Cadmium and its compounds
7. Persistent mineral oils and hydrocarbons of petroleum
origin
8. Persistent synthetic substances which my float, remain in
regarded as the most toxic heavy metal. Such suspension or sink and which may interfere with any use of
standards for EQS of waterborne contamin- the waters
ants may not necessarily reflect the potential a
Where certain substances in List II are carcinogens, they are
harm to the fauna in the sediments. For included in category 4 of this list.
example, mercury goes out of solution very
quickly as it is adsorbed onto suspended
particles, which then accumulate in sediments.
There is therefore a move toward future EQSs carefully controlled quantities. These priority
being based on sediment concentrations. It lists (Tables 8.4 and 8.5) are based on Annexes
should also be remembered, as discussed in of the 1992 Paris Convention (formerly the
earlier chapters, that the toxicity of any single linked Oslo and Paris Conventions, OSPAR)
compound may vary greatly according to envi- which has been adopted by many countries, to
ronmental factors within the estuary such as control land-based discharges and dumping
temperature and salinity, or may vary when into the NE Atlantic area. This approach is sim-
the single compound is part of a cocktail of ilar to the global London Convention (formerly
many chemicals. the London Dumping Convention) of which
A common theme in international policies to the United States is a signatory and thus has
regulate the discharge of toxic substances to adopted the substance of these lists. The Black
marine and estuarine waters is the recognition List is also List I of the European Dangerous
of two types of contaminants in order to prior- Substances Directive, and the Grey List is List II
itize control. First a Black List or List I of of that directive. These directives are the
substances, which should be banned or sub- parent directives setting out the framework
stantially eliminated from discharges, and for the control of waste discharges in Europe,
second a Grey List or List II of substances and are updated by the addition of daughter
the discharge of which may be permitted in directives as new compounds are identified as
168 T H E E S T UA R I N E E C O S YS T E M
Table 8.5 The Grey List. Substances whose discharge may be permitted in carefully controlled quantities. The list is of the
Paris Convention, and List II of the European Dangerous Substances Directive
List II contains: substances belonging to the families and groups of substances in List I for which the limit values (referred to in
Article 6 of the Directive) have not been determined; certain individual substances and categories of substances belonging to the
families and groups of substances listed below; and those which have a deleterious effect on the aquatic environment, which
can, however, be confined to a given area and which depend on the characteristics and location of the water into which they are
discharged.
Families and groups of substances referred to in the second indent
1. The following metalloids and metals and their compounds:
1. zinc 6. selenium 11. tin 16. vanadium
2. copper 7. arsenic 12. barium 17. cobalt
3. nickel 8. antimony 13. beryllium 18. thalium
4. chromium 9. molybdenum 14. boron 19. tellurium
5. lead 10. titanium 15. uranium 20. silver
2. Biocides and their derivatives not appearing in List I
3. Substances, which have a deleterious effect on the taste and/or smell of the products for human consumption derived
from the aquatic environment, and compounds liable to give rise to such substances in water
4. Toxic or persistent organic compounds of silicon, and substances that may give rise to such compounds in water,
excluding those that are biologically harmless or are rapidly converted in water into harmless substances
5. Inorganic compounds of phosphorus and elemental phosphorus
6. Non-persistent mineral oils and hydrocarbons of petroleum origin
7. Cyanides, fluorides
8. Substances which have an adverse effect on the oxygen balance, particularly: ammonia, nitrates
Table 8.7 Shellfish-rearing waters standards for Table 8.8 Environmental quality standards for Europe on
the State of Virginia, USA the quality of water required by designated shellfish waters
will be designed to use the best available a continued high effort by the dredging com-
technology (BAT), although this has been panies, or the beneficial use of the material. An
extended to the best available technology not example of the latter is to use dredged mate-
entailing excessive costs (BATNEEC). rial as beach nourishment or recharge onto
As shown in earlier chapters, dredging of intertidal areas as a means of countering sea-
navigation channels is a major operation in level rise and beach erosion.
estuaries and it has the potential to have a Other activities such as fisheries and aqua-
major impact on the estuary, hence its man- culture, mineral exploration and extraction,
agement is of central importance. Under and recreation are all managed to a lesser or
national and international legislation and greater extent albeit usually by different
agreements (see below), the quality of the organizations and certainly under different
material being dredged has to be analyzed for legislation and agreements. Where they pro-
the presence of pollutants. Highly contami- duce waste or polluting discharges, these are
nated material cannot be allowed to be dis- controlled by the relevant pollution abatement
persed into the estuarine or adjacent coastal legislation and procedures. For example,
waters and so needs to be contained. While organic matter produced by intensive aqua-
some of this may be put in secure landfill sites, culture can be controlled under licences either
whereby any runoff will not pollute ground- preventing dumping of pollution into the sea
waters, the degree of contamination in dredged or, as is the case in the United Kingdom, pol-
material from some estuaries has required lution from a land-based source. Aquaculture
large-scale solutions. For example, the Port of is subject to management of the persistent
Rotterdam on the estuary of the Rhine during chemicals that it uses, such as pesticides, the
the 1990s created two large bunded contain- use of medicines to combat disease and
ment areasthe Papasbiek for the most heav- through regulations controlling the use of
ily contaminated dredgings and the Slufter for genetically modified organisms or the intro-
the less-contaminated material. Both of these duction of alien species. Fisheries are man-
were land-claim areas although the former aged through by-laws, which may control the
was also sealed by a heavy duty lining to pre- timing, areas, and methods of fishing and
vent leaching of any pollutants. This solution increasingly fisheries legislation is designed to
to the problem had to be accompanied by a protect the ecosystem from the effects of fish-
reduction in pollution discharged into the eries, especially the taking of non-target sizes
Rhine such that in the long term, new dredg- and species of fishes and other organisms.
ings could safely be disposed to sea. Mineral exploration and extraction, such as
In many cases, dredgings are disposed of for oil, gas, and aggregates (sand and gravel
into the estuarine waters or out to sea with the for construction) is subject to the use of
management aim being to ensure that the licensed areas, controls on the use of the
material can be moved by the dredger faster seabed, and pollution abatement, ecosystem
than it is accumulating. More recently, nature protection and EIA legislation and agreements.
conservation bodies have been concerned that Hence, although estuarine areas may be
such a removal of sediment from an estuarine licensed for the extraction of oil and gas, the
system will have effects on the natural system, operations would not be allowed in sensitive
for example, removing sediment from a navi- areas or at sensitive times of the year, for
gation channel may draw down sediment example, when large numbers of wading
from intertidal mudflats. Because of this, estu- birds would be affected by any spillage of oil
arine managers have advocated either a greater following the exploration.
use of estuarine disposal, to keep the material Since the late 1980s, perhaps the largest dis-
in the estuarine circulation even if this requires cussion of estuarine management has centered
172 T H E E S T UA R I N E E C O S YS T E M
Camden
Sept. 1958
Chester 2
Table 8.10 Description of the five generations of Delaware Estuary water pollution control efforts. From Albert (1988)
10
Dissolved oxygen mean monthly (mg l1)
19826
8 197281
6 196271
5 Trenton Philadelphia Wilmington
geom. mean x 1000/100 ml
Fecal coliform annual
4
1973
2 1982
0
D F A J A O 1987
Months
0
0 80
Figure 8.3 Mean monthly dissolved oxygen concentration Miles
in the Delaware estuary at Chester, averaged for each
of three decades. Note the decrease in the duration of Figure 8.4 Annual geometric mean count of fecal
the critical low dissolved oxygen period with time. coliform bacteria in the Delaware estuary for 1973, 1982,
(After Albert 1988.) and 1987. (After Albert 1988.)
T H E M A N A G E M E N T O F E S T UA R I E S 175
diversity of fish increased from 16 resident in estuaries, which has been shown to be
species in the 1960s, to 36 by 1985. Improve- reversible in its effects. However, the problems
ments are still required, both to dissolved of toxic chemicals, especially those accumula-
oxygen levels, and to the control of specific toxic tive substances on the black lists, may be less
substances, but the Delaware Estuary has been readily reversible, and will continue to pose
established as a prime example of how problems to estuarine managers. The recovery
mankind can restore or repair the damage, of the ecosystem is largely dependent on
which he has caused to the estuarine ecosystem. the processes of sedimentation. Thus the
The Thames Estuary, UK, with the city of concentration of the pollutant in the water
London and approximately 10 million people column may decline rapidly after the cessation
on its banks, has experienced similar problems of discharge, but its concentration within
to the Delaware, and like it has now been dra- the sediment may continue to influence the
matically restored. On the Thames, there was a fauna and flora inhabiting the sediment. Any
steady decline in the oxygenation of the estuary re-suspension of the sediment, due, for example,
from 1920 until 1955, so that periods of total to dredging, will serve to liberate the pollutant
deoxygenation regularly occurred. Since that into the water again. The intertidal and other
date improvements, mainly to sewage-works, habitats of an estuary can utilize (or bury) many
have resulted in a substantial improvement of the pollutants that human activity introduces
in the condition of the middle reaches. The into estuaries, but such utilization (or burial)
recovery of the diversity of fish species in the depends on the maintenance of all aspects of a
Thames, as was shown in Fig. 6.9, amply shows healthy functioning ecosystem.
the success of the management program.
