Newell 1998

See discussions, stats, and author profiles for this publication at: https://www.researchgate.
net/publication/298415859
The impact of dredging works in coastal waters: a review of

the sensitivity to disturbance and subsequent recovery of
biological resources on the sea bed
Article in Oceanography and Marine Biology · January 1998
CITATIONS READS
394 3,924
3 authors, including:
Lindsay Jane Seiderer

Consonamus Limited
32 PUBLICATIONS 1,660 CITATIONS
SEE PROFILE
Some of the authors of this publication are also working on these related projects:
Benthic disturbance & recovery following marine aggregate extraction View project
Multi-disciplinary upwelling/kelp bed ecosystem project. University of Cape Town. 1980s. View project
All content following this page was uploaded by Lindsay Jane Seiderer on 04 July 2017.
The user has requested enhancement of the downloaded file.

Oceanography and Marine Biology: an An nual Review 1998,36,127-78
0 A. D. Ansell, R. N. Gibson and Margaret Barnes, Editors
UCL Press
THE IMPACT OF DREDGING WORKS IN

COASTAL WATERS: A REVIEW OF THE
SENSITIVITY TO DISTURBANCE AND
SUBSEQUENT RECOVERY OF BIOLOGICAL
RESOURCES ON THE SEA BED
R. C. NEWELL,1 L. J. SEIDERER1 & D. R. HITCHCOCK2

1
Marine Ecological Surveys Limited, West Country Office, Trewood Cottage,
Steeple Lane, St Ives, Cornwall TR26 2PF, UK
2
Coastline Surveys Limited, Bridgend Farmhouse, Stonehouse,
Gloucestershire GL1O 2AX, UK
Abstract The present review provides a framework within which the impact of dredging on biological
resources that live on the sea bed (“Benthic” communities) can be understood, and places in perspective some of
the recent studies that have been carried out in relation to aggregates dredging in European coastal waters. The
impact of dredging works on fisheries and fish themselves, and on their spawning grounds is outside the scope of
this review. We have, however, shown that empirical models for shelf waters such as the North Sea indicate that
as much as 30% of total fisheries yield to man is deriv ed from benthic resources, and that these become an
increasingly important component of the food web in near-shore waters where primary production by seaweeds
(macrophytes) and seagrasses living on the sea bed largely replaces that by the phytoplankton in the water
column. Because dredging works are mainly carried out in near-shore coastal deposits, and these are the ones
where benthic production processes are of importance in supporting demersal fish production, our review
concentrates on the nature of benthic communities, their sensitivity to disturbance by dredging and land
reclamation works, and on the recovery times that are likely to be required for the re-establishment of
community structure following cessation of dredging or spoils disposal.
Essentially, the impact of dredging activities mainly relates to the physical removal of substratum and
associated organisms from the seabed along the path of the dredge head, and partly on the impact of subsequent
deposition of material rejected by screening and overspill from the hopper. Because sediment disturbance by
wave action is limited to depths of less than 30m, it follows that pits and furrows from dredging activities are
likely to be persistent features of the sea bed except in shallow waters where sands are mobile. Recent studies
using Acoustic Doppler Current Profiling (ADCP) techniques suggest that the initial sedimentation of material
discharged during outwash from dredgers does not, as had been widely assumed, disperse according to the
Gaussian diffusion principles used in most simulation models, but behaves more like a density current where
particles are held together during the initial phase of the sedimentation process. As a result, the principal area
likely to be affected by sediment deposition is mainly confined to a zone of a few hundred metres from the
discharge chute.
Our review suggests that marine communities conform to well-established principles of ecological
succession, and that these allow some realistic predictions on the likely recovery of benthic communities
following cessation of dredging. In general, communities living in fine mobile deposits, such as occur in
estuaries, are characterized by large populations of a restricted variety of species that are well adapted to rapid
recolonization of deposits that are subject to frequent disturbance. Recolonization of dredged deposits is initially
by these “opportunistic” species and the community is subsequently supplemented by an increased species
variety of long-lived and slow-growing “equilibrium” species that characterize stable undisturbed deposits such
as coarse gravels and reefs.
Rates of recovery reported in the literature suggest that a recovery time of 6-8 months is charactristic
1
R. C. NEWELL, L. J. SEIDERER AND D. R. HITCHCOCK
of many estuarine muds where frequent disturbance of the deposits precludes the establishment of long-lived
components. In contrast, the community of sands and gravels may take 2-3 yr to establish, depending on the
proportion of sand and level of environmental disturbance by waves and currents, and may take even longer
where rare slow-growing components were present in the community prior to dredging. As the deposits get
coarser along a gradient of environmental stability, estimates of 5-10yr are probably realistic for development of
the complex biolo gical associations between the slow-growing components of equilibrium communities
characteristic of reef structures.
Most recent studies show, however, that biological community composition is not controlled by any
one, or a combination of, simple granulometric properties of the sediments such as particle size distribution. It is
considered more likely that biological community composition is controlled by an array of environmental
variables, many of them reflecting an interaction between particle mobility at the sediment-water interface and
complex associations of chemical and biological factors operating over long time periods. Such interactions are
not easily measured or analyzed, but the results suggest that the time course of recovery of an equilibrium
community characteristic of undisturbed deposits is controlled partly by the process of compaction and
stabilisation that occurs following deposition.
Biological community composition thus reflects changes in sediment composition, but is also in
equilibrium with seabed disturbance from tidal currents and wave action, both of which show spatial variations
and interactions with water depth. The processes associated with compaction and stability of seabed deposits
may, therefore, largely control the establishment of long-lived components of equilibrium communities and
account for the dominance of opportunistic species in the initial stages of colonization in unconsolidated deposits
of recently sedimented material after the cessation of dredging
Introduction
The importance of benthic communities in marine food webs leading to commercially

exploitable yields of fish has been widely recognized. Early models for the North Sea (see
Steele 1974) suggested that of net primary production by the phytoplankton,
approximately 80% was consumed by pelagic herbivores such as copepods and eup-
ausiids, and 20% fell to the seabed as a detrital input to the benthic community. At each
step of the food web, relatively large amounts (80-90%) of the material entering the
consumers is remineralized and returned to the water column to support further primary
production by the phytoplankton, leaving a small proportion incorporated into the biomass
of the consumer.
Because of the complexity of marine food webs, and the major dissipation of
energy at each step of the food chain, the empirical model proposed by Steele (1965,
1974) for the North Sea and shown in Fig. 1 indicates that out of 100gC m-2 yr-1 produced
at the sea surface as net primary production by the phytoplankton, only 0.3 g C m-2 yr-l
appears as yield to man through the pelagic food web, and approximately 0.13 g Cm-2 yr-1
from demersal fish. Despite the huge dissipation of materials that occurs at each step in
the food web, however, sufficient carbon evidently flows through the detrital food web,
even in plankton-based ecosystems such as the North Sea, for as much as 30% of total fish
production to be dependent on conversion through the community which lives on the sea
bed.
More recent analyses of the trophic structure and fluxes of carbon in shelf waters
of the North Sea by Joiris et al. (1982) suggest that as much as 50% of the annual phyto-
plankton production sinks to the sea bed as detritus and is supplemented by faecal
2
IMPACT OF DREDGING WORKS IN COASTAL WATERS
Figure 1 Simplified empirical carbon flow dia gram for the phytoplankton-based ecosystem of the
shelf waters of the North Sea. Note that, of the 100g C m-2 yr-1 of sea surface produced by the
phytoplankton, the yield to man through pelagic food webs and pelagic fish is estimated to be 0.3g
C m-2 yr-l where as that through benthic food webs is 0.13 g C m -2 yr-1, or about 30% of the total
exploitable fish yield to man. (Based on Steele 1965).
pellets of the zooplankton (see also Smetacek, 1984). The benthos is thus heavily
implicated in carbon flow in coastal systems, and becomes of increasing importance in
shallow waters where production by benthic algae (macrophytes) and seagrasses largely
replaces that derived from the phytoplankton (see also Taylor & Saloman 1968, Thayer et
al. 1975, Mann 1982, Moloney et al. 1986, Newell et al. 1988).
Benthic communities thus play a central role in the transfer of materials from
primary production by the phytoplankton, benthic macrophytes and coastal wetlands
through the detrital pool into higher levels in the food web, including commercially
exploitable fish. Most estimates suggest that even in phytoplankton-based systems such as
the North Sea, the yield to man through the benthos to demersal fish stocks is likely to
approach 30-40% of that derived through the pelagic system. Partly for this reason, the
populations of benthic communities which live on, and in, the deposits on the seabed have
been widely studied in investigations of the integrated effects of disturbance from a
variety of natural and other sources.
3
Early studies include extensive physiological-toxicological work on the potential

impact of suspended sediments on commercially significant target organisms (Loosanoff
1962, Sherk 1971, Sherk et al. 1972, 1974, Bright & Ellis 1989, Jokiel 1989, for review,
see Moore 1977). Such studies have been extended to include the potential impact of
dredging works on the ecology of biological communities in coastal embayments and
estuarine ecosystems (Jones & Ellis 1976, Morton 1977, Conner & Simon 1979, Johnston
1981, Ellis & Heim 1985, Ellis & Taylor 1988, Ellis & Hoover 1990, Giesen et al. 1990,
Onuf 1994).
Comprehensive studies of the impact of dredging for marine aggregates and sand
on marine communities in European waters have been carried out by Millner et al. (1977),
Pagliai et al. (1985), Sips & Waardenburg (1989), van Moorsel & Waardenburg (1990,
1991), and Kenny & Rees (1994, 1996). Reviews of the impact of sand and gravel
extraction include those of the International Council for the Exploration of the Sea (ICES,
1975, 1977, 1992a,b, 1993), Gayman (1978), de Groot (1986), Nunny & Chillingworth
(1986), Hurme & Pullen (1988), Lart (1991) and Charlier & Charlier (1992). A recent
review for the Minerals Management Service, US Department of the Interior containing a
number of specific case histories on the impact of marine mining has been given in a C-
CORE publication (1996; see also Ellis 1987).
Despite the work that has been carried out over the past 30 years, the non-
biologist could be forgiven for being bewildered by the diversity of the results and the
difficulties of making more than the most general predictions on the effects of dredging
activities including the extraction of marine aggregrates on biological resources.
Essentially, most studies show that dredging itself is usually accompanied by a significant
fall in species numbers, population density and biomass of benthic organisms. The rate of
recovery is, however, highly variable depending (among other factors) on the type of
community that inhabits the deposits in the dredged area and surrounding deposits, the
latitude and the extent to which the community is naturally adapted to high levels of
sediment disturbance and suspended particulate load.
In general, rapid rates of initial recolonization have been reported for some
coastal deposits where the organisms are mainly mobile “opportunistic” species that have
a rapid rate of reproduction and growth. Such organisms may also be able to recolonize
the deposits by migration of the adults (see McCall 1976, Conner & Simon 1979, Saloman
et al. 1982, Guillou & Hily 1983, Pagliai et al. 1985, van der Veer et al. 1985, Clarke &
Miller-Way 1992, Rees & Dare 1993, van Moorsel 1994). In contrast, long-lived and
slow-growing species, especially those in high latitudes may take several years before
larval recruitment and subsequent growth of the juveniles allows re storation of the
original community composition and biomass. The process of “recovery” following
environmental disturbance is generally defined as the establishment of a successional
community of species which progresses towards a community that is similar in species
composition, population density and biomass to that previously present, or at non-
impacted reference sites (C-CORE 1996; see also Ellis & Hoover 1990). Typically, values
range from up to one year in fine-grained deposits such as muds and clays (Ellis et al.
1995), although even in the fine deposits that characterize coastal ecosystems such as the
Dutch Waddensea, van der Veer et al. (1985) report that recolonization takes l-3 yr in
areas of strong currents but up to 5-10yr in areas of low current velocity. Longer recovery
times are reported for sands and gravels where an initial recovery phase in the first 12
months is followed by a period of several years before pre-extraction population structure
is attained (van Moorsel 1994, Kenny & Rees 1996).
4
Even longer times may be required for biologically-controlled communities that

characterize coarse deposits (see Garnett & Ellis 1995), although the evidence is
conflicting for coral reef communities. Some studies report long-term damage to coral
resources from sedimentation associated with dredging (Dodge & Vaisnys 1977, Bak
1978, Dodge & Brass 1984, Madany et al. 1987, Hodgson 1994; for review see Maragos
1991). Other studies suggest that corals themselves may be tolerant of short-term
increases in siltation associated with dredging (Marszalek 1981, Brown et al. 1990) but
that modification of community structure of other components of reef communities such
as fish species are detectable after multivariate analysis of species composition (Dawson-
Shepherd et al. 1992).
Recovery times following disturbance from a variety of sources, including
dredging work, may be extended in colder waters at high latitudes where communities
typically comprise large slow-growing species that may take many years for
recolonization and growth. In a Swedish fjord system, for example, a recovery that was
indistinguishable from natural variations was established only after 8 yr following closure
of a pulp mill (Rosenberg 1976), whereas de Groot (1979; see also Wright 1977, Aschan
1988) reports that recovery of communities within the Arctic Circle may take more than
12 yr compared with estimates of approximately 3yr for deposits off the coast of the
Netherlands. Similar extended timescales for recolonization by the benthic community
have been reported for Antarctic waters by Oliver & Slatterly (198 1).
The concept of “recovery” of biological resources is itself not an easy one to
define for complex communities whose composition can vary over time, even in areas that
remain undisturbed. Whether a community is identical in species composition and
population structure following cessation of dredging thus to some extent begs the question
of whether the biodiversity would have remained stable over that period in the absence of
disturbance by dredging. Probably a more practical approach to the question of “recovery”
will be the recognition of the establishment of a community that is capable of maintaining
itself and in which at least 80% of the species diversity and biomass has been restored.
This implies a substantial restoration of the carrying capacity of the benthic food
webs leading to fish, even though the precise species composition may not be identical to
that recorded in the pre-dredged system. This issue of whether biological resources have
been restored, and how this should be assessed, is of considerable importance in areas
such as Canadian coastal waters where recovery of seabed resources forms part of a
Statutory obligation following cessation of mining (D. V. Ellis pers. comm.).
Despite the complexity of the results for specific dredged areas, some firm
general principles governing community structure following environmental disturbance
have emerged in recent years and these appear to be generally applicable to a wide variety
of communities both on the land and on the sea bed. The application of such concepts to
coastal communities allows some credible predictions on the scale of impact of
environmental disturbance such as that imposed by dredging and dredged spoils disposal
and, more important, gives some insight into how long it might take for recovery in
dredged areas and the surrounding deposits once dredging has ceased.
The object of the present review is to provide a framework within which the bio-
logical impact and subsequent recovery of benthic resources can be understood, with
examples drawn mainly from the impact of dredging works in near-shore waters and
estuarine systems.
5
General features of community structure
Most general models of community structure are based on the concept that biological
communities do not form a series of distinct groups or assemblages along an
environmental gradient, but show a corresponding gradient in community composition.
Species that colonize habitats with unpredictable short-term variations in environmental
conditions at one end of an environmental gradient of stability are subject to frequent
catastrophic mortality. Such conditions occur in many shallow-water, intertidal and
estuarine habitats and are characterized by populations which tend to have a high genetic
variability that allows at least some components of the population to survive
environmental extremes (see Grassle & Grassle 1974, Guillou & Hily 1983). Such
organisms are thus selected for maximum rate of population increase, with high fecundity,
dense settlement, rapid growth and rather a short life cycle. They are well suited to rapid
invasion and colonization of environments where space has been left by a previous
catastrophic mortality, whether this has been induced by natural factors or disturbance by
man.
Such components have been designated “r-strategists” in a pioneer work by
MacArthur & Wilson (1967; see also Pianka 1970), although we prefer to use the term
“opportunists” for all such early colonizing species. Opportunists rely on a large
investment in reproductive effort, rather than on mobility, for success in colonizing
habitats made available by the catastrophic destruction of the previous community (see
Gadgil & Solbrig 1972, McCall 1976).
Many communities living in unstable environments may comprise small, highly
mobile species that are able to take advantage of recently created empty habitats quickly
and to colonize them with large populations. These mobile colonizers are often associated
with frequently-disturbed habitats (see Osman 1977). We distinguish these as “mobile
opportunists” (see also MacArthur 1960, Grassle & Grassle 1974). All such mobile
opportunists are r-strategists with life-cycle traits of small size, high fecundity, rapid
growth and high mortality.
Under the stable conditions that occur at the other end of the environmental
continuum, the community is controlled mainly by biological interactions, rather than by
extremes of environmental variability. Here the organisms have an “equilibrium strategy”
in which they are selected for maximum competitive ability in an environment that is
already colonized by many species and in which space for settlement and subsequent
growth is limiting. Such organisms are designated “K-strategists” or “ equilibrium species
” and devote a larger proportion of the resources to non-reproductive processes such as
growth, predator avoidance and investment in larger adults (MacArthur & Wilson 1967,
Gadgil & Bossert 1970, McCall 1976). Between these two extremes are communities
whose species may be intermediate between those that occur at the extremes of the
environmental gradient and have different relative proportions of opportunis tic r-
strategists and equilibrium K-strategists. The characteristics of r-selected and K-selected
equilibrium species are summarized in Table 1 (based on McCall 1976, Rees & Dare
1993), although it should be emphasized that the distinction is to some extent an arbitrary
one, and is blurred in habitats that are subject to only mild environmental disturbance.
6
Table 1 Table summarizing the population characteristics of r-selected

