HYDROLOGICAL PROCESSES

Hydrol. Process. (2012)

Published online in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/hyp.9671

Interactions between diatoms and ne sediment

J. I. Jones,1* C. P. Duerdoth,1 A. L. Collins,2,4 P. S. Naden3 and D. A. Sear4
1

School of Biological and Chemical Sciences, Queen Mary University of London, Mile End Road, London E1 4NS, UK
2
Soils Crops and Water, ADAS, Woodthorne, Wergs Road, Wolverhampton, West Midlands, WV6 8TQ, UK
3
CEH Wallingford, Maclean Building, Benson Lane, Crowmarsh Gifford, Wallingford, Oxfordshire, OX10 8BB, UK
4
Geography and Environment, University of Southampton, Higheld, Southampton SO17 1BJ, UK

Abstract:
Excessive mobilization and delivery of ne sediments to water bodies has detrimental impacts on those biotic elements used for
waterbody status classication, including macroinvertebrates, sh and macrophytes. The relationship between ne sediment and
diatoms is a reciprocal one, with diatoms inuencing the production and retention of ne sediments, as well as being impacted by
ne sediment derived from the catchment. Diatoms can increase the retention of ne sediments in benthic environments as a
result of various mechanisms, including shear stress modication, surface adhesion and bed clogging. Enhanced retention of
nes can have important implications for the transfer and fate of sediment-associated nutrients and contaminants. Excessive ne
sediment loadings impact diatom assemblages via shading, burial and scouring. Indirect impacts of increased ne sediment stress
can result from changes in habitat availability, herbivory or predator changes, which cascade down the food chain. Indices based
on the relative abundance of motile species have been proposed for using diatoms to assess waterbody status. However,
disentangling the potential confounding impacts of alternative environmental stressors on these simplistic indices remains a
signicant challenge. Coupling sediment pressure models, capable of predicting the potential impact of mitigation, with
meaningful diatom-based indices, remains a challenge for catchment planning for sediment abatement and the attainment of
improved, or protection of, ecological status. Existing targets for sediment management in river catchments are largely based on
relationships between sediment stress and impacts on sh, but these thresholds have been widely criticized. There remains a need
to develop generic modelling toolkits coupling sediment stress and impacts on a range of biological quality elements to support a
weight-of-evidence approach. Copyright 2012 John Wiley & Sons, Ltd.
KEY WORDS

diatoms; sediment management; bioassessment; siltation; BQE; nutrients

Received 15 March 2012; Accepted 20 November 2012

INTRODUCTION
As water moves through the landscape, it results in the
delivery of ne sediment to rivers and subsequent transport
to the oceans. This is a natural phenomenon that represents
an important pathway in the global geochemical cycle, a key
component of the global denudation system, and an
important measure of land degradation and the associated
reduction in the global soil resource (Walling and Fang,
2003). In recent years, there has been increasing concern
about the inuence of human activities on the amount of ne
sediment delivered to rivers. Here, we dene ne sediments
as particles smaller than 2 mm in size encompassing
inorganic sand (<2000 to >63 mm), silt (<63 to >4 mm)
and clay (<4 mm), as well as organic particles. Natural or
intrinsic levels of sediment in uvial systems are essential to
habitat heterogeneity and ecological functioning (Foster
et al., 2011), with rates of deposition and erosion dependent
upon hydrological conditions. However, activities such as
agriculture (e.g. Collins and Walling, 2007), forestry
operations (e.g. Davies and Nelson, 1993), construction
(e.g. Angermeier et al., 2004), mining (e.g. Turnpenny and

*Correspondence to: J. I. Jones, School of Biological and Chemical

Sciences, Queen Mary University of London, Mile End Road, London E1
4NS, UK
E-mail: j.i.jones@qmul.ac.uk
Copyright 2012 John Wiley & Sons, Ltd.

Williams, 1980) and the urbanization of catchments (e.g.