Fortunately not all polluted estuaries are, or
have been, as badly polluted as the Delaware 8.4.1 Monitoring for management
or the Thames, but it is also true to say that few
As an integral part of management practice,
have been as successfully treated as those two.
monitoring should occur in order to
The lesson of the Delaware and the Thames is
detect change and to determine whether the
that major improvements to sewage-works
management is having a desired effect. For
and improved treatment of industrial effluents
example, monitoring strategies in the Humber
can restore the oxygenation state of an estuary,
estuary, Eastern England, are designed to
so that the environmental quality objectives
reflect pollution abatement schemes by indus-
can be attained. In both cases the restoration of
try. As an integrated program, the Humber
the estuary has taken decades of effort, and
bioaccumulation monitoring program meas-
large sums of money. These recovery programs
ures the levels of trace metals in:
have also clearly shown that the further
downstream the discharges can be made, then a seaweed (Fucus vesiculosus) to reflect levels
the lesser will be the resultant pollution, in solution
since there will be a larger body of constantly an infaunal polychaete, the ragworm (Nereis
changing water to receive, dilute, and degrade diversicolor) to indicate the transfer of pollu-
the pollutants. In both of these estuaries the tants from a sink such as the sediments into a
discharges to the estuary have been reduced primary food source for the higher predators
to levels that were within the assimilation an epifaunal crustacean, the brown shrimp
capacity of the estuary, so that the normal (Crangon crangon), a major food source of
functioning of the estuarine ecosystem could fishes in the estuary
cope with the material entering it. an estuarine resident fish, the flounder
Both the Delaware and the Thames were (Pleuronectes flesus), to reflect uptake from
extreme problems of organic enrichment sediments and infauna
176 T H E E S T UA R I N E E C O S YS T E M
and a marine, commercially taken fish, the Resources Act 1991; it extends IPC, BATNEEC
Dover sole (Solea solea). replaces to some extent BPEO and Best Practical
Means, using the EQO/EQS approach; note
In addition, the sources of such pollutants are
that harbor authorities also have power to
measured in the water column, sediments,
establish by-laws to regulate dumping of pol-
riverine inputs, industrial discharges, and
luting substances under Rivers (Prevention of
domestic discharges. This monitoring is com-
Poll.) Acts 1951, 1961.
plemented by monitoring of the intertidal and
subtidal invertebrate communities, shrimp
EU legislation
populations, resident fish community and
The largest source of environmental legisla-
migratory salmonids, the water quality, inputs
tion for European countries is the EU and in
and ambient levels of persistent organics,
many ways the environmental direction mir-
hydrocarbons and nutrients, flow characteris-
rors that of other developed countries such as
tics, and effluent loadings.
the United States and Australia. There are
In addition to the above type of monitoring,
common themes throughout the environmen-
most major estuaries in North-west Europe
tal legislation of these large groupings of
are also subjected to a harmonized monitoring
member states and so the EU is given here as
scheme. This is under the auspices of the Paris
an example of environmental thinking amongst
Commission and is designed to produce an
a large block of autonomous states.
inventory of materials entering the European
The EU environmental policy is mainly
shelf sea areas.
through Regulations (which are directly applic-
able as law and are generally used to achieve
precise goals) and Directives (which establish
8.4.2 Legislativetypes of instruments
the goal and results to be achieved but allow
National flexibility and leave the method of imple-
Before managing an area, it is necessary to mentation to the state); thus the latter give
define the area of competence, that is, who is the some measure of discretion by each state (this
competent body responsible for implementing is termed subsidiarity). A Directive is appli-
legislation (such as the Environment Agency, cable in all member states (or those to which it
EA in England and Wales) and to define the was issued) and members must submit to the
area to be controlled (hence controlled waters EU Commission the national laws imple-
under COPA). In the United Kingdom, since mented to meet directives (this is called their
its implementation in 1985, the dominant piece enabling legislation). The member state is
of anti-pollution legislation is the Control of required to issue letters of compliance
Pollution Act 1974: Part II Pollution of Water detailing steps to incorporate the directive
(COPA II). This is aimed specifically at poison- into its national legislation. If the performance
ous matter and solid wastes and trade and of a member state is unsatisfactory then it is
sewage effluent to rivers, estuaries, and other reported to the European Court of Justice and
water courses. Its main provisions set down although member states can be stricter in envi-
the offence of pollution into relevant, specified, ronmental regulation than the EU they cannot
and controlled waters, it controls Trade be less strict. Even within this short descrip-
Effluents, deals with applications for consent to tion, North American readers will see the close
discharge; ensures publicity and freedom of similarities between the EU and the federal
information; and is enabling legislation for EC system in the United States.
Directives and OSPARCOM. It has been Tables 8.11 and 8.12 summarize the EU
updated and revised by the Water Act 1989, Directives of relevance to estuaries. Many of
Environmental Protection Act 1990, and Water the earlier directives had a sectoral basis, that
Table 8.11 Relevant EU Sectoral Directives and their impact on Estuaries
Dangerous A Framework Directive, provides for a system of licensing Reduction of inputs to catchments,
Substances ) by consent before discharge of listed substances; allows control on use of polluting
Daughters for Uniform Emission Standards (UES) and EQS Approaches; materials, prioritization of the
76/464/EEC etc. basis of List 1 and List 2 compounds and derivatives. most polluting materials; this
The directive requires an inventory of sources and probable links to the IPPC and Water
sources from users and sewers (data required on: max. conc. Framework Directives (see Table 8.3)
(g l!1), max. flow. (m3 day!1), max. Monthly load (kg), and will eventually be replaced
estimated load discharged (kg year!1), plant handling capacity
Urban Waste-water Provision of sewage treatment to minimum standards, Changes to estuarine discharges;
Treatment based on the population equivalent (PE), as a fixed often long sea outfalls following
91/271/EEC amount of organic matter from the population and only preliminary screening from
as a means of converting organic industrial STW for larger urban areas
discharges to a PE and type of receiving area
Integrated Basis in controlling waste from prescribed industrial Ensures that all major chemical plants
Pollution processes to all media (land, water, and air) and alongside estuaries have
Prevention and acknowledges a transfer of pollutants across media; aim Environmental Management Systems
Control 96/61/EC to ensure that industries bring in BAT to control waste and control pollutants from all sources
Titanium Dioxide A single industry relevant Directive, designed to reduce The largest European TiO2 plants are on
78/176/EEC polluting discharges, to ensure a licensing system, estuaries such as the Humber and Tees
inventory of discharges and monitoring system estuaries in the United Kingdom
Shellfish Growing Designed to monitor and improve waters in which This ensures that areas are designated
Waters and shellfish grow, ensures monitoring of water quality, within estuaries and coastal water
Health especially aesthetic and polluting aspects, in areas bodies and then any polluting
79/923/EEC suitable for shellfish (to be repealed by the Water discharges within those areas are
Framework Directive) treated
Shellfish Areas designated as Class A (shellfish can be taken and Has led to areas designated in many
Harvesting sold for consumption without treatment), Class B (shellfish estuarine areas, for example,
91/942/EEC require placing in purification waters (recirculating Morecambe Bay in the United
system) before being suitable for selling), Class C (shellfish Kingdom, where mussels are taken
require re-laying for a given period in order to improve
quality), Class D (shellfish too contaminated or waters
too polluted to allow consumption)
Quality of Set quality standards for recognized bathing areas; set Ensures that estuarine beaches are
Bathing Waters out frequency of sampling and procedures to be used; sampled on 20 occasions during
76/160/EEC and define circumstances in which directive may be waived; the bathing season (e.g.