opportunists and K-selected equilibrium species (based on Pianka 1970,
McCall 1976, Rees & Dare 1993, Holt et al. 1995).
Changes in the structure and physical size of the infauna along a gradient of
environmental conditions have been described in relation to organic pollution by Pearson
& Rosenberg (1978) and in relation to physical disturbance by Rhoads et al. (1978),
Oliver et al. (1980) and by Gray & Pearson (1982). These are illustrated in a schematic
diagram in Fig. 2. Essentially, such studies show that community composition of benthic
infauna (those that live within the deposits) along an environmental gradient is the result
of a complex interaction between physico-chemical factors that operate at one end of the
gradient and biologically-controlled interactions under the more uniform environmental
conditions that occur in deeper waters (see Sanders 1969, Boesch & Rosenberg 1981).
7
Figure 2 Pictorial diagram showing the ecological succession that characterizes benthic com-
munities through a gradient of environmental disturbance. Note that in highly disrupted
environments (on the left side of the diagram) few organisms may be capable of survival. In
polluted or semi-liquid muds the sediments are colonized by few (resistant) species but which can
attain very high population densities. As the stability of the environment increases, these
opportunistic r-selected species are replaced by increased species variety, including slower growing
K-selected species. Finally in environments of high stability the community is dominated by
equilibrium species with complex biological interactions between members of the community.
(Based on Pearson & Rosenberg 1978, Rhoads et al. 1978).
8
The large species that comprise the burrowing infauna of stable habitats and those with
low organic content maintain oxygen levels in the deposits down to considerable depths
(see Flint & Kalke 1986) and often have complex interactions with neighbouring species,
including smaller species whose survival depends on their association with large
burrowing components (Fig. 2). The importance of bioturbation in both enhancing species
diversity and in exclusion of potentially competitive species has been widely documented
(Gray 1974, Rhoads 1974, Lee & Swartz 1980, Carney 1981, Rhoads & Boyer 1982,
Thayer 1983). Comprehensive reviews by Pearson & Rosenberg (1978) and Hall (1994)
summarize the impact of disturbance by a wide variety of factors including storms,
dredging, fishing and biological activities on benthic community structure.
Biological interactions may also control community composition on the surface
of the deposits. The presence of surface-dwelling bivalves, for example, may allow
colonization by barnacles, ascidians and other epifaunal species that would not otherwise
occur in the surface of the sediments. In other stable habitats, the activities of suspension-
feeding mussels produce consolidated silt deposits that then allow deposit feeders such as
the polychaete Amphitrite, the burrows of which in turn provide specialized shelter for the
commensal scale worm, Gattyana (Newell, 1979).
Several studies have shown that the activities of the infauna may also inhibit,
rather than facilitate, the occurrence of potential competitors for space. In an important
study by Rhoads & Young (1970) it was shown that the benthic environment may be
significantly modified by the burrowing and feeding activities of deposit-feeding
organisms. This bioturbation results in the production of an uncompacted surface layer of
faecal material that may result in the transfer of fine material to the sediment-water
interface by turbulent mixing (Wildish & Kristmanson 1979, Snelgrove & Butman 1994)
and may lead to the exclusion of potential competitors by deposit feeders (Woodin 1991,
Woodin & Marinelli 1991). This inhibition of one type of population by the activities of
another has been termed “amensalism” by Odum & Odum (1959) and has since been
described in many habitats (Aller & Dodge 1974, Nichols 1974, Driscoll 1975, Eagle
1975, Johnson 1977, Myers 1977a,b, Brenchley 1981, de Witt & Levinton 1985, Brey
1991, Flach 1992).
Loss of these “key species” in K-dominated equilibrium communities following
dis -turbance by dredging or other activities can lead to a collapse of the entire
biologically-accommodated community even though individual species may be apparently
tolerant of environmental disturbance. The colonial polychaete Sabellaria spinulosa for
example, provides a complex habitat that is associated with a wide variety of species
which would not otherwise occur (see Holt et al. 1995). This polychaete undergoesa
natural cycle of accretion and decay along with the associated community with a
periodicity of from 5-10yr (Wilson 1971, Gruet 1986). Disturbance of communities that
are dominated by K-strategists may therefore take many years for recovery of their full
community composition even though recolonization by individual components may occur
comparatively rapidly.
As the amount of organic matter in the sediments increases along a gradient
towards the fine silts and muds that characterize estuarine habitats, the larger species and
deep-burrowing forms are replaced by large numbers of relatively inactive small
suspension-feeding and surface deposit feeders including polychaete worms, bivalves and
holothurians. This reduction in the species diversity and extent of sediment bioturbation
results in an increased sediment stability and a restriction of the oxygenated layer to the
surface of the sediments. Species in the intermediate parts of the environmental gradient
shown in Figure 2 are thus relatively smaller than their counterparts in deeper waters and
9
comprise a “transitional community” that is confined to a restricted habitat in the surface

oxygenated layer of sediment and includes components that have many intermediate
characteristics between typical r-selected opportunists and K-selected equilibrium species.
Because the K-selected components in the community live for longer, the
individuals must be able to tolerate short-term changes in environmental conditions
including siltation. They therefore have generally wider limits of physiological tolerance
than r-selected opportunistic species that respond to environmental change by selection of
genetically adapted components of the population during each of the many reproductive
cycles per year.
The transitional community comprises more species than the equilibrium
community shown in Figure 2 because of invasion by opportunistic species, but the
species variety and mean size rapidly decline as the organisms are increasingly crowded
into the upper oxygenated layer at the sediment - water interface. The region between this
transitional community and those dominated by large populations of a restricted variety of
small opportunists has been referred to as the “ecotone point” by Pearson & Rosenberg
(1978) and is shown in Figure 2.
Finally at the extreme end of the physical gradient shown in Figure 2, there is a
further restriction of habitat space to the upper oxygenated layer of sediment. This results
in a progressive elimination of species and to communities dominated by opportunistic r-
strategists that are selected for small size, high fecundity and an ability to recolonize
rapidly following catastrophic mortality (see Pearson & Rosenberg 1978, Gray & Pearson
1982). Very high population densities of these r-selected opportunists can occur (the
“Peak of Opportunists” in Figure 2) before these decline as organic pollution or high
environmental disturbance eliminates even these rapid colonizers.
A useful tool for determining the extent of impact of environmental impact from
a variety of sources is a plot of the proportional contribution of each species in the
community to the overall population density of the assemblage as a whole. These curves
have been designated “K-dominance curves” by Lambshead et al. (1983) and have been
widely used in environmental impact studies in recent years (Warwick 1986, Clarke &
Warwick 1994). Obviously the equilibrium communities characteristic of undisturbed (or
unpolluted) environments have a high species diversity and each com-ponent species
makes a relatively small contribution to the overall population density. Conversely, as a
point source of disturbance is approached the (sensitive) species are replaced by large
numbers of those (resistant) members of the community that are capable of survival. This
can lead to as mu ch as 80-90% of the population being dominated by only one or two
opportunists or r-selected species at the Peak of Opportunists shown in Figure 2.
A typical set of results taken from one of our surveys of coastal communities in
the eastern English Channel is shown in Fig. 3 (Newell & Seiderer, 1997c). From this it
can be seen that as much as 78% of the community in unstable, unconsolidated, mobile
deposits at Site 1 was represented by just one species, the opportunist amphipod
crustacean Ampelisca brevicornis, and that additional species each made only a
relatively small contribution to the population. Further along the gradient of sediment
stability in mixed sands and shells at Site 2, the dominance by one species (Sabellaria sp.)
alone was approximately 45%. Finally, in the stable environmental conditions of coarse
gravels and boulders at Sites 3 and 4 there was a very large species variety of over 300
and a relatively uniform species distribution with dominance values of only 12-15%.
10
Figure 3 A set of typical K-dominance curves showing the proportional contribution of individual
species to the o verall community in fine mobile muds, in mixed sands and shells and in stable
habitats of coarse gravels and boulders off the Kent coast at West Varne in 1996. The fine mobile
muds are dominated by the opportunist amphipod crustacean, Ampelisca brevicornis , whereas the
more stable deposits have a higher total species complement each of which makes a relatively small
contribution to the overall population density. (Based on Newell & Seiderer 1997c).
Estimation of K-dominance curves adjacent to dredging works and other point

sources of environmental disturbance is thus potentially useful because it can be used as a
relatively simple index to define the area of immediate impact. It can also be used to
determine whether this is enlarging or decreasing with time, without the necessity of the
complex analysis of community structure that is required for interpretation of the wider
impact on community structure in the transition zone.
These distinctions between the lifestyles and adaptive strategies of opportunists
and K-selected equilibrium species are of fundamental importance because they go some
way towards accounting for the differences in the rate of recovery that has been recorded
for biological resources following disturbance by episodic events such as dredging.
Clearly, the species composition and rate of recovery of biological communities following
cessation of dredging will depend to a large extent on whether the original communities
were dominated by opportunists or equilibrium species, and on the time that is required to
develop the complex associations which characterise interactions between the K-
dominated equilibrium community. Knowledge of the key fauna1 components and their
lifestyle thus allows some predictions on the impact of dredging and spoils disposal
11
on biological resources and on the subsequent rate of recovery of marine community

composition following cessation of dredging.
Ecological succession and the recolonization process
These general features of the structure of benthic communities apply not only to
successional stages along a gradient of environmental variability, but also to the
successive sequence of populations that recolonize deposits after the cessation of
environmental disturbance. McCall (1976) and Rees & Dare (1993) have recognized the
occurrence of three main types of benthic components of marine communities based on
the distinction between r-selected opportunistic species and K-selected equilibrium
species. Group I species comprise those that colonize first after a community has been
removed by disturbance. They comprise large populations of small sedentary tube-
dwelling deposit feeders that have rapid development, many generations per year, high
settlement and death rates. Examples include the polychaete worms Streblospio, Capitella
capitata, and Owenia fusiformis as well as the amphipod Ampelisca. That is, the Group I
community comprises mainly r-strategist opportunistic species.
Group II species comprise mainly bivalve molluscs such as Tellina, Nucula and
Abra, the tube worm Lagis (=Pectinaria) and the common starfish (Asterias rubens).
There is no absolute distinction between this community and the primary colonizers, but
the components attain a lower peak abundance than the smaller opportunistic species and
have slower recruitment and growth rates. Finally Group III species comprise larger slow-
growing K-strategist equilibrium species such as the polychaete Nephtys, the reef-forming
Ross worm (Sabellaria), razor shell (Ensis), sea urchins such as Echinocardium and
Echinus, scallops (Pecten), the ocean quahog (Arctica islandica), the edible crab (Cancer
pagurus) and larger burrowing crustaceans such as Nephrops and Callianassa.
The changes in species variety, abundance of individuals and biomass during the
recolonization process are shown in Fig. 4. Inspection of this figure shows that initially
the sediments are almost devoid of benthic macrofauna.
The initial colonizing species are few, but the number of individuals (population
density) increase rapidly with time to a peak of (Group I) opportunistic species. As time
passes, the short -lived opportunistic species (r-strategists) decrease in numbers and
biomass as more species invade the area. This transition point where the community is
poor in population density and biomass is the same ecotone point shown on the spatial
gradient in Figure 2.
12
Figure 4 Schematic diagram showing a colonization succession in a marine sediment following

cessation of environmental disturbance. Initial colonization is by opportunistic species which reach
a peak population density generally within 6 months of a new habitat becoming available for
colonization after the catastrophic mortality of the previous community. As the deposits are invaded
by additional (larger) species, the population density of initial colonizers decliners. This ecotone
point marks the beginning of a transitional community with high species diversity of a wide range of
mixed r-s elected opportunistic and K-selected equilibrium species. This period may last for l-5 yr
depending on a number of environmental factors, including latitude. Provided environmental
conditions remain stable, some members of this transition community are eliminated by competition
and the community as a whole then forms final equilibrium community comprising larger, long-
lived and slow growing species with complex biological inter-actions with one another. (Based on
Pearson & Rosenberg 1978).
Prior to this, the community is characterized by large populations of a few small

opportunistic species; after this time the species variety increases, as does the biomass, but
the population density declines. This Group II community is a transitional one where the
maximum number of species has invaded the newly-available space, and is followed by a
phase where some species are eliminated by competition and the community returns to the
(somewhat lower) species composition and biomass characteristic of the undisturbed
Group III community.
The sequence shown in Figure 4 indicates that colonization is likely to follow a
definite time course of progressive invasion by large numbers of opportunistic species in
the first instance, followed by a wider species diversity during the transitional phase and
finally by a consolidation phase when competition between the K-strategist equilibrium
species for the limited space available results in the elimination of some of the transitional
colonizers (see also Warwick et al. 1987). The biological diversity in any particular
community will then reflect the frequency of disturbance and represent a balance between
13
invasion and subsequent growth of colonizers, and losses by extinction and displacement
(see Huston 1994). In areas where environmental disturbance is unevenly distributed, this
may lead to a mosaic of communities, each at different stages of the successional
sequence shown in Figure 4 (see Johnson 1970, Grassle & Sanders 1973, Whittaker &
Levin 1977, Connell 1978), and may partly account for the patchiness of marine
communities in dredged areas.
The time taken for recovery of the full species composition and for subsequent
exclusion of some of the transition community following the growth of larger K-strategist
equilibrium species in a particular area will depend largely on the components that occur
under natural conditions. In shallow water and estuarine conditions, where the community
is in any case dominated by opportunistic species, recovery to the original species
composition may be very rapid and coincide with the Peak of Opportunists point in Figure
4. In the stable environmental conditions of deeper waters, the replacement of the initial
colonizers in the transitional community following complex biological interactions
between the K-selected equilibrium species may take several years.
The physical impact of dredging