Hogg and Norris, 1991) have the potential to enhance the
mobilization and delivery of ne sediment to rivers, with the
quality, quantity and timing of the sediment loads received
by rivers being dependent on key sources and delivery
pathways. Increased inputs of ne sediment can lead to
marked physical modications of the river environment
(Owens et al., 2005) with consequent hydrological and
ecological impacts (Waters, 1995; Wood and Armitage,
1997; Wood and Armitage, 1999; Bilotta and Brazier,
2008). In particular, the biological elements used to assess
the ecological quality of waters, as driven by the EU Water
Framework Directive (WFD), may be affected, including
sh, macroinvertebrates and macrophytes (Kemp et al.,
2011; Jones et al., 2012a,b). Here, we review the relationships between ne sediment and diatoms.
Introduction to diatoms

The diatoms or Bacillariophyta are a division of

unicellular photosynthetic eukaryotic algae that are characterized by a siliceous cell wall (frustule) comprised of two
shells (valves) held together by a series of overlapping
hoops (girdle bands). The complex and exquisitely beautiful
architecture of diatom frustules led to a urry of activity by
early microscopists in the mid-19th century (Ktzing, 1833,
1844; Ehrenberg, 1838, 1840; Ralfs, 1843a,1843b; Grunow,

J. I. JONES ET AL.

1860, 1884; Schmidt, 1874; Cleve 1894a,1894b), with these

early works concentrating on description of species. The
separation of species of diatom is largely on features of the
valves, which, as they are formed of inorganic material,
readily preserve in sediments as fossils. Consequently, if the
ecologies of the species are known, the fossil assemblage
can be used to estimate change in past environmental
conditions over time (e.g. Smol and Stoermer, 1999).
The taxonomy of diatoms is not fully resolved (Mann,
1999). Traditionally, they are divided into those that are
radially symmetric, the centric diatoms, and those that are
bilaterally symmetric, the pennate diatoms. The Round
et al. (1990) classication divides diatoms into three
classes; the centric diatoms are grouped in the order
Coscinodiscophyceae, whereas the pennate diatoms
comprise two orders, the Fragilariophyceae, which lack
a raphe (the organelle that diatoms use to move), and the
Bacillariophyceae, which posses a raphe. A further
revision (Medlin and Kaczmarska, 2004) more precisely
reects the evolution of the group. Here, diatoms are
divided into two subphyla: Coscinodiscophytina (the
radial centrics) and Bacillariophytina (the polar centrics
and the pennates), with the latter divided in turn into two
classes, the Mediophyceae (the polar centrics) and
Bacillariophyceae (the pennates including both raphid
and araphid pennate diatoms).
The raphe is a linear ssure through the valve, usually
running the length of the long axis and often bisected by
a central restriction. Mucopolysaccharide strands are
extruded through this slit for attachment or, by further
secretion of the continuous strand and moving the strands
along the raphe, pushing the diatom forward (Edgar and
Pickettheaps, 1983). The raphe is strongly associated with
motility, although some centric and araphid diatoms are
capable of moving themselves to some extent (Round et al.,
1990); raphid diatoms are able to propel themselves over
substrates at up to 35 mm s 1 (Hay et al., 1993). In marine
systems, this motility is associated with vertical migrations
through sediments timed to coincide with tidal and diurnal
cycles (Faur-Fremiet, 1951; Palmer and Round, 1967). In
freshwaters, circadian rhythms in migration through sediments have been observed (Harper, 1976), with movement
to the surface after dawn followed by departure from the
surface after mid-day.
Diatoms occur virtually everywhere that is wet and lit and
probably account for 20% of the global photosynthetic
xation of carbon (Mann, 1999). Cells can be solitary, or in
multi-cell chains or colonies. Some species directly adhere
to substrates, others produce stalks or mucilaginous pads to
raise the cells from the substrate and others produce attached
tubes within which they move (Round, 1965; Barber and
Haworth, 1981). Many species are free oating and form a
large proportion of both marine and freshwater phytoplankton. Other species are either loosely or rmly attached to
substrates, in communities of (mostly) autotrophic microorganisms collectively termed periphyton. The occurrence
of diatom species and their abundance is closely tied to
hydrological conditions (Reynolds, 2000, 2006; Hilton
et al., 2006).
Copyright 2012 John Wiley & Sons, Ltd.