revisions require reporting at regular intervals. Parameters: total MaySeptember) and any breaches are
coliforms, faecal coliforms, enteroviruses, pH, transparency, tackled by improvements in sewage
Salmonella, color, mineral oils, surface-active substances discharges
(foam), phenols, odors
Quality of Requires member states to designate waters, which need Of minor interest to estuaries
Freshwaters protection or improvement in order to support fishes; it except that it is designed to
to support Fishes is primarily concerned with salmonid waters and cyprinid protect the waters receiving or
78/659/EEC, amend (coarse fish) waters; (to be repealed by the Water delivering Diadromous fishes from
91/662/EEC Framework Directive) or into estuaries
Deliberate release Controls the use of genetically modified microorganisms Aquaculture in estuaries and
and use of GMO and the deliberate release into the environment other transitional waters
90/219/EEC, of genetically modified organisms; using genetically modified
90/220/EEC repealed the latter including those used in salmonids and shellfish
by 2001/18/EC aquaculture will be controlled
178 T H E E S T UA R I N E E C O S YS T E M
Environmental On the assessment of the effects of certain public and All major estuarine developments
Impact private projects on the environmentEIA to identify, will be subject to EIA and local
Assessment describe, and assess in an appropriate manner the planning regulations. This will
(85/337/EEC and direct and indirect effects of a project on the following: require adequate studies,
97/11/EC) (i) humans, fauna, and flora; (ii) soil, water, air, climate, mitigation, and compensation
and the landscape; the interaction between (i) and (ii); measures
material assets and the cultural heritage
Conservation of Basis in habitat protection by creating a network of Annex 1 habitats include estuaries,
natural habitats designated sites (Special Areas of Conservation, SAC) intertidal sand and mudflats and
and of wild fauna throughout Europe; SACs were to be proposed by 1995 subtidal sandbanks, salt marsh and
and flora (termed but by early 2000s sites were still being proposed and sea grass beds amongst others.
Habitats Directive) adopted as SACs. Annexes set out why and where SAC Annex 2 species include several
92/43/EEC to be designatedAnnex I gives the priority habitats estuarine fish species: sea and river
and Annex II gives the species of concern; sites are to lamprey, shads, salmon, and
have schemes and committees of management; enabling sturgeon
legislation in Member States, competent authorities
designated to use existing functions, relevant authorities
to use their powers, but cannot interfere with legitimate
users and uses
Conservation of Member States are required to take measures including Estuaries supporting internationally
Wild Birds the designation of Special Protected Areas; designation important populations of wading
79/409/EEC to maintain sufficient habitat for the support and birds have been designated as SPA
protection of all bird species, also conservation measures
for rare, vulnerable, and migratory species. SPA will be
combined with SAC to give the Natura 2000 network
of sites across Europe
Nitrateslevel, With relevance mainly to agricultural sources of nitrogen The aim is to prevent estuaries
control, use compounds, includes controls on nitrate use and runoff, from developing symptoms of
91/676/EEC aims to combat actual and potential eutrophication eutrophicationespecially
and health aspects due to excess nitrates in water macroalgal mats, toxic and nuisance
courses and groundwater; means of tackling diffuse algal blooms
nutrient inputs in order to ensure surface waters
Strategic Adopted to provide for a high level of protection of the Will ensure a more integrated
Environmental environment and to contribute to the integration of assessment of environmental
Assessment environmental considerations into the preparation and consequences due to estuarine
2001/42/EC adoption of plans and with a view to promoting developments
sustainable development
Water Framework Main Objectives of Sustainable Water policy: provision Resulting in definitions of types
Directive of drinking water, provision of water for other economic of estuaries, called Transitional
2000/60/EC requirements, protection of the environment, alleviation Waters in the Directive,
of the impacts of floods and droughts, will provide classification of quality, reference
overall framework for European, national and regional conditions against which other
integrated and coherent water policies. estuaries can be compared, and
(This Directive will repeal several others) the determination of good
ecological status for estuaries
Environmental Aim: to address effectively and efficiently site To Link with the Habitats and
Liability for contamination and the loss of biodiversity, to establish Water Framework Directives
preventing and a framework whereby environmental damage would be and related to biodiversity and
remedying prevented or remedied. Mode of achieving this to be human health
T H E M A N A G E M E N T O F E S T UA R I E S 179
is, they were designed to tackle a particular Zone Management (ICZM) strategy. The
problem such as the quality of bathing beaches Marine Strategy published in 2003 will inte-
or the discharge for a specific industry such as grate many of the marine environmental
titanium dioxide production (Table 8.11). Recent aspects given through EU policy and
years have, however, shown a movement to a legislation.
greater holistic, integrated, and ecosystem
approach with very wide ranging directives
International obligations
(Table 8.12). In addition, community-wide
Most countries have agreed, as signatories, to
agreements such as the Common Agriculture
wider international and in many cases, global
and Common Fisheries Policies now have a
declarations. These include:
wider environmental protection role. For exam-
ple, the Common Fisheries Policy now includes The UN Convention on Environment and
the protection of marine habitats and species in Development (UNCED, the so-called Earth
the management of fish stocks. Summits) in 1992 in Rio de Janeiro, in 1995 in
Within Europe, the Commission of the New York and 2002 in Johannesburg. These
European Union has been particularly active laid down the general principles for ensuring
in issuing directives for the control of pollu- sustainable ecosystems and the 1992 conven-
tion, based on the UES approach, and has in tion led to Agenda 21, which aimed to set the
recent years issued a series of standards philosophy for the twenty-first century by
through their directives. The directives are encouraging states and individuals to think
community law, but each needs a nations global, act local. The UNCED led to the
own law to enable them to be adopted, and Convention on Biological Diversity (CBD),
the nation has the choice of using the UES which provides a legal framework for biodi-
or EQS approach. In addition there are other versity conservation through the develop-
wide-ranging EU Agreements, which affect ment of national action plans to halt the
estuaries such as the Integrated Coastal worldwide loss of animal and plant species.
180 T H E E S T UA R I N E E C O S YS T E M
The International Maritime Organization the North Sea Ministerial Conferences have
(IMO), especially through its MARPOL taken place approximately every 5 years since
agreements, aims to control litter and pollu- the early 1980s. These have led their member
tion in seas, the use of antifouling paints on states to agree to estuarine and coastal
vessels and the introduction of alien organ- management measures to combat environ-
isms in ballast waters. mental deterioration and excessive use. How-
ever, there is some discussion over the value of
The UN through its Convention on the Law
these commissions in areas where they overlap
of the Sea (UNCLOS) also lays down general
with other collections of member states such as
principles on the management of the marine
the EU (see Ducrotoy and Elliott 1997).
areas and the prevention of harm to those sys-
As well as being signatories to regional agree-
tems. Amongst others, the UN Food and
ments, countries will agree to global conven-
Agricultural Organization has adopted and
tions. For example, all developed states have a
promoted several underlying legal principles
legislation and administrative system to control
to support (ICAM) Integrated Coastal Area
the dumping of wastes into the marine or estu-
Management: the precautionary principle, the
arine waters. While in Europe, the dumping of
principle of preventative action, the polluter
industrial waste was banned in 1992 and
pays principle, the responsibility not to cause
sewage sludge in 1998, the disposal of dredged
trans-boundary environmental damage, the
material requires a licence. Those countries are
rational and equitable use of natural resources
signatories to the global London Dumping
and the involvement of the public (www.fao.
Convention (now London Convention) on the
org/docrep/w8440e00.htm).
Prevention of Marine Pollution by Dumping of
Many countries are also signatories to
Wastes and Other Matter. Hence they need
regional environmental conventions and are
enabling legislation to adopt agreements under
thus members of commissions. These include
that convention. Part IV of Canadas Environ-
the Oslo and Paris Commission for the NE
mental Protection Act 1988, the Republic of
Atlantic including the North Sea (OSPAR,
Irelands 1981 Dumping at Sea Act, and the
now combined into the 1992 Paris Com-
United Kingdoms 1985 Food and Environ-
mission), the Helsinki Commission for the
mental Protection Act (FEPA), for example, pro-
protection of the Baltic (HELCOM) and the
vide for the licensing and control of materials
Barcelona Convention for the protection of
dumped into estuarine and coastal waters.
the Mediterranean (MEDPOL). Although each
of these originally was formed to prevent pol-
lution both from land and vessels, their remit
8.4.3 Administrative/organizational
has been extended since the 1990s to include
management framework
an ecosystem approach. These conventions
have produced harmonized monitoring Given the plethora of legislation illustrated
schemes to measure pollutants coming thro- above, it is necessary to consider which
ugh their estuaries and more recently have administrative bodies are responsible for
produced Action Plans and Quality Status implementing it. The estuarine management
Reports of their areas. OSPAR, notably, has organizations will implement a framework
produced strategies to combat eutrophication based on national and international legisla-
and pollution by hazardous substances within tion, agreements and protocols such as those
its estuaries and coastal regions. Similarly, sea illustrated above. This in turn has created,
areas are likely to have groupings of adjoining in most countries, an excessively complex
states in order to control activities in them, administrative and organizational structure
their estuaries and catchments. For example, for managing estuaries. The Box 8.5 illustrates
T H E M A N A G E M E N T O F E S T UA R I E S 181
Box 8.5 Case studyEstuarine and coastal management framework in Scotland (2003),
as an example of the legal complexities of estuarine management. Similar situation in
other countries
this complexity for one country, Scotland, other stakeholders but which have to link
but it is emphasized that this feature is together and cooperate in order to bring about
encountered in many others (e.g. Fernandes sustainable management. This system also
et al. 1995). Although this structure in many relies on temporal and spatial coordination
countries changes often, and usually follow- the aim to achieve the optimal phasing of
ing government changes, the structure gives management actions across places within
the main instruments for controlling and the estuary and at different times (see also
managing human activities. www.fao.org docrep w8440e00.htm).