Impact within the dredged area
The increased exploitation of marine deposits and the physical impact of dredging works
has been widely reviewed (see Dickson & Lee 1972, Shelton & Rolfe 1972, Cruikshank &
Hess 1975, Eden 1975, Millner et al. 1977, de Groot 1979, van der Veer et al. 1985,
Glasby 1986, Lart 1991, Gajewski & Uscinowicz 1993, ICES 1993, Land et al. 1994,
Whiteside et al. 1995, Hitchcock & Drucker 1996). Most of the sea-going aggregate
dredgers are self-contained and use a centrifugal suction pump to lift the aggregates from
the sea bed into a hopper where the material is screened before being transferred to the
hold. The in situ reserves for economic exploitation normally range from 15-55% gravel.
Unless the material is otherwise suitable for direct use as a beach feed or landfill (see Hess
1971), the sand : gravel ratio in the final cargo is adjusted to between 50 : 50 and 65 : 35
depending on customer requirements, local geology and ship performance (A. Hermiston
pers.comm).
A proportion of the dredged deposits may therefore be returned to the sea bed
through reject chutes when there is a larger proportion of fine material than is required for
a commercially viable cargo. In most aggregate deposits the fines comprise only l-2% of
the total and are dominated by the silt fraction although significant quantities of sand may
also be discharged in the immediate vicinity of the dredger to increase the gravel
component of the cargo. Overspilling of water via spillways from the hopper will also
contain some fine sands that are maintained in suspension by the turbulence within the
hopper.
Essentially the physical impact of dredging works is dependent partly on the
method of dredging, and partly on the amount and grade of deposits rejected by screening
(if used) and overspill from the hopper. Two main methods of dredging are used for
gravels extraction in European coastal waters. These are anchor hopper dredging and
trailer suction hopper dredging and are illustrated in Fig. 5. In anchor dredging thevessel
is stationary and dredges the deposits from a sequence of specific points on the sea floor
14
Figure 5 Diagram showing the two principal methods of dredging for marin e aggregates in
European coastal waters. Upper diagram shows the furrows left on the sea bed by trailer suction
dredging while the vessel is under way. In this case the sea bed is crossed by a series of tracks which
are 2-3m wide and up to 50cm deep. Lower diagram shows the pits left on the sea bed by anchor
dredging. In this case the vessel is anchored and the dredged pits may reach as much as 75 m in
diameter and 20 m in depth. (Based on Dickson & Lee 1972).
and can therefore leave pits or depressions on the sea bed that may reach as much as 20 m
in depth and 75 m in diameter (Cruikshank & Hess 1975, Dickson & Lee 1972).
15
Because the deposits required for marine aggregates are coarse, and sediment
disturbance by wave action is in any case limited mainly to depths of less than 30m even
during storm conditions, it follows that not only is the fauna likely to be removed in
patches from the dredged areas, but such pits are likely to be persistent features of the sea
bed topography for several years except in areas where the sands are mobile (Eden 1975).
Dickson & Lee (1972) studied the recovery of test pits dug by anchor dredge in gravel
deposits in the Hastings Shingle Bank off the south east coast of England. They found that
the pits were very slow to fill and were still visible after two years. In another study, van
der Veer et al. (1985) described the recovery of pits in sandy sub-strata in the Dutch
Wadden Sea. They showed that in this instance pits in channels with a high current
velocity filled within one year, but those in the lower current velocities which occur in
tidal watersheds took 5-10yr to fill whereas those in tidal flat areas were still visible after
15 yr.
Such sediment movement as does occur is mainly through slumping of the sides
of the furrows and subsequent infilling by fine particulates transported by tidal currents
into the furrows that reduce current velocity and act as sediment traps. This can lead to
heavily anoxic sediments and to colonization by a community which differs considerably
from that in the original deposits (Dickson & Lee 1972, Shelton & Rolfe 1972, Kaplan et
al. 1975, Bonsdorff 1983, Hily 1983, van der Veer et al. 1985, Hall 1994).
A second method of marine aggregate dredging is for the ship to extract deposits
by suction through one or two pipes deployed while the vessel is slowly under way
(Figure 5). In this case, side-scan sonar records show that the seabed within the
boundaries of licenced extraction areas in the southern North Sea is crossed by a series of
dredge tracks that are 2-3 m wide and up to 50 cm deep (van Moorsel & Waardenberg
1990, Kenny & Rees 1994) although deeper troughs of up to 2m were recorded from areas
where the dredge head had crossed the area several times. Davies & Hitchcock (1992)
reported dredge cuts of between 20-55cm depth and 3-3.8m width in commercially
exploited deposits of the Bristol Channel. Somewhat deeper troughs of up to 70cm were
reported for the Baltic (Gajewski & Uscinowicz 1993). In this case removal of the surface
0.5 m of deposit would be sufficient to eliminate the benthos from the deposits in strips,
the total removal depending on the intensity of dredging at a particular worked site.
Despite the shallower depth of removal, the evidence suggests that infilling of the
troughs from trailer suction dredging takes at least 12 months in the Baltic and is achieved
partly by slumping from the sides and partly by transport of fine material by bottom
currents into the sediment traps formed by the dredged furrows (Kaplan et al. 1975, Hily
1983, van der Veer et al. 1985, Gajewski & Uscinowicz 1993). Progressive removal of the
original sandy gravel and its replacement by fine sand has also been reported for the
sediments off Dieppe by Desprez (1992). In the case of experimental furrows dredged by
trailer suction in gravel deposits of the southern North Sea off the Suffolk coast of
England, even shallow depressions of only 20-30cm depth were still visible on side-scan
sonar records made up to 4yr later (Millner et al. 1977). In contrast, dredge furrows in the
Bristol Channel have been reported to disappear within 2 to 3 tides because of high
sediment mobility (pers. obs.).
Rather unexpectedly, Kenny & Rees (1994, 1996) found an increase in the
particle size of deposits in the dredged areas, possibly reflecting the exposure of coarse
deposits at depth below the surface gravel layers. In this study, which was carried out in
the southern North Sea, the dredged furrows were visible with side-scan sonar even after
2yr. Similar results have been reported for dredging tracks off the French coast at Dieppe
(Desprez 1992) although winter storms obliterated tracks within a few months on the
16
Klaver Bank in the Dutch sector of the North Sea (Sips & Waardenberg 1989, van
Moorsel & Waardenberg 1990, 1991). In general, dredge tracks will persist for varying
times depending on the rate of local sediment fluxes, recent measurements suggesting l-4
days only for the Norfolk Banks, but periods as long as l-4yr for more stable deposits off
the Owers to the east of the Isle of Wight (A. Hermiston pers.comm.).
Thus both anchor dredging and trailer suction dredging have an important
potentialimpact on the biology of the dredged areas, since no benthos is likely to occur
below the dredged depth. This can be expected to lead to a patchy distribution of
organisms, reflecting the differences between the dredged furrows and the intervening
undredged surfaces. Such recolonization as occurs within the dredged areas is likely to be
by migration of adults through transport on tidal currents (Rees et al. 1977, Hall 1994); by
transport in sediments slumping from the sides of the pits and furrows (McCall 1976,
Guillou & Hily 1983); by the return of some undamaged components through outwash
from the chutes and spillways (see Lees et al. 1992, Ministry of Agriculture, Fisheries and
Food 1993); and by colonization and subsequent growth of larvae from neighbouring
populations. In this case, a clear succession of colonizing species is to be anticipated,
leading to the establishment of definite clusters or patches in benthic community
composition, depending on the type of deposits that have infilled the dredged areas and
the time since the recolonization sequence started.
Impact adjacent to the dredged area
Although a good deal of concern has been expressed about the possible impact of marine
aggregate extraction on coastal resources (see ICES 1992a,b), the possible scale of impact
outside the immediate dredged area from the settlement on the sea bed of fine material
temporarily suspended by marine aggregates dredging is poorly understood.
Hitchcock & Drucker (1996) have summarized values for material lost through
the hopper overflow spillways and from the reject chutes during the screening process on
a typical modern trailer suction dredger of 4500 t hopper capacity operating in UK waters
off the coast of East Anglia. Table 2 shows the size distribution for the material lost
through the reject chute and spillway, while the total screened load quantities are
summarized in Table 3. These show that during a recorded average loading time of 290
min, 12 158 t of dry solids and 33 356 t of water were pumped by the dredge pump .
17
The data show that 4185 t of dry solids are retained as cargo, while 7223 t of dry solids
are returned overboard because of rejection by screening, and a further 750t from
overspill.
It is also clear that some 1338 t of material >2.0mm (representing 18.5% of the
7235 t in the reject chute - see Table 2) is lost overboard through the reject chute and a
further 126 t (representing 16.9% of the 750 t in the spillway - see Table 2) from the
spillway. This equates to a loss rate of 76.9 kgs-1 of particles >2.0mm from the screening
reject chute and 7.2 kg s-1 from overspill. Assuming that the dredger moves at an average
speed of 1 knots, the flux of material >2.0mm entering the water column is 39.6 kg s-1 m-1
from the screening reject chute and 3.7 kg s-1 m-l from the spillways. Much of this material
is in the size range 2.0-10.0mm and falls rapidly to the seabed with little horizontal
displacement during screening. Video recordings during normal loading operations show
that such material deposits on the seabed directly under the dredge vessel (Davies &
Hitchcock, 1992).
Finer sand and silt fractions discharged during dredging and screening amount to
5824 t from the reject screening chutes and 338 t from overspill. This is equivalent to a
deposition rate of 334 kg s-1 and 19.4 kg s-1 , respectively. In addition to the sand fraction,
up to 213 t (12.2 kg s-1 ) of muddy sediment (<0.063 mm diameter) may be lost through
the rejection process and 285 t (16.4 kg s-1 ) from overspill. The material may be expected
to settle more slowly than the sand-sized fraction and has a typical settling velocity of 0.l-
l.0mm s -1 .
In its simplest form, the settlement velocity and residence time of such particles
in the water column can be estimated from Stoke’s Law. If the residence time of particles
in the water column is known, the duration and speed of currents will then determine the
excursion pattern before settlement. Estimates of dispersion of fine material based on
these Gaussian diffusion principles suggest that very fine sand particles may travel up to
11 km from the dredge site, fine sand up to 5 km, medium sand up to 1 km and coarse
sand less than 50m (H. R. Wallingford 1994, cited in Hitchcock & Drucker 1996).
Similar estimates based on the settlement velocity of fine silt-sized particles (
<0.063 mm diameter) suggest that this material could remain in suspension for up to 445
tidal cycles and be carried for as much as 20 km on each side of a point source of
discharge. Most recent studies made on the dispersion of sediment plumes generated from
dredging operations suggest, however, that the area of impact of outwash from dredging
activities is smaller than estimates based on Gaussian diffusion models, especially where
the proportion of silt and clay in the deposits is low. This appears to be due to complex
cohesion properties of the discharged sediment particles that settle to the seabed as a
density current and do not conform to settlement rates based on the specific gravity and
size of the component particles themselves.
18
A detailed study by Gajewski & Uscinowicz (1993) in relation to trailer dredging

in the Baltic showed that the width of the plume, as determined by the light extinction in
the water, did not exceed 50m. Settlement of the suspended matter onto the sea floor was
measured by a series of sediment traps deployed approximately 1 m above the sea bed.
When the dredger was discharging mainly fine sand (0.25-0.125 mm) at a concentration
of approximately 11000 mg l-1 , it was found that deposition of 7500- 15 000g m-2 were
recorded in the troughs during dredging. The settlement in traps deployed 50m away from
the dredger’s route was, however, less than 1220gm-1 . At distances greater than 50m the
amount of material settling on the sea floor decreased rapidly (Fig. 6).
More recently, Acoustic Doppler Current Profiling (ADCP) techniques have
been used to determine plume dispersion in relation to spoils dispersal from both
commercial aggregate dredgers (Hitchcock & Dearnaley 1995, Hitchcock & Drucker
1996) and in relation to capital dredging works and sand mining (Land et al. 1994,
Whiteside et al. 1995). Remote airborne and satellite imagery has also proved to be a
useful tool in defining the contours of sediment dispersal (Whiteside et al. 1995).
Figure 6 Diagram showing the settlement of sand during dredging operations from trailer dredging
in the Baltic. Particle size profiles fo r the sediments deposited in the track of the dredger and 50m
on each side of the dredger are also shown. Note that the main deposition of sand from flow-off
from dredging operations was confined to distances within 150m on each side of the track of the
dredger. (After Gajewski & Uscinowicz 1993).
19
These studies confirm that the initial sedimentation of material discharged during outwash
from dredgers does not, as had been widely assumed, disperse according to the Gaussian
diffusion principles used in most simulation models, but behaves more like a density
current where particles are held together by cohesion during the initial phase of the
sedimentation process. As a result, the principal area likely to be affected by sediment
deposition is much less than the “worst case” scenarios predicted from conventional
Gaussian diffusion simulation models, and is mainly confined to a zone of a few hundred
metres from the discharge chutes.
A recent study by Whiteside et al. (1995; see also Johanson & Boehmer 1975,
Gayman 1978) has shown that the behaviour of plumes discharged during sand dredging
can best be regarded as comprising an initial “dynamic phase” during which the sediment-
water mixture descends rapidly to the sea bed as a density current jet at a rate that depends
on the overflow density, the diameter of the discharge pipe, the water depth, the velocity
of discharge and the speed of the dredger. During its passage through the water column
and following impact with the sea bed the sediment is dispersed into the water and forms a
well-defined plume astern of the dredger. This second longer phase has been referred to as
the “passive phase” of dispersion by Whiteside et al. (1995) and starts approximately
10min after outflow. During this phase the material behaves in a relatively simple settling
mode according to Stokes’ Law, the plume then decaying to background levels after a
period of 2-3 h.
Their study showed that approximately 100 m (corresponding to approximately 3
min from the overflow) astern of a dredger working in Hong Kong waters the plume
surface sediment concentrations were from 75-150mgl-1 . Levels were halved in 10 min
and reduced to 20-30 mg l-1 after 30 min. This approached the recorded background
suspended solids concentration of l0-15 mgl-1 and indicated that only a relatively small
proportion of the fines category (< 63 µm) remained in the water column at the start of the
passive phase of dispersion 10min after discharge. Even then, their data suggest that the
settlement rate of the plume continued to be more rapid than simple particle settlement
would suggest.
A plume dispersion model developed by Whiteside et al. (1995) for the surface
layer (the upper 8 m of the water column) for up to 40 min after discharge is shown in Fig.
7 and compares well with summed plume decay measurements in the vicinity of the
dredger. The contours for sediment deposition evidently remain as a narrow band
extending for approximately 100m on each side of the track of the dredging vessel, much
as recorded by Gajewski & Uscinowicz (1993) for Baltic waters.
Very similar rapid rates of deposition and decay of sediment outwash plumes
have been recorded by Hitchcock & Drucker (1996) who studied plume generation and
decay from four dredge vessels ranging in capacity from 2000-5000 t during normal
loading operations off the coast of East Anglia, UK. During the plume tracking exercise
peak current velocities reached 0.6 m s-l and the water depth was approximately 22m. The
concentration of total suspended sediment discharged was approximately 2500mg -1 l
comprising mainly sand-sized material and with < 30mg l-1 mud (< 0.063 mm diameter).
The total concentration of suspended solids in the water column at different
depths and distances from the dredger measured by water sampling and optical
transmis someters is summarized in Fig. 8. The corresponding values for silt-sized material
(<0.063 mm) are shown in Fig. 9. These data show that concentrations of sand-sized
material are reduced to background levels over a distance of only 200-500m from the
point of release into the water column and that the concentration of even silt-sized
particles is reduced to background values of 2-5 mg 1-l over a similar distance. This very
20
Figure 7 Diagram showing contours for suspended sediment concentrations astern of a trailer
dredger operating in Hong Kong waters. Upper diagram shows the contours as measured in the
upper 8m of water across the plume at various arbitrary time intervals during field studies of a
sediment plume from 16-19min after discharge. Lower diagram shows the output of a simulation
model developed for sediment dispersion based on rapid initial sedimentation during a dynamic
phase and a second longer passive phase which starts approximately 10min after outflow. (After
Whiteside et al. 1995).
rapid reduction in suspended sediment concentrations is similar to that reported by