Because of the diversity of species and the habitats they

persist in, diatoms fall into two biological quality elements
(BQE) as dened by the EU WFD (European Parliament,
2000), namely phytoplankton and phytobenthos. Under the
Directive, assessments of both BQE are required for lakes,
whereas only phytobenthos is required for rivers. When
present in the phytoplankton, diatoms are generally
considered along with other algae and cyanobacteria.
However, when in the phytobenthos, diatoms are often
considered alone, despite the other phytobenthos potentially
providing substantial information, particularly when nutrient concentrations are high (e.g. Schaumburg et al., 2004).
The difference in approach between these BQE is largely
due to the methods of sample preparation: for the
identication of benthic diatoms, all the organic matter is
chemically digested to leave cleaned frustules, whereas
planktonic algae are preserved and stained (e.g. with
Lugols iodine) before identication as whole cells.
Difculties in the identication of other Divisions of
freshwater benthic algae to species and a subsequent lack
of autecological information have further contributed to the
focus on diatoms in the phytobenthos (Pan et al., 2000).
Typically, changes in the diatom assemblage (as either
phytobenthos or phytoplankton) are used to assess the extent
of stress from eutrophication (nutrient pollution as dissolved
inorganic phosphorus or to a lesser extent dissolved inorganic
nitrogen (e.g. Kelly and Whitton, 1995; Kelly et al., 2001)
acidication (Battarbee et al., 1999) or salinization (van Dam
et al., 1994). Yet, increased delivery of ne sediment has the
potential to impact diatom assemblages and confound the
interpretation of assessments.
The relationship between ne sediment and diatoms is a
reciprocal one, with diatoms potentially inuencing the
production and retention of ne sediments, as well as being
impacted by ne sediment loads derived from the
catchment. In this paper, we deal with these two reciprocal
effects separately (for a summary, see Table I).

THE IMPACT OF DIATOMS ON FINE SEDIMENTS

Fine sediments come from a variety of sources (Collins
et al., 2011); one potential source is the settlement of algal
particles produced within the water column. Whereas other
planktonic algae largely decompose to organic matter,
diatoms precipitate dissolved silicates in their frustules in
addition to producing organic matter through photosynthesis, contributing to both the organic and inorganic
components of ne sediments. Diatoms are generally
negatively buoyant and will tend to settle where hydraulic
conditions permit (Reynolds, 2006). Hence, both live and
dead diatoms can contribute substantially to the bed load of
ne sediment. Planktonic diatoms can persist once settled,
with a proportion of cells re-entering the phytoplankton on
resuspension. Indeed, as part of their life cycle, certain
species produce resting spores through sexual reproduction,
which sink to the bed and re-enter the phytoplankton when
appropriate conditions occur (Round et al., 1990); it has
been estimated that the spores of the marine diatom
Hydrol. Process. (2012)
DOI: 10.1002/hyp

Copyright 2012 John Wiley & Sons, Ltd.

Indirect
effects

Deposited
material

Suspended
material

Potential cascading effects from

sh

Reduced herbivory due to

clogging of zooplankton

Reduced due to shading

Increased load due to suspended

and depositing diatoms

Phytobenthos

Habitat changes through effects on

macrophytes
Changes to invertebrate community
Potential cascading effects from sh

Reduced due to shading

Physical abrasion particularly by
saltating (ow driven) sand particles
Burial
Erosion of unstable substrate
Impact on nutrient availability
(dependent on nature of sediment)

Increased load due to resuspension of

benthic diatoms
Increased settlement of suspended
particles
Reduced resuspension of deposited
particles

*The Water Framework Directive does not require phytoplankton to be assessed in rivers.

(b)
The impacts of ne
sediment on diatoms

(a)
The impact of diatoms
on ne sediments

Phytoplankton*

Rivers

Potential cascading effects from sh

Impact on nutrient availability

(dependent on nature of sediment)
Reduced herbivory due to clogging
of zooplankton

Reduced due to shading

Increased load due to suspended and

depositing diatoms

Phytoplankton

Phytobenthos

Habitat changes through effects on

macrophytes
Changes to invertebrate community
Potential cascading effects from sh

Reduced due to shading

Physical abrasion particularly by
saltating (wave driven) sand particles
Burial
Erosion of unstable substrate
Impact on nutrient availability
(dependent on nature of sediment)