The integrated management of estuaries is Coastal Zone Management has been
included in ICZM, which relies on hierarchical defined as A dynamic process in which a
organization, that is, where different stake- coordinated strategy is developed and imple-
holders and players link with each other. This mented for the allocation of environmental,
relies firstly on vertical coordination, which socio-cultural, and institutional resources to
is aimed at creating cooperation amongst vari- achieve the conservation and multiple use of
ous hierarchical levels of government and the coastal zone. (Gubbay 1990). Similarly,
administrationat national and central, reg- Coastal Zone Management typically is con-
ional and local. Second, it relies on horizontal cerned with resolving conflicts among many
coordinationcreating cooperation within a coastal uses and determining the most appro-
specific level of hierarchy, for example, at the priate use of coastal resources. (Sorensen
local level among and across local govern- et al. 1984). In the United States the Environ-
ment, all sectors and various stakeholders, mental Protection Agency (EPA), and the
each of which have no responsibilities for the Coastal Zone Management Act (CZMA) have
T H E M A N A G E M E N T O F E S T UA R I E S 183
been the main federal instruments for the policies as well as collate and disseminate
protection of coastline environments. Since environmental information.
1972, all 35 coastal states have participated in
the CZMA program. Each individual state has National
produced legislation for its own shoreline, Estuaries are the sites of many activities, uses
and inevitably it has proved hard to maintain and users, each of which requires to be man-
Federal consistency, so that standards for aged and consequently, this requires many
estuarine management do vary from state to organizations to be involved. See Box 8.6 as an
state. Each state also has a State EPA together example of one English estuary, the Humber.
with county environmental regulations such In many countries, the overall environmental
that a developer may need permits from control machinery starts with high level parlia-
several levels in a hierarchy. mentary of advisory bodies. For example in the
UK, the Parliamentary Select Committee on
Global and supranational the Environment, Food and Rural Affairs will
The international agreements indicated above scrutinize Government decisions and policy,
are usually created by a Conference or and inform and respond to public debate. In
Convention attended by states and then lead- turn, the Government Department of the
ing to a more permanent Commission, the Environment, Food, and Rural Affairs (DEFRA)
latter having its own Secretariat. The final is a central body with overall responsibility for
declaration, protocol, or agreement is then environmental matters within England
applicable to its signatories although the suc- mainly policy but including discretionary,
cess of the implementation of these relies on political decisions including water policy.
member states bringing in enabling legislation. There is other involvement by the Department
In addition to this, states are increasingly part of Trade and Industry (DTI), Department of
of larger, supranational blocks such as the EU Transport, Local Government and the Regions
and so it is valuable to consider the nature of (DTLR), and the Office of the Deputy Prime
the latter as an example. Minister (ODPM). These departments have the
It is arguable that the EU is the largest power to approve actions of regulatory bodies
driver of environmental protection for the and make appointments to regulatory bodies.
countries which are either members, wish DEFRA control the funding of regulatory bod-
to become members, or by choice follow the ies, including the Environment Agency, and
EU policies. The Environment and Fisheries has responsibility for the conservation agencies
Directorates, part of the Commission, together whereas, perversely, DTLR and ODPM con-
with the relevant Councils of Ministers, which trols aggregate extraction and offshore wind-
are drawn from each member state, and the power. The devolved Governments in Scotland
Parliament, create and agree the EU statutes, and Northern Ireland have created the Scottish
principally Directives and Regulations. The Executive Rural Affairs Department (SERAD)
Commission will also create and implement and in Northern Ireland Environment and
EU policies and initiatives (e.g. environment Heritage Service and Department of Agri-
action programs). The European Court of culture and Rural Development (DARDNI).
Justice, the judiciary, then will interpret the These government departments oversee the
meanings of treaties and other legislation and respective environment protection agencies
decide whether states are adhering to letter and statutory nature conservation bodies of the
and/or spirit of the legislation. In addition to countries of the United Kingdom. This complex
this, the European Environment Agency structure has been criticized for creating frag-
(EEA), which is not a body of the EU, plays a mentation, duplication, and confusion in envi-
role in helping to frame EU environmental ronmental control (e.g. Carter 1988).
184 T H E E S T UA R I N E E C O S YS T E M
Box 8.6 Case Study: Organizations involved in the management of one estuary,
The Humber Estuary, Eastern England
management, local plans, economic development, foreshore and seabed (obtain revenue from it;
emergency planning ownership)
Regional Development AgenciesRegional Associated British Portsvessel control and
integrated aerial, water, and land waste contact, encourages, and advises for economic
management, health and safety development
Internal Drainage Boardsmaintenance and Othersindustry, farmers, landowners, Sailing
The United Kingdom also has a Royal EPA, Canada and Sweden have EPA, and in
Commission on Environmental Pollution the United Kingdom, each country has an
(RCEP) which is an independent body with a environment protection body. In Australia,
nominated and appointed membership, which each state or territory has an Environment
meets on an ad hoc basis to consider and advise Protection Authority or Agency or Department.
on specific topics. The RCEP started in 1970 but In the case of England and Wales, the Environ-
previous Royal Commissions have met since ment Agency is an independent corporate
late 1800s. The 1972 Report of the RCEP con- body, which is managed by an executive board
sidered Estuary Quality and included the first nominated by the Secretary of State and funded
discussion of EQO while other annual reports by direct grant and a charging scheme. Its main
include Tackling PollutionExperiences and roles are in water resources protection and
Prospects (1984) and BPEO (1988). It is of note management, flood defence, fisheries and nav-
that the EQO adopted for the Humber Estuary, igation, pollution control-waste management
Eastern England, were based on those pro- licensing, control over transport of waste, IPC
posed by Royal Commission on Environmental processes, and the investigation and clean up
Pollution, third report 1972. of contaminated land. It aims to promote
Many countries now have EPAs, for example, conservation and the enhancement of amenity,
in the United States there are state and Federal and to assess and prevent, minimize, remedy,
T H E M A N A G E M E N T O F E S T UA R I E S 185
or mitigate the effects of pollution. Most guidelines via the National Water Quality
notably, and in contrast to its forerunners, it has Management Strategy in Australia and the New
a costbenefit duty, that is to consider the costs Zealand Resource Management Act. These
and wider societal benefits when seeking to countries have moved toward an adoption of
achieve its general duties. best practice and cooperative best management.
The UK environment agencies, under the This has required a change from control to
Polluter Pays Principle and based on the prevention, from focusing on prescriptive
Water Act 1989 and it successors, can set a regulation to that on outcomes, and an empha-
charging scheme in which each discharger is sis on cooperation rather than direction. Their
to pay an annual charge for attendance (cost Water Quality Guidelines encourage coopera-
of visits, sampling, irrespective of distance), tion between all stakeholders to maintain or
compliance monitoring (analysis and report- improve water quality. Hence they use tools
ing, reliant on complexity and determinands such as the creation of memoranda of under-
in effluent), environmental monitoring of the standing, impact assessment, catchment man-
receiving waters, and for the application and agement plans, and monitoring.
issuing of the licence. The level of monitoring
is decided by the agency and so greater costs Local
are levied when there is a more complex waste Given the plethora of organizations involved
stream, the greater effort required in setting in management of estuaries, there is the need
a licence and in monitoring compliance. As to achieve coordination. For example, regional
an extension of this Principle, monitoring is and municipal bodies are often charged
increasingly being carried out by and directly with coordinating the many bodies involved
paid for by the discharger. Hence, it becomes with an estuary as well as the need to conserve
economically viable to minimize waste dis- and preserve the environment. They may use
charged into the nearby surface waters. structure plans to foster environmentally
The UK environment protection agencies sustainable development of natural and man-
(Environment Agency in England and Wales, made resources and economic regeneration,
Scottish Environmental Protection Agency but to take into account sea-level rise, and to
(SEPA) and EHS in Northern Ireland) are minimize adverse effects on the environment.
responsible for issuing consents for individual By their nature, the structure plans have to
land-based discharges to air, land, and water. consider water quality, coastal flooding,
Nature conservation is the statutory remit of undeveloped coasts, natural and cultural her-
the country agenciesEnglish Nature, Scottish itage, special habitats, and hazardous industry.