Whiteside et al. (1995).
21
Figure 8 The total concentration of suspended solids in the water column at different depths and
distances from the dredger measure d by water sampling and optical transmissometers. (After
Hitchcock & Drucker 1996).
Although suspended sediment concentrations in the plume were not significantly different
from background levels beyond 400-500 m from the point source of discharge using
conventional water sampling and optical transmissometer techniques, it is interesting to
note that it is possible to track the plume using ADCP techniques over a distance of up to
3.5 km. A series of typical sections across a discharge plume at varying distances from
80m up to 3335 m away from a dredger is shown in Fig. 10. This shows clearly the decay
of the plume to background levels at approximately 400m, but that a residual impact is
detectable up to 3.5 km by ADCP methods.
Figure 9 The concentration of silt -sized material (<0.063 mm) in the water column at different
depths and distances from the dredger measured by water sampling and optical transmis someters.
(After Hitchcock & Drucker 1996).
22
Figure 10 Acoustic Doppler Current Profiler (A DCP) acoustic backscatter across the plume of a
dredger at varying distances downstream (m). Water speed 100cm s -1. Note that although
sedimentation is achieved within 880m, the relative backscatter suggests an additional “plume”,
perhaps representing entrained air bubbles, biochemical precipitates or organic matter released from
the sediment, which extends up to 3380m astern of the dredger. (After Hitchcock & Drucker 1996).
23
Whether this residual impact is caused by air bubbles or organic matter becoming
entrained into the water column during the dredging operation is at present unknown.
However, it is noteworthy that there is now a good deal of evidence that suggests that
disturbance of marine sediments may release sufficient organic matter into the water
column to enhance benthic production.
These results for the dispersion of sediment in the water column thus suggest that
sedimentation is rapid and is confined to the immediate vicinity of the discharge. They
confirm earlier studies of Poiner & Kennedy (1984; see also Willoughby & Foster 1983)
who reported that sediment deposition generated from dredging activities in Moreton Bay,
Queensland was confined to the immediate vicinity of the dredging works. Concentrations
of suspended sand-sized material were reported to decay to background levels over a
distance of only 200-500m from the point of release into the water column from a
commercial aggregate dredger. They estimated that the sediment deposition 500m outside
the boundary of the dredged area was 29.6 kgm-2 (23mm m-2 ). At 1 km deposition was
21.2kg m-2 (16mm m-2 ), a t 1.5km it was 15kg m-2 (12mm m-2 ), at 2km it was 10.7kg m-2
(8mm m-2 ) and finally at 2.5km from the boundaries of the dredged area the estimated
deposition was less than 7.6kg m-2 (6mm m-2 ).
There is a good deal of evidence from other surveys that disturbance of
sediments by dredging may release sufficient organic materials to enhance the species
diversity and population density of organisms outside the immediate zone of deposition of
particulate matter. Disturbance of the sediments may thus enhance benthic production
outside the immediate zone of deposition provided that contaminants from polluted
sediments are not associated with the disposal of spoils. Stephenson et al. (1978) and
Jones & Candy (1981) both document the enhanced diversity and abundance of benthic
faunas near to dredged channels. Poiner & Kennedy (1984) showed that there was an
enhancement of benthic biota close to dredged areas at Moreton Bay, Queensland and that
the level of enhancement decreased with increasing distance from the dredged area up to a
distance of approximately 2 km. They ascribe this to the release of organic nutrients from
the sediment plume, a process which is well known from other studies (Ingle 1952, Biggs
1968, Sherk 1972, Oviatt et al. 1982; Walker & O’Donnell 1981).
The results reviewed above thus suggest that the impact of dredging activities
mainly relates to the physical removal of substratum and associated organisms from the
seabed along the path of the dredge head and to the impact of subsequent deposition of
sediment from outwash during the dredging process. The evidence from direct studies on
the sedimentation of particulate matter suggests that the impact of sedimentation on
biological resources on the seabed is likely to be confined to distances within a few
hundred metres of the dredger where the deposits are sands and gravels. It should be
remembered, however, that discharge of dredge spoils from maintenance and capital
dredging works in estuaries may result in much larger dispersion plumes that reflect the
dominantly fine particles and strong current flows that occur in estuaries, and that the
same processes, which result in the release of dissolved organic matter, can also result in
the release of bound surface contaminants from the sediments into the water column.
24
The impact of dredging on biological resources
Sensitivity to Disturbance
The impact of disturbance by the dredge head during marine aggregate dredging has been
reviewed on pp. 14-17. The effects of sediment deposition and spoils disposal outside the
immediate boundaries of dredged areas in coastal waters has also been widely studied and
includes extensive physiological-ecological work on a wide variety of animals including
plankton, benthic invertebrates and fish species (for reviews, see Sherk 1971, Moore
1977). Early studies by Loosanoff (1962) showed that different species of commercially
significant filter-feeding molluscs were differently affected by suspended sediment.
Subsequent studies by Sherk (1971) and Sherk et al. (1974) included both plankton and
fish species. They showed that, as in the case of bivalves, fish species have varying
tolerances of suspended solids, filter-feeding species being more sensitive than deposit
feeders and larval forms being more sensitive than adults (see also Matsumoto 1984).
Many of the macrofauna that live in areas of sediment disturbance are well
adapted to burrow back to the surface following burial (see Schafer 1972). Studies by
Maurer et al. (1979) showed that some benthic animals could migrate vertically through
more than 30cm of deposited sediment, and this ability may be widespread even in
relatively deep waters. Kukert (1991) showed, for example, that approximately 50% of the
macrofauna on the bathyal sea floor of the Santa Catalina Basin were able to burrow back
to the surface through 4-10cm of rapidly deposited sediment. A good deal of the apparent
recolonization of deposits following dredging or spoils disposal may therefore reside in
the capacity of adults to migrate up through relatively thin layers of deposited sediments
(see also Ellis & Heim 1985) or to migrate in during periods of storm-induced disturbance
(see Hall 1994).
There is good evidence that the activities of filter-feeding bivalves, in particular,
can play an important part in controlling the natural phytoplankton and seston loads in the
water column (Cloern 1982) to an extent that food may become a limiting resource in the
benthic boundary layer at the sediment-water interface (Wildish & Kristmanson 1984,
Frechette et al. 1989, 1993; see also Dame 1993, Snelgrove & Butman 1994) as well as on
coral reef flats and in cryptic reef habitats (Glynn 1973, Buss & Jackson 1981). Because
the suspension-feeding component is evidently highly effective in removing particulate
matter from sea water, the release of large quantities of suspended matter can lead to a
loss of suspension-feeding components through clogging of the gills. This has led to a
corresponding increase in the community of deposit feeders in some areas such as St
Austell Bay off the southwest coast of England (Howell & Shelton 1970).
In general, however, most recent studies of filter feeders that live in coastal
waters show that bivalves, in particular, are highly adaptable in their response to increased
turbidity such as can be induced by periodic storms, dredging or spoils disposal and can
maintain their feeding activity over a wide range of phytoplankton concentrations and
inorganic particulate loads (Shumway et al. 1985, 1990, Newell et al. 1989, Newell &
Shumway 1993 Iglesias et al. 1996, Navarro et al. 1996, Urrutia et al. 1996). Although
these studies on the physiology of individual species can give some insight into the
differing susceptibilities of the macrofauna to increased turbidity, or to burial from
dredger outwash, in general it is difficult to make predictions of the impact of dredging on
whole communities from the results of studies on individual species. Partly for this
25
reason, and because the interactions between the components of natural populations are
complicated in space and time, most recent studies on the impact of spoils disposal and
dredging works have been carried out on whole communities, rather than individual
species. Such studies have concentrated on three main features of benthic communities,
namely the number of individuals (population density), number of species (the diversity)
and the biomass (to give an index of the growth following recolonization).
Sampling is conventionally carried out by means of a grab that allows collection
of a sediment sample from a known area of seabed deposits, which are then eluted through
a l-mm mesh sieve to extract the macrofauna. Sediment samples from fine deposits such
as occur in coastal embayments, lagoons and estuaries are relatively easy to obtain by
means of equipment such as the van Veen and Smith-McIntyre grabs, the Ponar grab
(Ellis & Jones 1980), or the Day grab whose jaws are held closed by the tension of the
wire from which the grab is suspended rather than by a spring-loaded mechanism (see
Holme & McIntyre 1984). Sampling of coarser gravel deposits is complicated, however,
by the fact that the larger stones become trapped between the jaws of conventional grabs,
leading to extensive losses through “washout” from the grab. Partly because of this
problem, most work on coarser deposits has been carried out with semi-quantitative
dredges such as the Anchor dredge (Forster 1953, Holme 1966, Kenny et al. 1991) or the
Ralier du Baty dredge used by Davoult et al. (1988).
More recently, however, Sips & Waardenburg (1989) and Kenny & Rees (1994)
have used a Hamon grab for quantitative studies on the fauna of gravels and sands. This
grab takes a scoop out of the seabed deposits, rather than relying on the closure of
opposing jaws (see Holme & McIntyre 1984). This greatly reduces the problem of fauna
losses through “washout” during the sampling process and the Hamon grab is now widely
used in the quantitative evaluation of the benthos in coarse sands and gravels.
Such studies emphasize that the macrofauna may vary considerably even over
relatively short distances, and that a proper understanding of the distribution of benthic
communities is necessary if damage to potentially important communities is to be avoided
during dredging operations. Figure 11 shows, for example, the distribution of two
important members of the benthic community in mixed gravel, sand and muddy deposits
off the coast of East Anglia in August 1996 (Newell & Seiderer 1997b). Inspection of this
figure shows that the main population of the reef-building tubeworm, Sabellaria spinulosa
(called Ross by the fishermen) occurs in the northwestern part of the survey area, and
corresponds with a localized patch of coarse stones and cobbles that give sufficient
stability to support a rich reef community. This species may be predated upon by the pink
shrimp (Pandalus) and is potentially important as a feeding ground for a variety of
demersal fish species (see Warren, 1973).
In contrast, the populations of the comb worm (Lagis = Pectinaria koreni) occur
in mobile muddy sands in the southwest of the survey area. This species is an important
prey item for sole (Solea solea), dab (Limanda limanda and plaice (Pleuronectes platessa)
(see Lockwood 1980, Basimi & Grove 1985, Carter et al. 1991, Horwood 1993, also Peer
1970) and therefore represents a food resource within the survey area that requires
conservation.
26
Figure 11 Schematic diagram of a survey area in the southern North Sea off Suffolk showing the
distribution of the comb worm (Lagis koreni) in fine deposits of the survey area, and that of the
colonial “Ross” worm (Sabellaria spinulosa ) in areas where coarse boulders provide a stable
environment for development of reef-forming species. Population density in numbers of individuals
per 0.25 m2 Hamon grab sample. (Newell & Seiderer 1997b).
27
Impact of dredging on diversity and abundance
The impact of dredging on benthic communities varies widely, depending, among other
factors, on the intensity of dredging in a particular area, the degree of sediment
disturbance and recolonization by passive transport of adult organisms (see Hall 1994)
and the intrinsic rate of reproduction, recolonization and growth of the community that
normally inhabits the particular deposits.
Some examples of the impact of dredging on the species variety, population
density (number of individuals) and biomass of benthic organisms from a variety of
habitats ranging from muds in coastal embayments and lagoons, to oyster shell deposits,
and to sands and gravel deposits in the southern North Sea are summarized in Table 4.
This shows that both maintenance dredging and marine aggregates dredging can be
expected to result in a 30-70% reduction of species diversity, a 40-95% reduction in the
number of individuals, and in a similar reduction in the biomass of benthic communities
in the dredged area.
Despite the major impact of dredging on benthic community composition within
dredged areas, there is little evidence that deposition of sediments from outwash through
the chuteways during the dredging process has a significant impact on the benthos outside
the immediate dredged area. Poiner & Kennedy (1984) showed that the population density
and species composition of benthic invertebrates adjacent to dredging works on sandbanks
in Moreton Bay, Queensland, Australia increased rapidly outside the boundaries of the
dredged area, as might be anticipated from the relatively small amounts of sediment that
are deposited beyond a few hundred metres of the dredger trail (see Figs 6 and 7 and p.
24). The population density and species diversity recorded from a transect across a
dredged area in Moreton Bay in July 1982 by Poiner & Kennedy (1984) is shown in Fig.
12.
28
Figure 12 Diagram showing the number of individuals and species of benthos recorded in July
1982 on a transect crossing a dredged area on a sublittoral sandbank in Moreton Bay, Queensland,
Australia. Note that species variety and population density increased rapidly outside the immediate
boundaries of the dredged area. (Based on Poiner & Kennedy 1982).
Other than this study, there is surprisingly little detailed information on the
precise boundaries of biological impact or “footprint” surrounding areas that have been
dredged for sands and gravels. The circumstantial evidence from the boundaries of
sediment deposition suggest, however, that biological impact is likely to be confined to
the immediate vicinity of the dredged area.
One of the problems with assessing the impact of dredging works and the
recovery of benthic communities over time is that biological communities are often
subject to major changes in population density and community composition, even in areas
that are apparently unaffected by dredging. Variations in the population density and
species composition of the large bivalve population recorded between 1988 and 1991 in
the sand and gravel deposits of the Klaver Bank in the Dutch sector of the North Sea by
van Moorsel (1994) are shown in Fig. 13. This shows the major change in population of
the bivalve, Dosinia exoleta between the summer of 1988 and the spring of 1989 and the
loss of the large bivalve, Arctica islandica from the deposits even before aggregate
extraction had taken place.
29
Figure 13 Diagram summarizing the changes in population density and species composition of
large bivalves on the Klaver Bank in the southern North Sea between 1988 and 1991. Pre -dredging
values in 1988 show major seasonal changes in density and species composition. After dredging in
the summer of 1989, large differences in population density and species composition emerged
between dredged and control areas, and these differences persisted for at least two years. (After van
Moorsel 1994).
A short period of aggregate extraction was carried out in the study area on the Klaver
Bank in the summer of 1989. Thereafter, clear differences emerged between the large
populations of bivalves in control areas outside the dredged zone and those within the
dredged area, despite the natural variations in species composition and population density
that evidently occurred in the deposits of the survey area. These differences persisted until
the end of the survey period in autumn of 1991, suggesting that this slow-growing
component of the benthos remains impacted for at least 2yr after cessation of dredging.
30
The process of recolonization and recovery

These complex changes in community structure following dredging, and which occur
during the recovery process, are difficult to assess by mere inspection of the data for
species composition, population density and biomass. Most recent studies on community
structure in relation to environmental gradients, therefore, whether these are natural or
induced by man, use relatively sophisticated analytical techniques that incorporate the
type of species as well as their individual population densities and biomass to assess
changes in community structure. The use of these techniques is beyond the scope of this
review, but useful accounts for the biologist are given in Kruskal (1977), Hill (1979),
Field et al. (1982), Heip et al. (1988), Magurran (1991), Warwick & Clarke (1991), Clarke
& Ainsworth (1993), Clarke & Warwick (1994, and references cited therein).
Probably the most widely-used method is detrended correspondence analysis
(DECORANA), an ordination technique that arranges stations along axes according to
their similarity in species composition (Hill 1979). This is often used in association with
two-way indicator species analysis (TWINSPAN) to identify species that characterize
particular parts of an environmental gradient such as might be imposed, for example, by
dredging or spoils overspill, or communities in relation to wider spatial gradients (see
Eleftheriou & Basford 1989).
A second approach is the use of non-parametric multivariate analyses of
community structure as outlined by Field & McFarlane (1968), Field et al. (1982) and
Clarke & Warwick (1994). This procedure has recently become available in a convenient
software package PRIMER (Plymouth Routines in Multivariate Ecological Research) and
is now widely used in the analysis of benthic community structure in European coastal
waters.
Despite problems in the interpretation of long-term studies on the abundance and
composition of marine communities, studies that are carried out over even relatively short
time periods can give important information on the recovery process following cessation
of dredging. The most comprehensive analysis of the impact of dredging on community
composition and on the process of recolonization and recovery in mixed gravel deposits is
that of Kenny & Rees (1994, 1996). They carried out an intensive dredging programme by
suction trailer dredger in an experimental area off Lowestoft, Norfolk in the southern
North Sea and subsequently monitored the recovery over a period of 8 months in the first
instance, although this was increased to 2yr in an extended study of the recolonization
process (Kenny & Rees 1996). Dredging occurred in April 1992, during which the suction
dredger SAND HARRIER removed a total of 52 000 t of mixed aggregates from an area
measuring 500m by 270m, an estimated 70% of the surface deposits down to an average
depth of 0.3 m having been removed from the experimental area. The species variety,
population density and biomass in the exp erimentally dredged site was then compared
with that in a reference site nearby over the 8-month period between March and December
1992.
The results from their study are summarized in Fig. 14. This shows that the
number of species in the dredged site declined from 38 to only 13 species following
dredging, whereas the number of species remained at about 35 during the 8-month period
at the reference site. The number of species in the dredged area subsequently increased
somewhat in the following 7 months, suggesting that some recolonization occurs even
over this relatively short time.
31
Figure 14 Graphs showing the mean values for the abundance of individuals (No. per m 2) from
five Hamon grab samples each of 0.25 m 2 taken in a dredged site and at a reference site. Dredging
occurred in April, and samples were taken in the pre -dredged deposits in March 1992 and through to
December 1992. Values for the biomass are expressed as g AFDW m -2 from five Hamon grab
samples. The average number of species in each of the five Hamon grab samples is also shown. The
95% confidence limits are indicated as bars. Note that there was a significant increase in species
variety and abundance during the 7-month post-dredging period, but that the biomass increased only
slowly. This indicates that recruitment was mainly of small individuals by larval settlement. Despite
this recolonization, it is clear that population density, biomass and species variety had not recovered
at the end of the 7-month post-dredging period. (After Kenny & Rees 1994).
32
The average population density for all taxa of 2769 individuals recorded by
Kenny & Rees (1994) prior to dredging was reduced after dredging to only 129 ind. m-2 ,
compared with a relatively uniform invertebrate population density of 3300 ind. m-2 in the
reference site. Again, the population density showed a significant increase in the 7 months
after dredging had ceased.
Inspection of Figure 14 shows that the high biomass of 23 g AFDW m-2 was
reduced to only lg AFDW m-2 after dredging. This reflects the removal of relatively large
macrofaunal species, such as the mussel Modiolus modiolus, from the dredged sediments
and was followed by a slower rate of increase in the post-dredging period than that
recorded for population density. This implies that recolonization was initially by small
individuals that then grew relatively slowly during the 7 months after dredging had
ceased.
Figure 15 shows the output of a non-metric multidimensional scaling (MDS)
ordination (see Kruskal 1977, Kruskal & Wish 1978, Field et al. 1982) of the data for the
macrofauna sampled in gravel deposits before dredging of the experimental site off
Norfolk, and in the 7 months after dredging (after Kenny & Rees 1994). Their multi-
variate analysis of community structure prior to dredging and in the months following
dredging shows a number of important features of the recolonization process that
Figure 15 Two -dimensional multidimensional scaling (MDS) ordination for the benthic
communities in a survey area off Norfolk in March 1992 prior to dredging, and in May, August and
December 1992. No te that dredging of the experimental area resulted in an initial impact on
community structure which differed from that in control areas and to that in the deposits prior to
dredging. In the following months community structure became more similar to that in the
undredged deposits, but was still distinct at the end of the 7-month post-dredging period. (After
Kenny & Rees 1994).
33
highlight the general principles of succession outlined in Fig. 2 (p. 8).