Natural Heritage, Countryside Council for As an example, coastal and estuarine
Wales and Environment and Heritage Service regional municipalities and local authorities in
in Northern Ireland, who are coordinated the United Kingdom have a role to consider
through the Joint Nature Conservation and refine Structure Plan policies for their
Committee (JNCC). These bodies declare estu- coastlines. For example, within the Humber
arine sites as nature reserves, or sites of special estuary, Eastern England, the municipalities
scientific interest (SSSIs), and manage the have adopted policies in line with managing a
European designationsSPA and SAC. low lying area subject to increasing sea-level
In the Netherlands, a strong government rise and isostatic rebound (i.e. land sinking
policy on estuarine and coastal zone manage- since the last Ice Age). There is a presumption
ment is in operation, implemented by the against new development in areas liable to
Rijkswaterstaat, especially the Tidal Waters flooding or in areas which may require an
Division. Similarly, Australia and New Zealand enhanced level of protection, and they have to
have collaborated to produce water quality ensure that the development of land bordering
186 T H E E S T UA R I N E E C O S YS T E M
the estuary does not result in the loss of different namesfor example, defining
irreplaceable environmental resources such as Reference Conditions within the EU Water
mudflats. The body should ensure that the Framework Directive, Favourable Conservation
demand for coastal estuary protection does Status within the EU Habitats Directive,
not lead to the need for flood defences, which Ecological Quality Objectives (EcoQOs EcoQS)
would have a detrimental effect on estuarial as defined by OSPAR, and Reference sites under
environments. This is against a background of the Australian NZ Water Quality Guidelines.
a governmental strategy for flood defence and An estuary or its parts are often classified by
coastal erosion, that discourages building in managers in order to (a) describe the overall
areas susceptible to flooding erosion, and an quality of the system, (b) indicate which areas
Environment Agency strategy to protect exist- need action to be improved, restored, or reha-
ing facilities with coastal defences but not to bilitated, or (c) indicate the effectiveness of
impede the natural hydrography. any actions taken. While there are many such
classification schemes in operation world-
NonGovernmental Organizations wide, the example adopted by the UK envi-
In most countries, nongovernmental organiza- ronment protection agencies (Box 8.7) gives
tions (NGOs) play a role in public awareness many of the main features. Within the scheme,
and are often a non-statutory consultee in points are awarded for each biological, aes-
developments. For example in the United thetic, and water quality feature within each
Kingdom, there are many strong conservation- part of the estuary and then the points are
based NGOs such as the Royal Society for the totalled for each part. The points are then
Protection of Birds (RSPB), the Wildlife Trusts, related to a Class indicating good quality
the Worldwide Fund for Nature (WWF), the (Class A), fair quality (Class B), poor quality
Campaign for the Protection of Rural England (Class C) to bad quality (Class D).
(CPRE), Greenpeace and Friends of the Earth. Following the classification of an area,
These bodies may promote nature conservation where the area within each class can be
and natural resources, monitor and comment summarized for an estuary, targets can then
on local and structure plans, provide advice on be set for future improvements or remedia-
conservation issues, purchase or lease land for tion. Similarly, if the scheme is used at
nature reserves, monitor conservation value repeated intervals, say 5 years, then it should
and promote the need for further research and indicate the effectiveness or even costbenefit
cooperative effort. analysis of protection measures employed. For
These bodies in general have an underlying example, in the Forth estuary, Scotland, an
theme of working toward sustainable devel- area around the Grangemouth oil refinery was
opment, that is, the use of natural resources in originally designated as Class C because of the
order to meet existing demands but not to changes to the benthos resulting from the
jeopardize future uses. refinery effluent discharged to the intertidal
area. The building of a wastewater treatment
Estuarine classification schemes and plant and extension to the effluent pipe to
standards/objectives: Deviation from normality increase the dilution of the waste then resulted
A central theme in the management of estuar- in the quality of the area being improved to
ies is the use of standards and objectives, Class B such that the refinery company and
which indicate what is the normal situation the local EPA were able to quantify in area the
and to measure departure from that normality benefits of spending money on the new plant.
and then to bring in management options to As a wider form of classification systems
restore normality. This occurs throughout and geographical assessments, there is an
many policies and instruments and may have increasing use of Quality Status Reports (QSR)
Box 8.7 Estuarine Classification Scheme (Produced by the Environment and Heritage Service, Northern Ireland)
Biological Quality: points awarded for the passage of migratory fish, the support of a usual and expected fish community, the presence of a typical benthic
community, the ability to meet microbiological standards, and the absence of toxic/tainting substances in biota;
Aesthetic Quality: points based on the easily observed signs of estuarine polluting inputs sufficient to interfere with estuarine usage or to cause aesthetic pollution;
includes the presence on beaches and in the waters of litter, sewage solids, foaming, slicks, etc;
Water quality: points based on the ability to meet agreed (nationally or at EU level) chemical standards such as ammonia or dissolved oxygen concentrations exceeding
specified saturation values, levels of persistent pollutants.
Summation of points in each category to produce the Estuary Classification Scheme:
Class Description Aesthetic Fish Resident Resident Persistent Dissolved UK Red list
condition migration biota fish substances oxygen and EC
and/or (Biota) (DO) in dangerous substances
bioassay water in water
A given area of estuary is classified by allocating it to the highest class to which all of its criteria conform. An estuarine are satisfying Class B aesthetic and chemical
criteria but which is Class C on the basis of one of the biological criteria would be classified as Class C overall (i.e. a default system). If there are only limited biological
and chemical data then estuaries will be classified using information for known discharges, pollution complaints, etc. If there are none of the latter then the area will be
Class A.
188 T H E E S T UA R I N E E C O S YS T E M
such as those produced by the OSPAR for the reclamation and other engineering works
North Sea and the NE Atlantic area. These may, in the long run, prove to be the most
QSR summarize the nature of these sea areas, intractable in managing our estuarine ecosys-
their natural and anthropogenic features, and tems. Management has to include an assess-
the threats to their resources. ment of the impact of loss of area, not only as
a food loss for birds or fish, but also for the
loss of a free sewage-works area, which can
8.5 Final remarks
cope with the various pollutants discussed
Although the discussion here has indicated above. It is essential to ask appropriate ques-
methods for dealing with pollution, and espe- tion and to understand the functioning of all
cially that from point sources such as dump- aspects of the area before undertaking any
ing from vessels or from pipelines, the more irreversible activities (Fig. 8.5).
intractable problems requiring management The conservation interest in estuaries is
of estuaries are diffuse pollution, such as from focused on two aspects. First the maintenance
farmland, and the loss of habitats. The prob- of fisheries, either by the harvesting of fish
lems of the destruction of estuarine habitat by and shellfish, or by the use of estuaries as
nursery areas for marine fish stocks, or for fish interest. Persistent chemicals, which accumu-
such as sprat and flounder caught within the late in the food chain must, however, be
estuary, or for estuaries as routes for migra- strictly controlled. Excessive reclamation,
tory species to pass through. The conservation especially if it takes away intertidal areas,
of fish populations is inhibited by poor water which are often more productive than subtidal
quality (usually low dissolved oxygen, but areas, poses probably the greatest single threat
also, e.g., high ammonia levels), which acts as to estuarine conservation, destroying forever
a barrier to migratory fish, as well as levels of the habitat.
persistent pollutants, and high nutrient levels, The management of estuaries is thus largely
which after the collapse of a phytoplankton the protection of biological systems, even if the
bloom can lead to low dissolved oxygen lev- biology is that of mankind. The future man-
els. Second, the conservation interest is cen- agement of estuaries as a resource for wildlife
tered on the abundant bird populations, as well as for mankind depends very much on
which feed on the intertidal areas, attracted by the successful implementation of legislation
the rich and available food supplies. To pro- for pollution control and habitat protection.
tect these interests, many estuaries have been As emphasized throughout the chapters here,
declared as nature reserves, primarily the options for sustainable and cost-effective
because, as stated in the original paradoxes, management have to fulfil six tenets: being
they can be some of the least disturbed natural environmentally sustainable, economically
areas with rich natural animal populations, viable, technologically feasible, legally permis-
and also to help protect them against pollution sible, socially desirable, and administratively
or reclamation, which can destroy them for- achievable. Because of this, an estuarine
ever. The conservation of wildlife in estuaries scientist working on real environmental prob-
need not exclude other users. Industries are lems cannot ignore all the aspects mentioned
compatible with wildlife, provided that here; similarly, managers, legislators, and
organic effluents and power-station dis- policy-makers need an understanding of
charges are well diluted and carefully sited. the science of estuaries in order to interpret
Indeed small amounts of organic effluents, and implement the available tools. Only by
and well-controlled power-station discharges taking such a wide view can our estuaries be
may actually serve to increase the productiv- managed properly for the benefit of the
ity of an estuary, and enhance its conservation wildlife and Man.
Reading list
The reading list for this book consists of first a Bergman, H. L., Kimerle, R. A. and Maki, A.W. (eds.)
selection of general reference books, multi-author (1986). Environmental Hazard Assessments of Effluents.
works and journals which have applicability to Pergamon Press, Oxford and New York, 366 pp.
all of the book, or to several chapters, followed Bird, E. C. F. (1993). Submerging coasts The Effects of
by references to individual papers which are a Rising Sea Level on Coastal Environments. John
arranged in chapter groupings. If a reference Wiley and Sons, Chichester, 194 pp.
relates to more than one chapter it is listed in the Bird, E. C. F. (1996). Beach Management. John Wiley
first chapter in which appears. and Sons, Chichester, 281 pp.
Bird, E. C. F. (2000). Coastal Geomorphology: An
Introduction. John Wiley and Sons, Chichester,
322 pp.
Books and Multi-author works
Birnie, P. W. and Boyle A. E. (1992). International
Allanson, B. R. and Baird, D. (eds.) (1999). Estuaries Law and the Environment, Oxford University
of South Africa. Cambridge University Press, Press, Oxford, 563 pp.
Cambridge, 340 pp. Carter, R. W. G. (1988). Coastal Environments
Aller, J. Y., Woodin, S. A., and Aller, R. C. (eds.) An Introduction to Physical, Ecological and
(2001). Organism-sediment Interactions. University Cultural Systems of Coastlines. Academic Press,
of South Carolina Press, Columbia, S.C. Belle W London, 640 pp.
Baruch Library in marine science, Number 21, Chapman, D. (ed.) (1996). Water Quality Assessment:
403 pp. A Guide to the Use of Biota, Sediments and Water
Attrill, M. J. (ed.) (1998). A Rehabilitated Estuarine in Environmental Monitoring. (2nd edition) E. and
Ecosystem. Kluwer, Dordrecht. 254 pp. F. N. Spon, London, 626 pp.
Baretta-Bekker, J. G., Duursma, E. K. and Chapman, V. J. (ed.) (1977). Wet Coastal Ecosystems.
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Berlin, 357 pp. the Sea. (3rd edition) Manchester University Press,
Barnes, M. and Gibson, R. N. (eds.) (1990). Trophic Manchester, 494 pp.
Relationships in the Marine Environment. Aberdeen Cicin-Sain, B. and Knecht, R. W. (1998). Integrated
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NW Europe. Cambridge University Press, Clark, R. B. et al. (2001). Marine Pollution (fifth edi-
Cambridge, 287 pp. tion). Oxford University Press, Oxford, 237 pp.
Barnes, R. S. K. and Hughes, R. N. (1999). An Constable, A. J. and Fairweather, P. G. (eds.) (1999).
Introduction to Marine Ecology, (third edition) Ecology of estuaries and soft-sediment habitats.
Blackwells, Oxford, 286 pp. Australian Journal of Ecology, 24, 289476.
Bayne, B. L., Clarke, K. R. and Gray, J. S. (eds.) Davidson, N. C. et al. (1991). Nature Conservation and
(1988). Biological Effects of Pollutants. Marine Estuaries in Great Britain. Nature Conservancy
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Law (fifth edition of Ball and Bell). Blackstone Press, lar reference to southern Africa. Balkema books,
London, 726 pp. Rotterdam, 400 pp.
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READING LIST 191
Day, J. W. et al. (1989). Estuarine Ecology. John Wiley, Jones, N. V. and Wolff, W. J. (eds.) (1981). Feeding
New York, 558 pp. and Survival Strategies in Estuarine Organisms.
De Leeuw, C C. and Backx, J. J. G. M. (2001). Naar Plenum Press, New York. 304 pp.
een hertsel van estuariene gradienten in Nederland. Kay, R. and Alder, J. (1999). Coastal Planning and
Rijkswaterstaat rapport nr. 2000.044, 167 pp. Management. E. and F. N. Spon (Routledge),
Department of the Environment (1993). Coastal London, 375 pp.
Planning and Management: A Review. HMSO Kennedy, V. S. (ed.) (1980). Estuarine Perspectives.
Publications, London, 178 pp. Academic Press, New York, 533 pp.
Dickson, K. R., Maki, A. W. and Brungs, W. A. (eds.) Kennedy, V. S. (ed.) (1982). Estuarine Comparisons.
(1987). Fate and Effects of Sediment-bound Chemicals Academic Press, New York, 709 pp.
in Aquatic Systems. Pergamon Press, New York, Kennedy, V. S. (ed.) (1984). The Estuary as a Filter.
449 pp. Academic Press, New York.
Dyer, K. R. (1986). Coastal and Estuarine Sediment Kennish, J. R. (1986). Ecology of Estuaries (Volume 1,
Dynamics. Wiley, Chichester, 342 pp. Physical and Chemical aspects). CRC Press, Florida,
Dyer, K. R. (1997). Estuaries A Physical Introduction 254 pp.
(2nd Edition). John Wiley, Chichester, 199 pp. Kennish, M. J. (ed.) (2000). Estuary Restoration and
Dyer, K. R and Orth, R. J. (eds.) (1994). Changes Maintenance. CRC Press, Florida.
in fluxes in estuaries: implications from science to Ketchum, B. H. (ed.) (1983). Estuaries and Enclosed
management. Proceedings of the ECSA22/ERF Seas. Elsevier, Amsterdam, 500 pp.
Symposium. Olsen and Olsen, Fredensborg, Kinne, O. (ed.) (1978, onwards). Marine Ecology
Denmark. 485 pp. (5 volumes). John Wiley, New York.
Dyke, P. (1996). Modelling Marine Processes. Prentice- Kjerfve, B. (ed.) (1978). Estuarine Transport Processes.
Hall, London, 152 pp. Belle W. Baruch Library in Marine Science, No. 7,
Elliott, M. and Ducrotoy, J-P. (eds.) (1991). Estuaries University of South Carolina Press, Columbia,
and Coasts: Spatial and Temporal Intercomparisons. S.C., 331 pp.
Olsen and Olsen, Fredensborg, Denmark. 390 pp. Knox, G. A. (1986). Estuarine Ecosystems (2 volumes).
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Environmental Perspectives for the Northern Lalli, C. M. and Parsons, T. R. (1997). Biological
Seas. Marine Pollution Bulletin, 29, 253660. Oceanography: An Introduction. (second edition)
French, P. (1997). Coastal and Estuarine Management. Butterworth Heinemann, Oxford, 314 pp.
Routledge, London, 251 pp. Lauff, G. H. (ed.) (1967). Estuaries. American
Goksyr, A. (ed.) (1998). Pollution Responses Association for the advancement of science,
in Marine Organisms (PRIMO9). Marine Washington, D.C. Publication number 83, 755 pp.
Environmental Research, Special Issue, 46, 1607. Lawrence, A. J. and Hemingway, K. L. (eds.) (2003).
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192 READING LIST
Forth. Proceedings of the Royal Society of Edinburgh, Postma, H. and Zijlstra, J. J. (eds.) (1988).
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McLusky, D. S., de Jonge, V. N. and Pomfret, J. Raffaelli, D. and Hawkins, S. (1996). Intertidal
(eds.) (1990). North Sea Estuaries Interactions. Ecology. Chapman and Hall, London, 356 pp.
Hydrobiologia, 195, 1221. Raymont, J. E. G. (1980). Plankton and Productivity
Meire, P and Vincx, M. (eds.) (1993). Marine and in the Oceans (Volume 1: Phytoplankton). (second
estuarine gradients. Netherlands Journal of Aquatic edition). Pergamon Press, Oxford, 489 pp.
Ecology, 27, 71496. Raymont, J. E. G. (1983). Plankton and Productivity in
Moreira, M. H., Quintino, V. M. S. and Rodrigues, the Oceans (Volume 2: Zooplankton). (second edition).
A. M. J. (eds.) (1995). Northern and southern Pergamon Press, Oxford, 824 pp.
estuaries and coastal areas. Netherlands Journal of Reise, K. (1983). Tidal Flat Ecology An Experimental
Aquatic Ecology, 29, 199471. Approach. Springer-Verlag, Heidelberg, 191 pp.