The first point that is clear from their results is that the community within the
dredged site prior to dredging in March 1992 formed a small “cluster” on the MDS
ordination. This indicates that the communities sampled within the experimental site were
similar to one another, and were also evidently very similar to those in the reference site
since they are close together on the MDS ordination.
The experimental area was again sampled in May 1992, 1 month after
completion of dredging. At this stage it can be seen from Fig. 15 that dredging had
resulted in two important changes in community structure. First, the communities in all
the samples from the dredged site were well separated in the MDS plot from those in
March and from those in the reference site. This implies a major change in community
composition following dredging. Secondly, the communities at each of the sampling sites
within the dredged area were different from one another. This is indicated by the fact that
they have an increased derived variance (S2) and no longer form a tight “cluster” on the
MDS ordination shown in Fig. 15 (see also Warwick & Clarke 1993). This increased
variance would be expected when some samples were taken from the dredged furrows
themselves whereas others were from areas between furrows.
One of the interesting features of this study is that it shows that much of the
initial process of colonization of the gravel deposits off the Norfolk coast was
accomplished within the following 7-month period. Inspection of Fig. 15 shows that the
community in the dredged area became more similar to those in the surrounding deposits
of the reference area and to those in the pre-dredged site, and also had a closer internal
similarity to one another (S2 reduced to 0.09) in the months following cessation of
dredging. This shows that many of the commoner species present in the dredged area in
March 1992 prior to dredging had recolonized by December 1992. The clear difference
from both the reference site and the community prior to dredging suggests, however, that
many of the rarer components of the community had not yet colonized the dredged area in
the following 7 months.
The study was then extended to include data for a 2-yr period following
dredging. These results are reported by Kenny & Rees (1996). They showed that although
recruitment of new species, especially r-selected species such as the barnacle Balanus
crenatus and the ascidian, Dendrodoa grossularia had occurred by December 1992, even
at the end of a 2-yr period both the average species abundance and biomass for the
dredged area were lower than those in the reference site.
It is also clear from their work that the community composition in the dredged area was
not restored even 2yr after dredging, Inspection of Figure 16 shows the tightly clustered
samples from the reference site and from the pre-dredged experimental site in March
1992. The marked shift in community composition and the increased variation between
samples taken in May 1992 shortly after dredging is shown, as well as data collected in
May 1993, 1 yr post-dredging and in May 1994, 2 yr after dredging. It is apparent from
Figure 16 that despite the significant recolonization that had evidently occurred within 7
months of dredging, the community in the dredged area remained distinct from that in the
reference area and from that in the deposits prior to dredging, even after 2yr. Whether this
reflects residual differences in the nature of the deposits following dredging, or the long
time period required for establishment of the rarer components of the original community
is not yet known.
34
Figure 16 Two -dimensional multidimensional scaling (MDS) ordination for the benthic
communities in a survey area off Norfolk in March 1992, and for following 2yr post-dredging. Note
that despite the increasing similarity of the community in the dredged area to those in the
surrounding sediments over the 2yr post-dredging period, recovery had not been fully accomplished
even after 2 yr. (After Kenny & Rees 1996).
The results that have been reviewed above thus show that the process of recolonization
involves two distinct phases; first, recolonization of species composition and population
density by settlement of small individuals as larvae and juveniles; secondly, a period of
growth during which the biomass approaches that in the undisturbed deposits. Inspection
of Figure 16 shows, however, that in the gravel deposits of the southern North Sea this
process had only entered its initial phase of partial restoration of community structure in
the 7-month period that followed cessation of dredging, and that full recovery may take
several years, much as would be anticipated for typical equilibrium communities on the
seabed (see Figure 4, p. 13).
The rate of recovery of biological resources
The rates of recovery of biological resources following capital and maintenance dredging,
disposal of dredged spoils and marine aggregate dredging have been widely studied in
other habitats and conform with the general principles of ecological succession shown in
Figures 2 and 3. That is, communities that inhabit fine semi-liquid and disturbed muds
35
comprise opportunistic r-selected species that have a high rate of recolonization and which
can reach high population densities within weeks or months of a catastrophic mortality.
Conversely, communities that inhabit less disturbed deposits of deeper waters or coarse
substrata have complex associations and are characterized by large slow-growing species
that are selected for maximum competitive advantage in a habitat where space is already
crowded. These large, slow-growing, K-selected equilibrium species recolonize only
slowly following disturbance and may take several (or many) years for recovery of full
species composition and biomass.
Table 5 shows the rates of recovery of the benthic fauna following dredging in
various habitats. We have included semi-liquid muds from freshwater tidal areas and have
arranged the data along a gradient of increasing environmental stability and predictability
through estuarine and coastal muds to sands and gravels and coral reef assemblages.
Inspection of the data summarized in Table 5 shows that recovery of the benthic fauna in
highly disturbed semi-liquid muds can occur within weeks. This is associated with an
ability for species such as Limnodrilus spp., Ilyodrilus, Coelotanypus sp. and Procladius
to migrate through the surrounding deposits and to recolonize disturbed muds as adults.
Table 5 Table showing the rates of recovery of the benthic fauna following dredging in various
habitats. Note that highly disturbed sediments in tidal fresh waters and estuaries that are dominated
mainly by opportunistic (r-strategist) species have a rapid rate of recovery. Recovery times increase
in stable habitats of gravels and coral reefs that are dominated by long-lived components with
complex biological interactions controlling community structure. Longevity and slow growth are
also associated with slow recolonization rates in sub-arctic seas. Examples have been arranged along
a gradient from disturbed muds of freshwater-tidal estuarine conditions to stable reef assemblages.
36
A similar recolonization of disturbed deposits in dredged channels may also account for
the relatively fast recolonization of some muds and sands in near-shore waters, especially
those where tidal currents may transport juveniles into the dredged area (see Hall 1994).
Inspection of the recolonization rates reported in the literature and summarized in
Table 5 suggest that a period of 2-4yr is a realistic estimate of the time required
for recovery in gravels and sands, but that this time may be increased to more than 5 yr in
coarser deposits, including coral reef areas. Interestingly, the data for areas in Tampa Bay,
Florida that had been dredged for oyster shell, suggest that a period of as much as 1Oyr
may be required for recovery following complete defaunation whereas a recovery time of
only 6-12 months is required for recovery following partial dredging and incomplete
defaunation (see Benefield 1976, Conner & Simon 1979). This suggests that areas of
undisturbed deposits between dredged furrows may provide an important source of
colonizing species that enable a faster recovery than might occur solely by larval
settlement and growth (see also van Moorsel 1993, 1994).
Other more complex environmental factors also evidently affect the rate of
recovery of dredged areas. Studies in the Dutch Wadden Sea by van der Veer et al. (1985)
show that the recovery of species composition and biomass of benthic organisms was
related to the speed of infilling of dredged pits. These data are summarized in Table 6,
which shows that even 16yr after cessation of dredging no recovery of the benthos had
occurred on a tidal flat at Terschelling Sand. On a tidal watershed at Oosterbierum a
partial recovery of 85% of the species and 39% of the biomass had occurred after 4 years.
This is typical of recolonization by small individuals that were in the process of growth
towards the original biomass levels of the undisturbed deposits, a process which would
clearly take several more years.
In the tidal channels, both the rate of infill and recolonization were related to the
speed of currents. A partial recovery of 57% of the species and 67% of the biomass was
recorded after 3 yr in a tidal channel at Paesensrede (see Table 6), with greater recovery
and shorter time periods being recorded in areas of faster current. Even then, it will be
noted that the species composition had not recovered and that the biomass evidently
became dominated by fewer species of relatively large size compared with those in the
surrounding deposits.
The likely recolonization rates for the benthic community of estuarine muds,
sands, gravels and reef areas have been superimposed onto a generalized colonization
succession in Figure 17, which allows some predictions to be made on the rates of
recovery of deposits following dredging. The fine muds that characterize coastal
embayments, estuaries and lagoons are likely to be colonized by large populations of a
37
relatively restricted variety of opportunistic r-selected species, which are capable of rapid
colonization within months of space being made available for colonization and growth.
Because such deposits are subject to regular disturbance under natural conditions prior to
dredging, the ecological succession recovers to the colonization phase shown in Figure 17,
but does not proceed to the development of K-selected slow-growing equilibrium species
within the community. Recovery of the “normal” community in disturbed deposits such as
muds, therefore, can be achieved within months of cessation of dredging, or disposal of
spoils.
Figure 17 Schematic diagram showing the likely recolonization rates for the benthic community
of estuarine muds, sands and reef areas. The curves for recovery have been superimposed onto a
generalized colonization succession and allows some predictions to be made on the rates of recovery
of deposits following dredging. Note that the fine muds that characterize coastal embayments,
estuaries and lagoons are likely to be recolonized by a relatively restricted variety of opportunistic r-
selected species within months of space being made available for recolonization and growth.
Because such deposits are subject to regular disturbance, the succession recovers to the colonization
phase, but does not proceed to the development of long-lived slow-growing K-selected species. The
natural communities of gravels and sands, however; contain varying proportions of slow-growing
K-selected equilibrium species depending on the d egree of disturbance by waves and currents.
These communities are held in a transitional state by natural environmental disturbance and are
likely to recover within a period of 2-3yr after cessation of dredging. Finally, the recovery curve for
reef communities indicates that a period of 8-l0yr may be required for the long process of
establishment and growth of the long-lived and slow-growing K-selected species characteristic of
equilibrium communities.
38
The natural communities of gravel and sand deposits, however, contain varying
proportions of slow-growing, K-selected equilibrium species, depending, among other
factors, on the degree of disturbance by waves and the speed of tidal currents. In this case,
the tail of the sigmoid recovery curve becomes more pronounced because the rarer
components of the equilibrium community may take several years to recolonize the
deposits, even after the main components of the community have become established.
Where the deposits are sandy, periodic mortality of the long-lived components may result
in major seasonal changes in community composition such as occurs in the North Sea on
the Klaver Bank (van Moorsel 1994) and as has been reported for the sediments of
Liverpool Bay by Eagle (1975). Under these conditions, the community will be held in a
transitional state by natural environmental disturbance, and is likely to recover within 2-3
yr after cessation of dredging.
There is good evidence that disturbance of the deposits by man may result in a
shift from the equilibrium community characteristic of undisturbed deposits towards the
transitional community, which characterizes deposits in areas of natural environmental
disturbance. Studies by de Groot (1984) suggest, for example, that the increasingly heavy
bottom gear used by trawlers has been associated with a shift in community composition
of the benthos of the North Sea, and this also applies to the benthos ofthe Wadden Sea.
As might be anticipated from the successional sequence shown in Figure 17,
long-lived components such as molluscs and larger crustaceans in near-shore waters, such
as the Wadden Sea, have decreased in numbers and diversity over the years and have been
replaced by larger populations of rapidly growing polychaete species (Reise 1982, Riesen
& Reise 1982, Reise & Schubert 1987).
Finally, the community recovery curve for reef communities indicates that a
period of 8-10yr may be required for the process of establishment and growth of the long-
lived and slow-growing K-selected equilibrium species and for the development of the
biological interactions that are familiar to those who have observed the immense diversity
and complexity of life on undisturbed reef structures. This long process of establishment
of an equilibrium community reflects partly the time required for colonization by rarer
components of the community, but is also influenced by the nature and stability of the
substratum following cessation of dredging, and the time required for complex
stabilization processes involving both physical compaction and biological interactions.
The relationship between biological community structure, sediment composition and
seabed stability is considered in more detail below.
Community composition and seabed stability
The influence of sediment composition in controlling the nature of communities of

animals that live on the sea bed has been widely recognized since the pioneer studies of
Petersen (1913), Thorson (1957) and Sanders (1958). Most recent evidence suggests,
however, that the precise relationship between biological community composition and
specific properties of the sediments is poorly understood. In some estuaries and shallow
water coastal embayments, fine grained and silty deposits clearly support an entirely
39
distinct community compared with those from mo bile sands or on stable sub-strata such as
rocks and boulders.
On the other hand it is a matter of common observation that although very fine
mobile muds may be dominated by opportunistic species such as the amphipod Ampelisca
brevicornis or the polychaete Lagis koreni, the same silts can become consolidated into
clays and then support long-lived and sedentary equilibrium species such as the boring
piddock bivalves Pholas dactylus and Barnea parva as well as an epifauna of hydroids,
ascidians and other species more characteristic of reefs. Clearly, the stability of the
sediment, rather than particle size itself, is of importance in controlling community
structure. In other instances it is clear that the deposits on the seabed undergo a complex
process of consolidation or “armouring ” that allows the establishment of communities
that are more typical of rocks and reefs reflecting the complex relationships between the
physical deposits and biological activities of the animals themselves.
The relationship between community composition and sediment type in deeper
waters of the continental shelf is less well documented than that for estuaries and lagoons.
Some early studies suggest that macrobenthic communities can be distinguished on a basis
of sediment granulometry (Gltmarec 1973, Buchanan et al. 1978, Flint 1981) but other
studies have shown little correlation (Buchanan 1963, Day et al. 1971). Efforts to identify
what physical properties are of greatest importance in controlling the structure of marine
communities are often frustrated by the fact that most of the sediment variables obtained
from conventional sorting methods are interdependent since they are expressed as a
percentage of the total sample (see Weston 1988). A high percentage of silt, for example,
is inversely related to the percentage of the other sediment components. Again, many of
the physical properties of sediments are linked with other features such as depth of
disturbance by wave action, strength and duration of currents, and may themselves be
linked with complex biological interactions including the surface area available for
microbial food components, and the presence of species that can exclude potential
competitors.
Partly for this reason, most recent studies have concluded that the complexity of
soft-bottom communities defies any simple paradigm relating to a single factor, and that
there should be a shift towards understanding relationships between the distribution of
organisms in terms of a dynamic relationship between the sediments and their
hydrodynamic environment. According to this view, complex shear forces at the
sediment-water interface are considered to play a dominant role in controlling food
availability, settlement of larvae, microbial food availability, pore water flow and other
environmental features that affect the benthic organisms that inhabit marine deposits. It is
therefore considered unlikely that any one factor alone, or even a combination of single
granulometric properties, can account for the distribution of animals in most sedimentary
habitats (for review, see Snelgrove & Butman 1994).
Despite this emerging view that sediment granulometry itself is unlikely to
control the composition and distribution of biological communities on the sea bed,
concern has been expressed that dredging for marine aggregates can result in significant
changes in sediment composition. Studies off Dieppe, France have shown, for example, a
large increase in the proportion of fine sand in deposits that have been intensively worked
for marine aggregates (see Desprez 1992, ICES 1992, 1993). Again, the infill of pits and
grooves from dredging for marine aggregates is commonly dominated by the fine deposits
which are capable of mobilization by shear stress induced by waves and tidal currents
(Dickson & Lee 1972, Shelton & Rolfe 1972, Millner et al. 1977).
40
If sediment composition were of importance in controlling biological community