Moulder, D. S. and Williamson, P. (eds.) (1986). Reise, K. (2001). Ecological Comparisons
Estuarine and Coastal Pollution: Detection, of Sedimentary shores. Springer, Heidelberg,
Research and Control. Water Science and 387 pp.
Technology, 18, 1364. Rijkswaterstaat (1985). Biological Research in the
National Research Council (2000). Clean Coastal Ems-dollart Estuary. Rijkswaterstaat, The Hague,
Waters Understanding and Reducing the Effects of Netherlands, 182 pp.
Nutrient Pollution. National Academy Press, Ros, J. D. (ed.) (1989). Topics in Marine Biology
Washington, D.C., 405 pp. (Proceedings of 23rd EMBS). Scientia Marina, 53
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Estuaries. Advances in Ecological Research, 29. Salomons, W., Bayne, B. I. Duursma, E. K.
1295, Academic Press, London. and Forstner, U. (eds.) (1988). Pollution of the North
Neff, J. M. (2002). Bioaccumulation in Marine Sea An Assessment. Springer-Verlag, Berlin, 687 pp.
Organisms Effect of Contaminants from Oilwell Schmitt, R. J. and Osenberg, C. W. (1996). Detecting
Produced Water. Elsevier, Amsterdam, 452 pp. Ecological Impacts: Concepts and Applications in
Newman, P. J. and Agg, A. R. (eds.) (1988). Coastal Habitats. Academic Press, New York,
Environmental Protection of the North Sea. 401 pp.
Heinemann, Oxford, 886 pp. Schramm, W. and Nienhuis, P. H. (eds.) (1996).
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Oosterschelde Estuary (The Netherlands): A Case Effects of Eutrophication. Springer-Verlag, Berlin,
Study of a Changing Ecosystem. Kluwer Academic 470 pp.
Publishers, Dordrecht, 597 pp. Sheppard, C. (ed.) (2000). Seas at the Millennium: An
Open University Course Team (1999). Waves, Tides Environmental Evaluation. 3 volumes, Pergamon
and Shallow-water Processes. (second edition). Open Press, Oxford, 2400 pp.
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227 pp. (2001). Encyclopaedia of Ocean Sciences, 6 Volumes.
Orive, E., Elliott, M. and de Jonge, V. N. (eds.) Academic Press, London, 3399 pp.
(2002). Nutrients and Eutrophication in Estuaries Tait, R.V. and Dipper, F. A. (1998). Elements of Marine
and Coastal Waters. Kluwer Academic Publishers, Ecology (4th edition). Butterworth-Heinemann,
Dordrecht, 526 pp. Oxford, 462 pp.
Penning-Rowsell, E. et al. (1992). The Economics of Tenore, K. R. and Coull, B. C. (eds.) (1980). Marine
Coastal Management, Belhaven Press, London, Benthic Dynamics. Belle W. Baruch Library in
380 pp. Marine Science, No. 11, University of South
Perkins, E. J. (1974), The Biology of Estuarine and Carolina Press, Columbia, S.C., 451 pp.
Coastal Waters. Academic Press, London, 678 pp. Viles, H. and Spencer, T. (1995). Coastal Problems:
Pethick, J. S. (1984). An introduction to coastal Geomorphology, Ecology and Society at the Coast.
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Phillips, D. J. H. and Rainbow, P. S. (1994). Walker, C. H., Hopkin, S. P., Sibly R. M. and Peakall
Biomonitoring of Trace Aquatic Contaminants. D. B. (1996). Principles of Ecotoxicology. Taylor and
Chapman and Hall, London, 371 pp. Francis, London, 321 pp.
READING LIST 193
Whitehouse R., Soulby, R., Roberts, R. and would like details of these associations is invited
Mitchener, H. (2000). Dynamics of Estuarine Muds. to contact the authors who will forward the
Thames Telford Ltd., London, 210 pp. current address of the ECSA or ERF Secretary.
Wiley, M. (ed.) (1976). Estuarine Processes 2 volumes. These organizations produce or sponsor the key
Academic Press, London, 603 pp. journals in this subject area: Estuarine Coastal and
Wilson, J. G. and Halcrow, W. (eds.) (1985). Shelf Science and Estuaries.
Estuarine Management and Quality Assessment. In addition to the above journals, the following
Plenum Press, New York, 225 pp. journals contain a selection of relevant papers.
Wilson, J. G. (1983). The Biology of Estuarine Aquatic Ecology
Management. Croom Helm, London. 204 pp. Aquatic Science and Biology Abstracts
Wolfe, D. A. (ed.) (1986). Estuarine Variability. Coastal Zone Topics
Academic Press, Florida, 509 pp. Journal of Animal Ecology
Wolff, W. J. (ed.) (1980). Ecology of the Wadden Sea, Journal of Applied Ecology
3 volumes. Balkema Books, Rotterdam, 1300 pp. Journal of Experimental Marine Biology and Ecology
Journal of the Marine Biological Association of the U.K.
Handbooks on methods for Limnology and Oceanography
studying estuaries Marine Biology
Marine Ecology Progress Series
Bagenal, T. B. (1978). Methods for Assessment of Fish Marine Pollution Bulletin
Production in Fresh Waters. Blackwell Scientific Journal of Sea Research
Publications, Oxford. 3rd edition, 365 pp. Oceanography and Marine Biology. An Annual Review
Baker, J. M. and Wolff, W. J. (eds.) (1987). Biological Ophelia
Surveys of Estuaries and Coasts. EBSA handbook,
Cambridge University Press, Cambridge, 449 pp.
Dyer, K. R. (ed.) (1979). Estuarine Hydrography and
Chapter One
Sedimentation. EBSA handbook, Cambridge Anderson, F. E. (1983). The northern muddy inter-
University Press, Cambridge, 230 pp. tidal: a seasonally changing source of suspended
Head, P. C. (ed.) (1985). Practical Estuarine Chemistry. sediments to estuarine waters-a review. Canadian
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Index
Cerastoderma edule ($ Cardium edule) (Cockle) 26, 54, 61, Dredging 125129, 171
73, 86, 100, 116 Duplin River, Georgia 36
Charleston Harbor, USA 5
Chemical factors 12 Economic value 91
Chesapeake bay, USA 5, 12, 44, 46, 48, 64, 65, 66, 93 Ecosystem pathology 94
Chlorophyll 4546 Ecotone 102
Chromium 118, 119, 167, 168 Elbe, Germany 5, 14, 104
Chronic pollution 9597, 140 Embayments 4, 5
Circulation 5, 6, 7, 8, 9 Ems-Dollard, Netherlands/Germany 11, 14, 4345, 50,
Clyde estuary, Scotland 7, 64, 93, 97, 98 71, 99, 100, 130
Coastal lagoons 2, 7, 15 Energy budget 28
Coastal plain estuaries 4, 5 Energy subsidy 50
Coastal squeeze 124 Enteromorpha (filamentous algae) 34, 41, 99, 141, 143
Coastal Zone Management 182 Environmental Impact assessment (EIA) 153, 170
Columbia River, USA 46 Environmental management Systems 160
Community level assessments 147148 Environmental quality 137
Compensation for land-claim 123 Environmental quality objectives (EQO) 163170
Complex estuaries 4, 5 Environmental Quality standard (EQS) 163170
Compliance monitoring 139 Environmental Stress 137, 138
Conservation 136 Episodic pollution 96
Conservative wastes/ inputs 96 Essex, England 5
Contamination 96 Estuarine Classification Schemes 186188
Cooling water 130 Estuarine resident fish 75
Copper 118, 119, 167, 168 European Union (EU) 176180
Coriolis 6, 7 European-type estuaries 5, 52, 70
Cornwall, England 5 Euryhaline 19, 21, 25, 81
Corophium volutator (mud shrimp) 25, 26, 54, 59, 83, Euryoecious 25
86, 140 Eurytemora (Zooplankter) 64, 67
Crab larvae 67 Eutrophication 45, 57, 99, 100