composition, such changes following dredging could potentially prevent subsequent
recolonization by communities that were similar to those that occurred in the deposits
prior to dredging (see Windom 1976) and could by implication affect the nature and
abundance of food organisms for commercial fish stocks.
We have analyzed the relationship between biological community composition
and the sediment granulometry in undredged coastal deposits in the English Channel and
southern North Sea and find that both biological communities and the sediments fall into
relatively distinct groups or communities when analyzed by multivariate techniques
(Newell & Seiderer 1997d). However, there is little evidence of any correspondence
between the distribution of different sediment types and biological communities in the
survey areas. Analysis of the Spearman rank correlation between the similarity of
biological communities and any one, or a combination of, particle size indices show that
granulometric properties of the sediments are likely to account for a maximum of 45% of
the variability of the biological component, leaving approximately 55% determined by
other environmental factors.
The conclusion to be drawn from these results is that they support recent views
that biological community composition is not controlled by any one, or a combination of
simple granulometric properties of the sediments such as particle size distribution. It is
considered more likely that biological community composition is controlled by an array of
environmental variables, many of them reflecting an interaction between particle mobility
at the sediment-water interface and complex associations of chemical and biological
factors operating over long periods.
Such interactions are not easily measured or analyzed, but the results clearly
suggest that restoration of sediment composition after completion of dredging for marine
aggregates is not, within broad limits, a prerequisite for the establishment of marine
communities that are comparable with those that occurred in the deposits prior to
dredging. What is possibly of more importance in controlling the time course of recovery
of an equilibrium community characteristic of undisturbed deposits is the process of
compaction and stabilization. This will reflect changes in sediment composition, but is
also in equilibrium with seabed disturbance from tidal currents and wave action, both of
which show spatial variations and interactions with water depth. The processes associated
with compaction and stability of seabed deposits may therefore largely control the
establishment of long-lived components of equilibrium communities and account for the
dominance of opportunistic species in the initial stages of colonization of recently
sedimented material in unconsolidated deposits after the cessation of dredging.
Conclusions
At the outset of this review, we assessed the importance of the benthic community to
fisheries production and outlined our intention of providing an ecological framework
within which the impact of dredging can be understood. We have shown that systems
models for shelf waters such as the North Sea suggest that the flow of materials from
primary production by the phytoplankton passes partly through planktonic grazers, but
that 20-50% sinks to the sea-bed either from dead and decaying phytoplankton cells, or as
41
faecal material derived from the feeding activities of the grazing zooplankton (Steele
1974, Joiris et al. 1982, Newell et al. 1988). Such material then passes into the benthic
food web, whose production in turn forms an important food resource for demersal fish.
It has been estimated from empirical models developed for the North Sea that as
much as 30% of total fisheries yield to man is derived from benthic resources (see Fig. 1,
p. 3). Production by the benthos is therefore important, not only as a resource in itself, but
as a key food resource for demersal fish stocks. It becomes an increasingly important
component of the marine food web in near-shore waters where primary production by
larger macrophytes and seagrasses living on the seabed largely replaces that from the
phytoplankton in the water column (for review, see Mann 1982).
From this it is clear that reclamation of large areas of coastal wetlands, coastal
embayments or estuaries can have a potentially important effect on the supply of materials
and energy to marine food webs, and that even in plankton-based deeper water ecosystems
such as the North Sea, fish yields based on benthic production are sufficiently large to
warrant proper conservation of benthic resources. Our review has concentrated, therefore,
on the nature of benthic communities, their susceptibility to disturbance by dredging and
land reclamation works, and on the evidence that is available for the recovery times
required for the re-establishment of community structure following dredging or spoils
disposal.
Our review of the literature shows that the communities of near-shore habitats
are characterized by large populations of a relatively restricted variety of species that are
well-adapted to exploit space that has become newly available by episodic catastrophic
mortality. Such species are generally small, often mobile, and are selected for maximum
rate of population increase, with high fecundity, dense settlement, rapid growth and rather
a short life cycle. Such species have been designated “r-strategists” (see MacArthur &
Wilson 1967, Pianka 1970) and have been referred to in our review as opportunists. Their
population characteristics allow a rapid recovery of the initial community structure in
deposits that are naturally subjected to high levels of environmental disturbance. It is, not
surprising, therefore, to find that there are frequent reports in the literature of community
recovery times that range from a few weeks to several months for disturbed deposits such
as semi-liquid muds in tidal fresh waters, estuaries, lagoons and dredged channels (see
Table 5, p. 36 and Fig. 17, p. 38).
In deeper waters, or where the substratum is sufficiently stable to allow the long-
term survival of benthic organisms, the habitat tends to be crowded. Under these
conditions, organisms have an “equilibrium strategy” and are selected for maximum
competitive ability in an environment in which space for colonization and subsequent
growth is limiting. Such species have been designated “K-strategists” and devote a larger
proportion of their resources to non-reproductive processes such as growth, predator
avoidance, and investment in larger adults (MacArthur & Wilson 1967, Gadgil & Bossert
1970). Because the K-selected equilibrium species live longer, they tend to have wider
limits of physiological tolerance, which allows them to survive those variations in
environmental conditions that occur in their habitat. Many have active site selection
phases that include chemical recognition of the presence of adults of the same species. a
strategy which ensures that environmental conditions have been within the limits of
tolerance for long enough to allow survival of other members of the same species (for
review, see Newell 1979).
Such K-selected equilibrium species develop complex biological associations
with other long-lived components of the community, and may alter the environment in
such a way as to both allow the presence of many other species that would not otherwise
42
occur, and also inhibit other potential competitors for space. Biological interactions
between the components of equilibrium communities that are characteristic of stable
substrata thus lead to the development of complex communities that may take many years,
or decades, to re-establish following destruction. It is therefore not surprising to find that
as one moves along a gradient of increasing sediment stability from muds through sands
to gravels and reefs, there is a corresponding increase in the times reported for recovery of
community structure (Table 5, p. 36).
Knowledge of the components that comprise the benthic community on the sea
bed, whether these are r-selected opportunistic species or K-selected equilibrium species,
thus gives important information not only on key resources that may require protection,
but on the likely rate of recovery following dredging. Inspection of the schematic
colonization succession shown in Figure 17 (p. 38) suggests that a recovery time of 6-8
months is characteristic for many estuarine muds whereas sands and gravels may take
from 2-3yr depending on the proportion of sand and the local disturbance by waves and
currents. As the deposits become coarser, estimates of 5-10yr are probably realistic for the
development of the complex biological associations between the slow-growing
components of equilibrium communities characteristic of reef structures.
Our review suggests that processes associated with compaction and stabilization
of seabed deposits may largely control the time-course of recovery of these long-lived
components of equilibrium communities and account for the dominance of opportunistic
species in the initial stages of colonization of recently sedimented material in
unconsolidated deposits following the cessation of dredging.
Acknowledgements
This work was partly supported through a co-operative research initiative funded by the
Minerals Management Service, US Department of the Interior, Washington DC (Contract
# 14-35-001-30763 to D. R. Hitchcock) and supported by the following organisations:
Messrs ARC Marine Ltd, United Marine Dredging Ltd, South Coast Shipping Ltd, Civil
& Marine Ltd and H. R. Wallingford Ltd.
We are grateful for the helpful assistance with the literature given by Professor S.
E. Shumway of the University of Long Island, NY, Professor R. I. E. Newell of the
University of Maryland and Professor Derek V. Ellis of the University of Victoria,
Canada. We also acknowledge with thanks the advice and comments of Mr Tom
Matthewson of Oakwood Environmental, Wormley, Surrey, Mr A. Hermiston of ARC
Marine, Southampton, Dr A. J. Kenny and Dr H. L. Rees of the Centre for Environment,
Fisheries & Aquaculture Science (CEFAS), Burnham-on-Crouch, Essex. Messrs South
Coast Shipping and Oakwood Environmental kindly gave permission for use of
unpublished data obtained in baseline surveys carried out by us on their behalf off the
coasts of Kent and East Anglia during 1996.
43
References
Aller, R. C. & Dodge, R. E. 1974. Animal-sediment relations in a tropical lagoon. Journal of Marine Research
32(2), 209-32.
Aschan, P. 1988. The effect of Iceland scallop (Chlamys islandica) dredging at Jan Mayen and in the
Spitsbergen area. ICES CM 1988, No. 16.
Bak, R. P. M. 1978. Lethal and sublethal effects of dredging on reef corals. Marine Pollution Bulletion 9, 14-16.
Basimi, R. A. & Grove, D. J. 1985. Estimates of daily food intake by an inshore population of Pleuronectes
platessa L. off eastern Anglesey, North Wales. Journal of Fish Biology 27, 505-20.
Benefield, R. L. 1976. Shell dredging sedimentation in Galveston and San Antonio Bays 1964-69. Technical
Series, Texas Parks and Wildlife Department 19, l-34.
Biggs, R. B. 1968. Environmental effects of overboard spoils disposal. Journal of Sanitary Engineering Division
94, No. SA3, Proc. Paper 5979, June 1968. Contr. No. 328,477-87.
Boesch, D. F. & Rosenberg, R. 1981. Response t o stress in marine benthic communities. In Stress effects on
natural ecosystems, G. W. Barrett & R. Rosenberg (eds). Chichester: John Wiley & Sons, 179-98.
Bonsdorff, E. 1983. Recovery potential of macrozoobenthos from dredging in shallow brackish waters.
Fluctuation and succession in marine ecosystems. Proceedings of the 17th European Marine Biology
Symposium, Brest : France, Oceanologica Acta, 27-32.
Brenchley, G. A. 1981. Disturbance and community structure: an experimental study of bioturbation in marine
soft -bottom environments. Journal of Marine Research 39,767-90.
Brey, T. 1991. The relative significance of biological and physical disturbance: an example from intertidal and
subtidal sandy bottom communities. Estuarine Coastal and Shelf Science 33, 339-60.
Bright, D. A. & Ellis, D. V. 1989. Aspects of histology in Macoma carlottensis (Bivalvia: Tellinidae) and in situ
histopathology related to mine-tailings discharge. Journal of the Marine Biological Association of the
United Kingdom 69,447-64.
Brown, B. E., Le Tissier, M. D. A., Scoffin, T. P. & Tudhope, A. W. 1990. Evaluation of the environmental
impact of dredging on intertidal coral reefs at Ko Phuket, Thailand, using ecological and physiological
parameters. Marine Ecology Progress Series 65, 273-81.
Buchanan, J. B. 1963. The bottom fauna communities and their sediment relationships off the coast of
Northumberland. Oikos 14, 154-75.
Buchanan, J. B., Sheader, M. & Kingston, P. F. 1978. Sources of variability in the benthic macrofauna off the
south Nort humberland coast. 1971-1976. Journal of Marine Biological Association of the United Kingdom
58,191-209.
Buss, L. B. & Jackson, J. B. C. 1981. Planktonic food availability and suspension-feeder abundance: evidence of
in situ depletion. Journal of Experimental Marine Biology and Ecology 49, 151-61.
Carney, R. S. 1981. Bioturbation and biodeposition. In Principles of Benthic Marine Paleoecology, A. J. Boucot
(ed.). New York: Academic Press, 357-99.
Carter, C. G., Grove, D. J. & Carter, D. M. 1991. Trophic reso urce partitioning between two co-existing flatfish
species off the North coast of Anglesey, North Wales. Netherlands Journal of Sea Research 27,
325-35.
C-CORE. 1996. Proposed mining technologies and mitigation techniques: a detailed analysis with respect to the
mining of specific offshore mineral commodities. Contract Report for U.S Department of the Interior,
Minerals Management Service. C-CORE Publication 96-C15.
Charlier, R. H. & Charlier, C. C. 1992. Environmental, economic and social aspects of marine aggregates
exploitation. Environmental Conservation 19,29-37.
Clarke, D. & Miller-Way, T. 1992. An environmental assessment of the effects of open-water disposal of
maintenance dredged material on benthic resources in Mobile Bay, Alabama. National Technical
Information Service, Springfield, Virginia 22 161, USA.
Clarke, D. G., Homziak, J., Lazor, R., Palermo, M. R., Banks, G. E., Benson, H. A., Johnson, B. H., Smith-
Dozier, T., Revelas, G. & Dardeau, M. R. 1990. Engineering design and environmental assessm ent of
dredged material overflow from hydraulically filled hopper barges in Mobile Bay, Alabama. Miscellaneous
Paper D-90-4. Vicksburg, MS. US Army Engineer Waterways Experiment Station.
Clarke, K. R. & Ainsworth, M. 1993. A method of linking multivariate community structure to environmental
variables. Marine Ecology Progress Series 92,205-19.
Clarke, K. R. & Warwick, R. M. 1994. Change in Marine Communities: an approach to statistical analysis and
interpretation. Natural Environment Research Council.
Cloern, J. E. 1982. Does the benthos control phytoplankton biomass in South San Francisco Bay? Marine
Ecology Progress Series 9,191-202.
Connell, J. H. 1978. Diversity in tropical rain forests and coral reefs. Science 199, 1302-10.
44
Conner, W. G. & Simon, J. L. 1979. The effects of oyster shell dredging on an estuarine benthic community.
Estuarine and Coastal Marine Science 9,749-58.
Courtenay, W. R., Hartig, B. C. & Loisel, G. R. 1972. Ecological monitoring of two beach nourishment projects
in Broward County, Florida. Shore and Beach 40(2), 8-13.
Cruikshank, M. J. & Hess, H. D. 1975. Marine sand and gravel mining. Oceanus 19,32-44.
Dame, R. F. 1993. The role of bivalve filter feeder material fluxes in estuarine ecosystems. NATO ASI Series G
33, Bivalve Filter Feeders; In Estuarine and coastal ecosystem processes, R. F. Dame (ed.). Heidelberg
Springer-Verlag, 246-68.
Davies, C. M. & Hitchcock, D. R. 1992. Improving the exploitation of marine aggregates by the study of the
impact of marine mining equipment. Unpublished Report Ref. GR/G 20059. Marine Technology
Directorate UK.
Davoult, D., Dewarumez, J. M., Prygiel, J. & Richard, A. 1988. Benthic communities map of the French part of
the North Sea. Station Marin de Wimereux URA-CNRS. 1363.
Dawson-Shepherd, A., Warwick, R. M., Clarke, K. R. & Brown, B. E. 1992. An analysis of fish community
responses to coral mining in the Maldives. Environmental Biology of Fishes 33,367-80.
Day, J. H., Field, J. G. & Montgomery, M. P. 1971. The use of numerical methods to determine the distribution
of the benthic fauna across the continental shelf of North Carolina. Journal of Animal Ecology 40, 93-126.
de Groot, S. J. 1979. An assessment of the potential environmental impact of large -scale sand-dredging for the
building of artificial islands in the North Sea. Ocean Management 5,211-32.
de Groot, S. J. 1984. The impact of bottom trawling on benthic fauna of the North Sea. Ocean Managment 9,
177-90.
de Groot, S. J. 1986. Marine sand and gravel extraction in the North Atlantic and its potential environmental
impact, with emphasis on the North Sea. Ocean Management 10,21-36.
Desprez, M. 1992. Bilan de dix annees de suivi de l’impact biosedimentaire de l’extraction de graves marins au
large de Dieppe. Comparaison avec d’autres sites. Rapport Groupe d’tude des Milieux Estuariens et
Littoraux GEMEL. St Valery/Somme. (Cited in Report of the working group on the effects of extraction of
marine sediments on fisheries. ICES Report No. CM 1993/E:7 Marine Environmental Quality Committee,
51-67).
de Witt, T. H. & Levinton, J. S. 1985. Disturbance, emigration and refugia: how the mud snail Ilyanassa
obsoleta (Say) affects the habitat distribution of an epifaunal amphipod, Microdeutopus gryllotalpa
(Costa). Journal of Experimental Marine Biology and Ecology 92,97-111.
Diaz, R. J. 1994. Response of tidal freshwater macrobenthos to sediment disturbance. Hydrobiologia 278, 201-
12.
Dickson, R. & Lee, A. 1972. Study of the effects of marine gravel extraction on the topography of the sea bed.
ICES, C.M. 1972/E: 25.
Dodge, R. E. & Brass, G. W. 1984. Skeletal extension, density and calcification of a reef coral (Montastrea
annularis): St Croix, US Virgin Islands. Bulletin of Marine Science 34,288-307.
Dodge, R. E. & Vaisnys, J. R. 1977. Coral populations and growth patterns : responses to sedimentation and
turbidity associated with dredging. Journal of Marine Research 35,715-30.
Driscoll, E. G. 1975. Sediment-animal-water interaction, Buzzards Bay, M a s s a c h u s e t t s . Journal Marine
Research 33,275-302.
Eagle, R. A. 1975. Natural fluctuations in a soft bottom community. Journal of the Marine Biological
Association of the United Kingdom 55, 865-78.
Eden, R. A. 1975. North Sea environmental geology in relation to pipelines and structures. Oceanology
International 75, 302-9.
Eleftheriou, A. & Basford, D. J. 1989. The macrobenthic infauna of the offshore northern North Sea. Journal of
the Marine Biological Association of the United Kingdom 69, 123-43.
Ellis, D. V. 1987. A decade of environmental impact assessment at marine and coastal mines. Marine Mining
6,385417.
Ellis, D. V. & Heim, C. 1985. Submersible surveys of benthos near a turbidity cloud. Marine Pollution Bulletin
16(5), 197-203.
Ellis, D. V. & Hoover, P. M. 1990. Benthos recolonising mine tailings in British Columbia fiords. Marine
Mining 9,441-57.
Ellis, D. V. & Jones, A. A. 1980. The Ponar grab as a marine pollution sampler. Canadian Research June/ July
1980,23-25.
Ellis, D. V., Pedersen, T. F., Poling, G. W., Pelletier, C. & Horne, I. 1995. Review of 23 years of STD: Island
Copper Mine, Canada. Marine Georesources and Technology, 13(1-2), 59-99.
Ellis, D. V. & Taylor, L. A. 1988. Biological engineering of marine tailings beds. In Environmental Management
of Solid Wastes. W. Salomons & V. Forstner (eds). Heidelberg: Springer-Verlag.
Field, J. G., Clarke, K. R. & Warwick, R. M. 1982. A practical strategy for analyzing multispecies distribution
patterns. Marine Ecology Progress Series 8, 37-52.
45
Field, J. G. & MacFarlane, G. 1968. Numerical methods in marine ecology. I. A quantitative similarity analysis
of rocky shore samples in False Bay, South Africa. Zoologica Africana 3, 119-38.
Flach, E. C. 1992. Disturbance of benthic infauna by sediment reworking activities of the lugworm, Arenicola
marina. Netherlands Journal of Sea Research 30,81-9.
Flint, R. W. 1981. Gulf of Mexico outer continental shelf benthos. macrofaunal- environmental relationships.
Biological Oceanography 1, 135-55.
Flint, R. W. & Kalke, R. D. 1986. Biological enhancement of estuarine benthic community structure. Marine
Ecology Progress Series 31,23-33.
Forster, G. R. 1953. A new dredge for collecting burrowing animals. Journal of the Marine Biological
Frechette, M, Butman, C. A. & Geyer, W. R. 1989. The importance of the boundary layer flows in supplying
phytoplankton to the benthic suspension-feeder, Mytilus edulis L. Limnology and Oceanography 34,19-36.
Frechette, M., Lefaivre, D. & Butman, C. A. 1993. Bivalve feeding and the benthic boundary layer. NATO ASI
SERIES G 33. Bivalve Filter Feeders In Estuarine and Coastal Ecosystem Processes, R. F. Dame (ed.).
Heidelberg: Springer-Verlag, 325-64.
Gadgil, M & Bossert, W. H. 1970. Life historical consequences of natural selection. American Naturalist 104, l-
24.
Gadgil, M & Solbrig, 0. T. 1972. The concept of r- and K-selection: evidence from wild flowers and some
theoretical considerations. American Naturalist 106, 14-31.
Gajewski, L. S. & Uscinowicz, S. 1993. Hydrologic and sedimentologic aspects of mining marine aggregate
from the Slupsk Bank (Baltic Sea). Marine Georesources and Geotechnology 11,229-44.
Garnett, R. H. T & Ellis, D. V. 1995. Tailings disposal at a marine placer mining operation by West Gold,
Alaska. Marine Georesources and Geotechnology, 13(1-2), 41-57.
Gayman, W. 1978. Offshore dredging study: environmental ecological report. Ocean Management 4,51-104.
Giesen, W. B. J. T. van Katwijk, M. M. & den Hartog, C. 1990. Eelgrass condition and turbidity in the Dutch
Wadden Sea. Aquatic Botany 37,71-85.
Glasby, G. P. 1986. Marine minerals in the Pacific. Oceanography and Marine Biology: an Annual Review 24, 1
l-64.
Glemarec, M. 1973. The benthic communities of the European North Atlantic continental shelf. Oceanography
and Marine Biology: an Annual Review 11,263-89.
Glynn, P. W. 1973. Ecology of a Caribbean coral reef. The Porites flat biotope. Part II. Plankton community
with evidence for depletion. Marine Biology 22, 1-21.
Grassle, J. F. & Grassle, J. P. 1974. Opportunistic life histories and genetic systems in marine benthic
polychaetes. Journal of Marine Research 32,253-84.
Grassle, J. F. & Sanders, H. L. 1973. Life histories and the role of disturbance. Deep-Sea Research 20, 643-59.
Gray, J. S. 1974. Animal-sediment relationships. Oceanography and Marine Biology: an Annual Review 12,
223-61.
Gray, J. S. & Pearson, T. H. 1982. Objective selection of sensitive species indicative of pollution-induced change
in benthic communities. I. Comparative methodology. Marine Ecology Progress Series 9, 111-19.
Gruet, Y. 1986. Spatio -temporal changes of sabellarian reefs built by the sedentary polychaete Sabellaria
alveolata (Linne). P.S.Z.N.Z. Marine Ecology, 7, 303-19.
Guillou, M. & Hily, C. 1983. Dynamics and biological cycle of a Melinna palmata (Ampharetidae) population
during the recolonisation of a dredged area in the vicinity of the harbour of Brest (France). Marine Biology
73,43350.
Hall, S. J. 1994. Physical disturbance and marine communities: life in unconsolidated sediments. Oceanography
and Marine Biology: an Annual Review 32, 1 7 9 -2 3 9 .
Heip, C., Harman, P. M. J. & Soetaert, K. 1988. Data processing, evaluation and analysis. In Introduction to the
study of meiofauna, R. P. Higgins & H. Thiel (eds). Washington, DC: Smithsonian Institution, 197-231.
Hess, H. D. 1971. Marine sand and gravel mining industry in the United Kingdom. NOAA Technical Report
ERL 213-MMTCI. US Department of Commerce, Boulder, Colorado.
Hill, M. 0. 1979. DECORANA- A FORTRAN Program for Detrended Correspondence Analysis and Reciprocal
Averaging. Ithaca, Cornell University.
Hily, C. 1983. Macrozoobenthic recolonisation after dredging in a sandy mud area of the Bay of Brest enriched
by organic matter. In Fluctuation and Succession in Marine Ecosystems. Proceedings of the 17th European
Marine Biology Symposium, Brest. France. Oceanologica Acta 113-20.
Hitchcock, D. R. & Dearnaley, M. P. 1995. Investigation of benthic and surface plumes associated with marine
aggregate production in the United Kingdom: overview of year one. Proceedings of the XVth Information
Transfer Meeting, Gulf Coast Region. INTERMAR, New Orleans, USA.
Hitchcock, D. R. & Drucker, B. R. 1996. Investigation of benthic and surface plumes associated with marine
aggregates mining in the United Kingdom. In The global ocean - towards operational oceanography. Pro-
46
ceedings of Conference on Oceanology International. Spearhead Publications, Surrey Conference Pro-

ceedings 2,221-84.
Hodgson, G. 1994. The environmental impact of marine dredging in Hong Kong. Coastal Management of
Tropical Asia 2, l-8.
Holme, N. A. 1966. The bottom fauna of the English Channel. Part II. Journal of the Marine Biological
Holme, N. A. & McIntyre, A. D. (eds). 1984. Methods for the Study of Marine Benthos. International Biological
Programme Handbook. 2nd edn Oxford: Blackwell.
Holt, T. J., Jones, D. R., Hawkins, S. J. & Hartnoll, R. G. 1995. The sensitivity of marine communities to man-
induced change - a scoping report. Countryside Council for Wales. Contract No. FC 73-01-96. Port Erin
Marine Laboratory, Centre for Coastal Studies, University of Liverpool. Port Erin, Isle of Man UK.
Horwood, J. 1993. The Bristol channel sole (Solea solea (L): a fisheries case study. Advances in Marine Biology
29,215367.
Howell, B. R. & Shelton, R. G. J. 1970. The effect of china clay on the bottom fauna of St Austell and
Mevagissey Bays. Journal of the Marine Biological Association of the United Kingdom 50, 593-607.
Hurme, A. K. & Pullen, E. J. 1988. Biological effects of marine sand mining and fill placement for beach
replenishment: lessons for other uses. Marine Mining, 7, 123-36.
Huston, M. 1994. Biological diversity: the coexistence of species on changing landscapes. Cambridge: Cam-
bridge University Press.
ICES 1975. Report of the working group on effects on fisheries of marine sand and gravel extraction. Co-
operative Research Report No. 46.
ICES 1977. Second report of the ZCES working group on effects on fisheries of marine sand and gravel
extraction. Co-operative Research Report No. 64.
ICES 1992a. Report of the working group on the effects of extraction of marine sediments on fisheries. Marine
Environmental Quality Committee Report Ref C.M. 1992/E:7.
ICES 1992b. Report of the working group on the effects of extractio n of marine sediments on fisheries. Co-
operative Research Report No. 182.
ICES 1993. Report of the working group on the effects of extraction of marine sediments on fisheries. Marine
Environmental Quality Committee Report, Ref. CM 1993/E:7.
Iglesias, J. I. P., Urrutia, M. B., Navarro, E., Alvarez-Jorna, P., Larretxea, X., Bougrier, S. & Heral, M. 1996.
Variability of feeding processes in the cockle, Cerastoderma edule (L.) in response to changes in seston
concentration and composition. Journal of Experimental Marine Biology and Ecology 197, 121-43.
Ingle, R. M. 1952. Studies on the effect of dredging operations on fish and shellfish. State of Florida Board of
Conservation Technical Series No. 5. l-25.
Johanson, E. E. & Boehmer, W. R. 1975. Examination of predictive models for dredged material dispersion in
open water. In 1st International Symposium on Dredging Technology Canterbury: University of Kent.
BHRA Fluid Engineering, Bedford, Session D3, 33-46.
Johnson, R. G. 1970. Variations in diversity within benthic marine communities. American Naturalist 104, 285-
300.
Johnson, R. G. 1977. Vertical variation in particulate matter in the upper twenty centimetres of marine
sediments. Journal of Marine Research 35,273-82.
Johnston, S. A. 1981. Estuarine dredge and fill activities: a review of impacts. Environmental Management 5,
427-40.
Joiris, C., Billen, G., Lancelot, C., Daro, M. H., Mommaerts, J. P., Bertels, A., Bosicart, M., Nijs, J. & Hecq, J.
H. 1982. A budget of carbon cycling in the Belgian coastal zone: relative roles of zooplankton,
bacterioplankton and benthos in the utilisation of primary production. Netherlands Journal of Sea Research
16, 260-75.
Jokiel, P. L. 1989. Effects of marine mining dredge spoils on eggs and larvae of a commercially important
species of fish, the Mahimahi (Coryphaena hippurus). Marine Mining 8, 303-15.
Jones, A. A. & Ellis, D. V. 1976. Sub-Obliterative effects of mine-tailing on marine infaunal benthos. II. Effect
on the productivity of Ammotrypane aulogaster. Water, Air and Soil Pollution 5,299-307.
Jones, G. & Candy, S. 1981. Effects of dredging on the macrobenthic infauna of Botany Bay. Australian Journal
of Marine and Freshwater Research 32,379-99.
Kaplan, E. H., Welker, J. R., Kraus, M. G. & McCourt, S. 1975. Some factors affecting the colonisation of a
dredged channel. Marine Biology 32, 193-204.
Kenny, A. J. & Rees, H. L. 1994. The effects of marine gravel extraction on the macrobenthos: Early post -
dredging recolonisation. Marine Pollution Bulletin 28(7), 4 4 2 -7 .
Kenny, A. J. & Rees, H. L. 1996. The effects of marine gravel extraction on the macrobenthos: Results 2 years
post -dredging. Marine Pollution Bulletin 32(8/9), 615-22.
47
Kenny, A. J., Rees, H. L. & Lees, R. G. 1991. An inter-regional comparison of gravel assemblages off the
English east and south -east coasts: preliminary results. International Council for the Exploration of the
Sea (ICES) Report CM 1991/E:27.
Kruskal, J. B. 1977. Multidimensional scaling and other methods of discovering structure. In Mathematical
methods for digital computers. Vol. 3. Statistical methods for digital computers, K. Enstein et al. (eds).
New York: Wiley-Interscience, 296-339.
Kruskal, J. B. & Wish, M. 1978. Multidimensional scaling. Beverley Hills, CA: Sage Publications.
Kukert, H. 1991. In situ experiments on the response of deep sea macrofauna to burial disturbance. Pacific
Science 45, 95 only.
Lambshead, P. J. D., Platt, H. M. & Shaw, K. M. 1983. The detection of differences among assemblages of
marine benthic species based on an assessment of dominance and diversity. Journal of National History 17,
859-74.
Land, J., Kirby, R & Massey, J. B. 1994. Recent innovations in the combined use of acoustic doppler current
profilers and profiling silt meters for suspended solids monitoring. In Proceedings of 4th Nearshore and
Estuarine Cohesive Sediment Transport Conference. Hydraulic Research Wallingford, UK.
Lart, W. J. 1991. Aggregate dredging;fisheries perspectives. Sea Fish Industry Authority: Seafish Technology.
Seafish Report No. 404.
Lee, G. F. & Swartz, R. C. 1980. Biological processes affecting the distribution of pollutants in marine
sediments. Part II. Biodeposition and bioturbation. In Contaminants and Sediments, R. A. Baker (ed.). Ann
Arbor: Michigan, 555-605.
Lees, R. G., Kenny, A. & Pearson, R. 1992. The condition of benthic fauna in suction trailer dredger outwash:
initial findings. pp 2 2 -2 8 In Report of the Working Group on the Effects of Extraction of Marine
Sediments on Fisheries. International Council for the Exploration of the Sea. CM 1992/E:7.
Lockwood, S. J. 1980. The daily food intake of O-group plaice (Pleuronectes platessa L.) under natural
conditions. Journal du Conseil, Conseil International pour I’Exploration de la Mer. 39,745-69.
Loosanoff, V. L. 1962. Effects of tur bidity on some larval and adult bivalves. Proceedings of the Gulf and
Caribbean Fisheries Institute, Nov. 1961 80-94. Institute of Marine Science, University of Miami.
MacArthur, R. H. 1960. On the relative abundance of species. American Naturalist 94,25-36.
MacArthur, R. H. & Wilson, E. 0. 1967. Theory of island biogeography. Princeton NJ: Princeton University
Press.
Madany, I. M., Ah, S. M. & Akhter, M. S. 1987. The impact of dredging and reclamation in Bahrain. Journal of
Shoreline Management 3,255-68.
Magurran, A. E. 1991. Ecological diversity and its measurement, London: Chapman & Hall.
Mann, K. H. 1982. Ecology of coastal waters: a systems approach. Studies in Ecology Vol. 8. Oxford:
Blackwell Scientific Publications.
Maragos, J. E. 1979. Environmental surveys five years after offshore marine sand mining operations at Keauhou
Bay, Hawaii. International Report of US Army Engineer Division, Pacific Ocean, Fort Shafter, HI.
Maragos, J. E. 1991. Impact of coastal construction on coral reefs in the US-affiliated Pacific islands. Coastal
Management 21,235-69.
Marszalek, D. S. 1981. Impact of dredging on a subtropical reef community, Southeast Florida, USA. Pro-
ceedings of the 4th International Coral Reef Symposium Manila 1 , 147-53.
Matsumoto, W. M. 1984. Potential impact of deep seabed mining on the larvae of tunas and billfishes. US Dept
of Commerce, NOAA Technical Memorandum NMFS, NOAA-T M-NMFS- SWFC-44.
Maurer, D. L., Leathem, W., Kinner, P. & Tinsman, J. 1979. Seasonal fluctuations in coastal benthic invertebrate
assemblages. Estuarine and Coastal Marine Science 8, 181-93.
McCall, P. L. 1976. Community patterns and adaptive strategies of the infaunal benthos of Long Island Sound.
Journal of Marine Research 35, 221-66.
McCauley, J. E., Parr, R. A. & Hancock, D. R. 1977. Benthic infauna and maintenance dredging: a case study.
Water Research 11,233-42.
Millner, R. S., Dickson, R. R. & Rolfe, M. S. 1977. Physical and biological studies of a dredging ground off the
east coast of England. ICES C.M. 1977/E: 48.
Ministry of Agriculture, Fisheries and Food. 1993. Aquatic Monitoring Report No. 36. Lowestoft: Directorate of
Fisheries Research.
Moloney, C. L., Bergh, M. O., Field, J. G. & Newell, R. C. 1986. The effect of sedimentation and microbial
nitrogen regeneration in a plankton community: a simulation investigation. Journal of Plankton Research
8,427-45.
Moore, P. G. 1977. Inorganic particulate suspensions in the sea and their effects on marine animals.
Oceanography and Marine Biology: an Annual Review, 15,225-363.
Morton, B. 1996. The subsidiary impacts of dredging (and trawling) on a subtidal benthic molluscan community
in the southern waters of Hong Kong. Marine Pollution Bulletin 32(10), 701-10.
48
Morton, J. W. 1977. Ecological effects of dredging and dredge spill disposal: a literature review. US Fish and
Wildlife Service, Technical Papers, No. 94, l-33.
Myers, A. C. 1977a. Sediment processing in a marine subtidal sandy bottom community. I. Physical aspects.
Journal of Marine Research 35, 609-32.
Myers, A. C. 1977b. Sediment processing in a marine subtidal sandy bottom community. II. Biological
consequences. Journal of Marine Research 35, 633347.
Navarro, E., Iglesias, J. I. P., Camacho, P. A., & Labarta, U. 1996. The effect of diets of phytoplankton and
suspended bottom material on feeding and absorption of raft mussels (Mytilus galloprovincialis Lmk).
Journal of Experimental Marine Biology and Ecology 198, 175-89.
Newell, C. R. & Shumway, S. E. 1993. Grazing of natural particulates by bivalve molluscs: a spatial and
temporal perspective. NATO ASI Series G 33, In Estuarine and Coastal ecosystem processes, R. F. Dame
(ed.). Heidelberg: Springer-Verlag, 85-148.
Newell, C. R., Shumway, S. E., Cucci, T. L. & Selvin, R. 1989. The effects of natural seston particle size and
type on feeding rates, feeding selectivity and food resource availability for the mussel, Mytilus edulis Lin-
naeus, 1758 at bottom culture sites in Maine. Journal of Shellfish Research 8, 187-96.
Newell, R. C. 1979. Biology of intertidal animals. Faversham, Kent: Marine Ecological Surveys.
Newell, R. C., Moloney, C. L., Field, J. G., Lucas, M. I. & Probyn, T. A. 1988. Nitrogen models at the
community level : plant-animal-microbe interactions. In Nitrogen cycling in coastal marine environments,
T. H. Blackburn & J. Sorensen (eds). Chichester: SCOPE Publications, John Wiley, 3799414.
Newell, R. C. & Seiderer, L. J. 1997a. Benthic ecology off Lowestoft: Dredging Application Area 454. Report
prepared for Oakwood Environmental by Marine Ecological Surveys. Ref SCS/454/1.
Newell, R.C . & Seiderer, L. J. 1997b. Benthic ecology off Suffolk (Shipwash Gabbard): Dredging Application
Area 452. Report prepared for Oakwood Environmental by Marine Ecological Surveys. Ref SCS/452/1.
Newell, R. C. & Seiderer, L. J. 1997c. Benthic ecology of West Varne Licensed Dredging Areas 432/l & 432/2.
Report prepared for Oakwood Environmental by Marine Ecological Surveys. Report Ref SCS/432/1.
Newell, R. C. & Seiderer, L. J. 1997d. Influence of sediment granulometry on benthic community composition.
Report prepared for Oakwood Environmental by Marine Ecological Surveys.
Nichols, F. H. 1974. Sediment turnover by a deposit -feeding polychaete. Limnology and Oceanography 19, 945-
50.
Nunny, R. S. & Chillingworth, P. C. H. 1986. Marine dredging for sand and gravel. London: HMSO.
Odum, E. P. & Odum, H. T. 1959. Fundamentals of Ecology. Philadelphia: W. B. Saunders.
Oliver, J. S. & Slattery, P. N. 1981. Community complexity, succession and resilience in marine soft-bottom
environments. In 6th Biennial International Estuarine Research Co nference. Estuaries 4(3), 259.
Oliver, J. S., Slattery, P. N., Hulberg, L. W. & Nybakken, J. W. 1980. Relationships between wave disturbance
and zonation of be nthic invertebrate communities along a sub-tidal high energy beach in Monterey Bay,
California. Fisheries Bulletin of the United States 78,437-54.
Onuf, C. P. 1994. Seagrasses, dredging and light in Laguna Madre, Texas, U.S.A. Estuarine, Coastal and Shelf
Science 39, 75-91.
Osman, R. W. 1977. The establishment and development of a marine epifaunal community. Ecological
Monographs. 47,37-63.
Oviatt, C. A., Hunt, C. D., Vargo, G. A. & Kopchynski, K. W. 1982. Simulation of a storm event in marine
microcosms. Journal of Marine Research 39,605-18.
Pagliai, A. M. B., Varriale, A. M. C., Crema, R., Galletti, M. C. & Zunarelli, R. V. 1985. Environmental impact
of extensive dredging in a coastal marine area. Marine Pollution Bulletin 16(12), 483-8.
Pearson, T. H. & Rosenberg, R. 1978. Macrobenthic succession in relation to organic enrichment and pollution
of the environment. Oceanography and Marine Biology: an Annual Review 16,229-311.
Peer, D. L. 1970. Relation between biomass, productivity and loss to predators in a population of a marine
benthic polychaete, Pectinaria hyperborea. Journal of the Fisheries Research Board of Canada 27,243-53.
Petersen, C. G. J. 1913. Valuation of the sea. II. Animal communities of the sea bottom and their importance for
marine zoogeography. Report of the Danish Biology Station to the Board of Agriculture 21, l-44.
Pfitzemneyer, H. T. 1970. Gross physical and biological effects of overboard spoil disposal in Upper
Chesapeake Bay. N.R.1 Special Report No. 3 26-35. Chesapeake Biological Laboratory, Solomons,
Maryland. Contr. No. 397.
Pianka, E. R. 1970. On r- and K-selection. American Naturalist 104, 592-7.
Poiner, I. R. & Kennedy, R. 1984. Complex patterns of change in the macrobenthos of a large sandbank
following dredging. Marine Biology 78, 335-52.
Rees, E. I. S., Nicholaidou, A & Laskaridou, P. 1977. The effects of storms on the dynamics of shallow water
benthic associations. In The Biology of Benthic Organisms. B. F. Keegan et al (eds). New York: Pergamon,
465-74.
Rees, H. L. & Dare, P. J. 1993. Sources of mortality and associated life-cycle traits of selected benthic species: a
review. Fisheries Research Data Report, No. 33.
49
Reise, K. 1982. Long-term changes in the macrobenthic invertebrate fauna of the Wadden Sea: are polychaetes
about to take over? Netherland Journal of Sea Research 16,29-36.
Reise, K. & Schubert, A. 1987. Macrobenthic turnover in the subtidal Wadden Sea: the Norderaue revisited after
60 years. Helgoliinder. Meeresuntersuchungen 41,69-82.
Rhoads, D. C. 1974. Organism-sediment relationships on the muddy sea floor. Oceanography and Marine
Biology: an Annual Review 12,263-300.
Rhoads, D. C. & Boyer, L. F. 1982. The effects of marine benthos on physical properties of sediments: A
successional perspective. In Animal-sediment relations, P. L. McCall & M. J. S. Tevesz (eds). New York:
Plenum Press, 3-52.
Rhoads, D. C., McCall, P. L. & Yingst, J. Y. 1978. Disturbance and production on the estuarine seafloor.
American Scientist 66, 577-86.
Rhoads, D. C. & Young, D. K. 1970. The influence of deposit -feeding organisms on sediment stability and
community structure. Journal of Marine Research 28, 150-78.
Riesen, W. & Reise, K. 1982. Macrobenthos of the subtidal Wadden Sea: revisited after 55 years. Helgoliinder
Meeresuntersuchungen. 35,409-23.
Rosenberg, R. 1976. Benthic fauna1 dynamics during succession following pollution abatement in a Swedish
estuary. Oikos 27,414-27.
Saloman, C. H., Naughton, S. P. & Taylor, J. L. 1982. Benthic community response to dredging borrow pits,
Panama City beach, Florida. M. R. 82-3. US Army Corps of Coastal Engineers Research Center, Fort
Belvoir, Va.
Sanders, H. L. 1958. Benthic studies in Buzzards Bay. I. Animal sediment relationships. Limnology and
Oceanography 3,245-58.
Sanders, H. L. 1969. Marine benthic diversity and the stability-time hypothesis. In Brookhaven Symposia in
Biology (22) Diversity and Stability in Ecological Systems. Upton, New York: Brookhaven National Lab.
71-81.
Schafer, W. 1972. Ecology and Palaeoecology of Marine Environments. Chicago : University of Chicago Press.
Shelton, R. G. & Rolfe, M. S. 1972. The biological implications of aggregate extraction: Recent studies in the
English Channel. ICES C.M. 1972/E: 26.
Sherk Jr, J. A. 1971. The effects of suspended and deposited sediments on estuarine organisms. Contrib. No. 443
Chesapeake Biological Laboratory, Solomons, Maryland.
Sherk Jr, J. A. 1972. Current status of the knowledge of the biological effects of suspended and deposited
sediments in Chesapeake Bay. Chesapeake Science 13 (Suppl.), 137-44.
Sherk Jr, J. A., O’Connor, J. M. & Neumann, D. A. 1972. Effects of suspended and deposited sediments on
estuarine organisms. Phase II. Ref No. 72-9E Deptartment of Environmental Research, Chesapeake
Biological Laboratory, University of Maryland.
Sherk Jr, J. A., O’Connor, J. M., Neumann, D. A., Prince, R. D. & Wood, K. V. 1974. Effects of suspended and
deposited sediments on estuarine organisms. Final Report. Ref No. 74-20 Department of Environmental
Research, Chesapeake Biological Laboratory, Prince Frederick, Maryland.
Shumway, S. E., Cucci, T. L. Newell, R. C. & Yentsch, C. L. 1985. Particle selection, ingestion and absorption
in filter-feeding bivalves. Journal of Experimental Marine Biology and Ecology 91, 77-92.
Shumway, S. E., Newell, R. C., Crisp, D. J. & Cucci, T. L. 1990. Particle selection in filter-feeding bivalve
molluscs: a new technique on an old theme. In The Bivalvia- Proceedings of a Memorial Symposium in
Honour of Sir Charles Maurice Yonge, Edinburgh. 1986. B. Morton (ed.). Hong Kong: University Press,
151-65.
Sips, H. J. J. & Waardenburg, H. W. 1989. The macrobenthic community of gravel deposits in the Dutch part of
the North Sea (Klaverbank). Ecological impact of gravel extraction. Reports of the Bureau Waadenburgbv,
Culemborg, The Netherlands.
Smetacek, V. 1984. The supply of food to the benthos. In: Flows of energy and materials in marine ecosystems,
M. I. R. Fasham (ed.). 517-48. New York: Plenum Press.
Snelgrove, P. V. R. & Butman, C. A. 1994. Animal-sediment relationships revisited: cause versus effects.
Oceanography and Marine Biology: an Annual Review 32,ll l -7 7 .
Steele, J. H. 1965. Notes on some theoretical problems in production ecology. Memoire dell’lstituto Italiano di
Idrobiologia Dott Marco de Marchi 18 (Suppl.), 383-98.Steele, J. H. 1974. The Structure of Marine
Ecosystems. Cambridge, Mass: Harvard University Press.
Stephenson, W. W. T., Cook, S. D. & Newlands, S. J. 1978. The macrobenthos of the Middle Banks area of
Moreton Bay. Memoirs of the Queensland Museum 18,95-l 18.
Taylor, P. M. & Saloman, C. H. 1968. Some effects of dredging and coastal development in Boca Ciega Bay,
Florida. United States, Fish and Wildlife Service, Fishery Bulletin 67(2), 213-41.
Thayer, C. W. 1983. Sediment-mediated biological disturbance and the evolution of marine benthos. In Biotic
interactions in recent and fossil benthic communities. M. J. S. Tevesz & P. L. McCall (eds). New York:
Plenum Press, 479-625.
50
Thayer, C. W., Wolfe, D. A. & Williams, R. B. 1975. The impact of man on seagrass systems. American
Scientist 63, 288-96.
Thorson, G. 1957. Bottom communities (sublittoral or shallow shelf). Geological Society of American Memoir
67,461-534.
Urrutia, M. B., Iglesias, J. I. P., Navarro, E. & Prou, J. 1996. Feeding and absorption in Cerastoderma edule
under environmental conditions in the Bay of Marennes-Oleron (Western France). Journal of the Marine
Biological Association of the United Kingdom 76,431-50.
U.S. Army Corps of Engineers, 1974. Draft Environmental Statement - Oyster Shell Dredging Tampa &
Hillsborough Bays, Florida, USA. Army Engineer District, Jacksonville, Florida.
van der Veer, H. W., Bergman, M. J. N & Beukema, J. J. 1985. Dredging activities in the Dutch Waddensea:
effects on macrobenthic infauna. Netherlands Journal of Sea Research 19, 183-90.
van Moorsel, G. W. N. M. 1993. Long-term recovery of geomorphology and population development of large
molluscs after gravel extraction at the Klaverbank (North Sea). Rapport Bureau Waardenburg bv,
van Moorsel, G. W. N. M. 1994. The Klaver Bank (North Sea), geomorphology, macrobenthic ecology and the
effect of gravel extraction. Rapport Bureau Waardenburg and North Sea Directorate (DNZ), Ministry of
Transport, Public Works & Water Management, The Netherlands.
van Moorsel, G. W. N. M. & Waardenburg, H. W. 1990. Impact of gravel extraction on geomorphology and the
macrobenthic community of the Klaverbank (North Sea) in 1989. Rapport Bureau Waardenburg bv,
van Moorsel, G. W. N. M & Waardenburg, H. W. 1991. Short -term recovery of geomorphology and
macrobenthos of the Klaverbank (North Sea) after gravel extraction. Rapport Bureau Waardenburg bv,
Walker, T. A. & O’Donnell, G. 1981. Observations on nitrate, phosphate and silicate in Cleveland Bay, Northern
Queensland. Australian Journal of Marine and Freshwater Research 32, 877-87.
Warren, P. J. 1 9 7 3 . The fishery for the pink shrimp Pandalus montagui in the Wash. MAFF, Laboratory
Leaflet No. 28.
Warwick, R. M. 1986. A new method for detecting pollution effects on marine macrobenthic communities.
Marine Biology 92, 557-62.
Warwick, R. M. & Clarke, K. R. 1991. A comparison of methods for analysing changes in benthic community
structure. Journal of the Marine Biological Association of the United Kingdom. 71,225-44.
Warwick, R. M. & Clarke, K. R. 1993. Increased variabilty as a symptom of stress in marine communities.
Journal of Experimental Marine Biology and Ecology 172,215-26.
Warwick, R. M., Pearson, T. H. & Ruswahyuni 1987. Detection of pollution effects on marine macrobenthos:
further evaluation of the species abundance/biomass method. Marine Biology 95, 193-200.
Weston, D. P. 1988. Macrobenthos-sediment relationships on the continental shelf off Cape Hatteras, North
Carolina. Continental Shelf Research 8(3), 267-86.
Whiteside, P. G. D, Ooms, K. & Postma, G. M. 1995. Generation and decay of sediment plumes from sand
dredging overflow. Proceedings of the 14th World Dredging Congress. Amsterdam, The Netherlands:
World Dredging Association (WDA), 877-92.
Whittaker, R. H. & Levin, S. A. 1977. The role of mosaic phenomena in natural communities. Theoretical
Population Biology 12, 117-39.
Wildish, D. J. & Kristmanson, D. D. 1979. Tidal energy and sublittoral macrobenthic animals in estuaries.
Journal of the Fisheries Research Board of Canada 36, 197-206.
Wildish, D. J. & Kristmanson, D. D. 1984. Importance to mussels of the benthic boundary layer. Canadian
Journal of Fisheries and Aquatic Sciences 41, 1618-25.
Willoughby, M. A. & Foster, D. N. 1983. Middle Banks Humber River plume mapping and hydrographic
surveys. 18th June to 3rd July 1982. Technical Report, Univ esity of New South Wales Water Research
Laboratory, Australia, No. 83 1-13.
Wilson, D. P. 1971. Sabellaria colonies at Duckpool, North Cornwall, 1961-70. Journal of the Marine
Biological Association of the United Kingdom. 51, 509-80.
Windom, H. L. 1976. Environmental aspects of dredging in the coastal zone. Critical Reviews in Environmenal
Control 6,9 1-1 1 0 .
Woodin, S. A. 1991. Recruitment and infauna: positive or negative cues? American Zoologist 31,797-807.
Woodin, S. A. & Marinelli, R. 1991. Biogenic habitat modification in marine sediments:‘the importance of
species composition and activity. Symposia of the Zoological Society of London 63, 231-50.
Wright, D. G. (Coordinator) 1977. Artificial islands in the Beaufort Sea. A review of potential impacts.
Department of Fishery and Environment. Winnipeg, Manitoba. Sept 1977.
51
View publication stats