Crangon crangon (shrimp) 25, 73, 78, 79, 83, Export 35
89, 175
Crassostrea (oyster) 60 Factor complexes 15
Crustacea 5960, 118 Fairbridges definition 2
Ctenophores 64, 68, 83 False Bay, Washington 43
Current speed 9 Feeding guilds 55
Cytochrome p450 141 Fish 7482, 106, 144
Fisheries 105107, 170
Decision tree 155, 156 Fisheries effect of. 105108
Decomposers 34 Fjard 4, 5
Dee estuary, UK 5 Fjord 46
Definition of an estuary 2, 15 Flax Pond, New York State 36, 49
Degradable wastes/ inputs 96 Florida 5, 81
Delaware, USA 5, 7, 44, 45, 77, 172175 Flushing Time 8, 44, 65
Delta Plan 123124, 131 Food web 14, 27, 32
Denmark 3, 7, 122 Forth estuary, Scotland 5, 8, 51, 61, 62, 66, 69, 78, 79, 86,
Deposit feeding 53, 55, 72 87, 101, 106, 116, 122
Detoxification 141 Fourleague bay, USA 45
Detritus 34, 38, 4653, 70 Freshwater fish 75
Diadromous fish 75, 129 Freshwater-brackish interface (FBI) 13
Diffuse pollution 95, 96, 98, 100 Freshwater-seawater interface (FSI) 13, 15, 26, 66
Direct input 96 Fucus spp. (seaweeds) 34, 40, 41, 141, 143, 175
Dissolved Organic matter 12, 38, 47 Fundulus heteroclitus (killifish) 74, 76, 77, 89
Diversity 20
Diversity index 150 Gammarus spp. (sand-hopper) 25
Divisions of an estuary 8 Genetic techniques 144146
DPSIR approach 93, 94, 160 Georgia 36, 37, 55, 64
INDEX 211
Nanaimo, Canada 49 Patuxent River, USA 12, 38, 65, 66, 67, 74
Narragansett Bay, USA 64, 83 PCBs 119121, 168
National Pollutant Discharge Elimination Scheme Pearl River 44
(NPDES) 163, 164 PearsonRosenberg model 97, 103, 151
Negative estuary 2, 3 Pesticides 113
Nematodes 62, 63, 97 Petpeswick Inlet, Canada 56
Neomysis integer (Mysid shrimp) 55, 6467, 83 Petrochemical discharges 113117
Nephrops norvegicus (Norway lobster) 99 Petromyzon marinus (Sea lamprey) 162
Nephtys spp. (Cat-worm) 25, 54, 58, 73, 77, 83 pH 13
Netherlands 122, 124 Phosphates 14, 100
Neutral estuary 2, 3, 4 Photosynthetic rate 35, 43
New England 5 Physical features 4
New York 93 Physiological mechanisms 22
New York Bight 97 Physiological stress 2226
New Zealand 5, 169, 170 Phytoplankton 14, 34, 4346, 53, 72, 99, 100
Nickel 167, 168 Planning and designations 160172
Nigeria 116 Plant production 3452
Nitrogen 14, 47, 98 Platichthys flesus (flounder) 19, 73, 74, 77, 78, 89, 90,
Non-indigenous organisms 94 143, 175
North Carolina 5, 43, 49, 99 Platichthys americanus (winter flounder) 78
North Inlet, SC 49, 60 Pleuronectes platessa (plaice) 76
North Kent estuaries, England 5 Pluvialis squatarola (Grey plover) 85, 87
North Sea 14, 65, 76, 78, 121, 123, 131 Pollution 26, 9398
Norway 5 Pollution, definitions of 96
Numenius arquata (Curlew) 8587 Population responses 146147
Numerical analysis of data 149152 Posidonia (sea grass) 3941
Nutrients 14, 34, 38, 45, 47, 72, 94, 98100 Positive estuary 2, 3
Potamopyrgus jenkinsi (mud snail) 25
Oil 48, 95, 113117, 167, 168 Potassium-40 120
Oil extraction 116 Potomac River, USA 8, 21
Oil refinery 116 Power generation 128131
Oil spillage 114117 Practical salinity Scale 2, 3
Oligochaetes 55, 59, 101, 104, 116 Praunus flexuosus (Mysid) 25, 34, 67, 83
Oligohaline 19, 20 Predator effects 91
Omnivores 55 Primary consumers 32, 5372
Oosterschelde 4346, 72, 76, 77, 123, 124, 137 Primary producers 32, 3452
Opportunistic species 102 Pritchards definition 2
Organic enrichment 99103 Productivity 1, 28, 29, 32, 55, 71, 91, 146
Organic enrichment 103 Protozoan 64
Organic matter 94 Pygospio spp. 89
Organisms in estuaries 19 Pyramid of biomass 27, 28
Organohalogens 119, 120, 167, 168 Pyramid of numbers 27, 28
Osaka 92
Osmoregulation 24, 25 Radioactive discharges 94, 120121
Osmotic factors 24 Rance estuary, France 132, 135
Osmotic stress 45, 66 Reclamation See Land-Claim
Outwelling hypothesis 35 Recreation 136
Oxygen 13 Recycling of nutrients 14, 34
Oxygen depletion or sag 67, 99, 101, 127, 168, 173 Red list 168
Red Sea 3
P:B ratio 2831, 59, 147 Redox potential (Eh) 12, 101, 102
Pandalus montagui (shrimp) 19 Refractory matter 46
Partially mixed circulation 5, 6 Remanes curve 21, 26
Particulate Organic Matter (POM) 12, 34, 48, 52, 53, 70 Respiration 31, 32
Pathology of ecosystem 94 Responses to pollution 95
INDEX 213
Retusa obtusa (Gastropod) 77, 84 Spartina spp. 8, 3639, 46, 47, 52, 61, 70, 77, 86
Rhine 14 Special Protected Areas (SPAs) 163
Ria 4, 5 Speciation 25
Ria de Arosa, Spain 43 Species number 21, 22, 102, 149
Ria de Aveiro, Portugal 135 Spencer Gulf, Australia 3
Ribble estuary, England 7 Sprattus sprattus (sprat) 75
River discharge 35 Stability-time hypothesis 22
Rotterdam 126, 171 Stenohaline 1921, 25
Rubidium-87 120 Stokes law 10
Stress 96, 157
S-A-B curves 102, 151 Suaeda maritima (grass) 38
Salicornia (samphire) 38 Substrate 21
Salinity 2, 3, 8, 15, 19, 21, 23, 80 Surveillance (Condition monitoring) 139
Salinity gradient 15 Suspension feeding 53, 55, 72
Salmo salar (Salmon) 19, 74, 129, 162 Swartkops estuary, South Africa 69
Salmon farming 108110 Sweden 7, 78
Salt extraction 135 Synthetic substances (e.g. Plastic) 167
Salt flocculation 10
Salt marsh 34, 3639, 48, 70 Tadorna tadorna (Shelduck) 84, 87, 88
Salt wedge 5, 6 Tagus, Portugal 43
Sampling design 149 Tamar estuary, England 14
San Francisco Bay 5, 44, 48, 122 Tampa Bay, Florida 7
Sapelo Island, Georgia 49, 64 Tay estuary, Scotland 5, 7, 9, 20
Scheldt estuary, The Netherlands See Oosterschelde Tees estuary, England 7, 8587, 93, 99, 100, 122, 132
and Westerschelde Tellina tenuis (bivalve) 76
Scrobicularia plana (bivalve) 28, 54, 86 Texas 3
Scyphozoans 68 Thallasia (sea grass) 3941
Sea-level rise 124 Thames estuary, England 5, 7, 93, 98, 104, 124,
Secondary consumers 32, 7391 131, 175
Sediment testing 141 Thames barrier 132
Sediment triad approach 152 Tidal flat turntable 90, 91
Sedimentation 119 Tidal-surge barrier 124
Sediments 1, 912, 15, 48, 126 Tides 1, 35
Seine, France 7 Tokyo 92
Selective deposit feeders 55 Tolerance of pollution 95
Settling velocity 9 Tomales bay, USA 45
Severn estuary, UK 5, 7, 132 Toxicity tests 121, 140, 165
Sewage 35, 48, 57, 9597, 100, 173 Trace metals 94
Sewage sludge disposal 97, 98 Transition zone 1, 102
Shanghai 92 Transitional waters 16
Shannon, Ireland 46 Tringa totanus (Redshank) 39, 84, 86, 87
Sheepscot estuary, Maine 86 Trophic diagrams 71
Shellfish farming 111113, 169 Trophic relationships 27, 28, 33
Shellfish poisoning 99 Tropical fish 7982
Shoreline Management Plans (SMPs) 172 True estuarine species 19
Sieve size 55 Tubifex spp. (Worm) 55
Sippewisset, MA 36 Tubificoides spp. (Worm) 90, 97
Slocum river estuary, MA 76 Turbidity 11, 43, 46, 52, 65, 126
Solea solea (Dover Sole) 176 Turbidity maximum 11
Solway Firth, UK 5 Tyne estuary, England 7
Somateria m. mollissima (Eider duck) 84, 88 Types of estuary 4
Somme, France 7, 56, 69
South Africa 4, 5, 80 Uniform Emission Standard (UES) 163170
South Carolina 56 Users of estuaries 92136
Southampton water, England 122, 123, 129 Uses of estuaries 92136
214 INDEX