Newell 1998

Uploaded by

Copyright:

Available Formats

Newell 1998

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Newell 1998

Uploaded by

Copyright:

Available Formats

See discussions, stats, and author profiles for this publication at: https://www.researchgate.

The impact of dredging works in coastal waters: a review of

Article in Oceanography and Marine Biology · January 1998

Lindsay Jane Seiderer

The user has requested enhancement of the downloaded file.

THE IMPACT OF DREDGING WORKS IN

R. C. NEWELL,1 L. J. SEIDERER1 & D. R. HITCHCOCK2

The importance of benthic communities in marine food webs leading to commercially

Early studies include extensive physiological-toxicological work on the potential

Even longer times may be required for biologically-controlled communities that

General features of community structure

Table 1 Table summarizing the population characteristics of r-selected

comprise a “transitional community” that is confined to a restricted habitat in the surface

Estimation of K-dominance curves adjacent to dredging works and other point

on biological resources and on the subsequent rate of recovery of marine community

Ecological succession and the recolonization process

Figure 4 Schematic diagram showing a colonization succession in a marine sediment following

Prior to this, the community is characterized by large populations of a few small

The physical impact of dredging

Impact adjacent to the dredged area

A detailed study by Gajewski & Uscinowicz (1993) in relation to trailer dredging

rapid reduction in suspended sediment concentrations is similar to that reported by

The impact of dredging on biological resources

Impact of dredging on diversity and abundance

The process of recolonization and recovery

highlight the general principles of succession outlined in Fig. 2 (p. 8).

The rate of recovery of biological resources

Community composition and seabed stability

The influence of sediment composition in controlling the nature of communities of

If sediment composition were of importance in controlling biological community

ceedings of Conference on Oceanology International. Spearhead Publications, Surrey Conference Pro-

You might also like