Increased load due to resuspension of

benthic diatoms
Increased settlement of suspended
particles
Reduced resuspension of deposited
particles

Lakes

Table I. Summary of (a) the impacts of diatoms on ne sediment and (b) the impact of ne sediment on diatoms. Impacts summarized for both phytoplankton and phytobenthos in rivers and lakes

INTERACTIONS BETWEEN DIATOMS AND FINE SEDIMENT

Hydrol. Process. (2012)

DOI: 10.1002/hyp

J. I. JONES ET AL.

Chaetoceros can comprise up to 50% of the deposited

sediments in certain regions (Graham and Wilcox, 2000).
Diatoms are a signicant source of particles where
populations develop to high densities, particularly in lentic
waters; accumulation rates of 0.11 kg m 2 annum 1 of
algal material on the bed of lakes are not uncommon (Rose
et al., 2011). Although planktonic diatoms can develop
dense populations in rivers, this is largely conned to slower
owing or larger rivers, where loss rates downstream are
lower, or rivers where upstream lakes provide a source
population (Reynolds, 2000; Hilton et al., 2006).
Where shear stresses are sufcient, benthic diatoms can
enter the water column (termed tychoplankton accidental
plankton) and thus form another potential source of
suspended ne sediments. However, the presence of a layer
of diatoms (and other algae) increases settlement and
stabilizes ne sediments that have settled (whatever their
source), thereby reducing the likelihood of resuspension
except during higher magnitude extreme events (Biggs
et al., 1998b, 1999a; Gerbersdorf et al., 2008). Several
mechanisms assist the resistance to resuspension, including
shear stress modication, surface adhesion and bed
clogging. Boundary layer thickness increases with increasing thickness of periphyton (Jones et al., 2000a), and
turbulence and ow are altered (Nikora et al., 1997, 1998).
The extracellular polysaccharides produced by diatoms
(for movement or attachment) bind to deposited
particles, thus increasing cohesion and increasing the
resistance to erosion and mobilization (Holland et al.,
1974; de Brouwer et al., 2005; Gerbersdorf et al., 2008).
Although much work has been undertaken in marine and
estuarine systems, where diatoms play a critical role in
the initial stabilization of mud ats and succession to salt
marsh (Holland et al., 1974; Madsen et al., 1993; Austen
et al., 1999; de Brouwer et al., 2005), recent application
of the understanding to freshwaters found similar effects
of diatoms in the stabilization of deposited ne
sediments (Gerbersdorf et al., 2008; Droppo, 2009;
Grabowski et al., 2011). The polysaccharides that
diatoms produce and bind diatom mats together also
bind suspended particles encouraging settlement; this is
an attribute of all diatom mats, not just those on ne
sediments (e.g. on rocks or plants). Despite assertions
that the settlement of ne particles in areas of high
velocity is negligible (e.g. Hynes, 1970), Graham (1990)
found substantial deposition of clay in turbulent, high
velocity patches of glacial rivers associated with growths
of periphyton. Here, the mass of inorganic silt particles
increased at three times that of the organic matter
(Graham, 1990): evidently, substantial amounts of ne
sediments can be trapped by algal layers attached to hard
substrates (Jowett, 2002: see Figure 1). Such effects are
more pronounced with increasing thickness of the algal
layer, particularly when chain-forming taxa dominate the
assemblage. The amount of retained material is proportional to the biomass of algae over a wide range of
ydraulic and sediment conditions (Figure 1). The
composition and structure of layers of periphyton
inuences the impact on ne sediments: layers of diatoms
Copyright 2012 John Wiley & Sons, Ltd.

Figure 1. Relationship between the organic (Ash Free Dry Mass) and
inorganic contents (Ash Mass) of periphyton samples either of mixed
communities dominated by diatoms or of green algae from eld and
experimental studies covering a range of stream types, hydraulic
conditions, substrates and sediment regimes. The dotted line represents
equivalence, i.e. organic = inorganic mass. (Redrawn from Salant (2011).
Data from Graham (1990), van Dijk (1993), Jowett and Biggs (1997),
Collier (2002), Yamada and Nakamura (2002), Kiffney et al. (2003),
Runck (2007) and Salant (2011)).

have distinctly different effects on hydraulics, deposition

and inltration of suspended solids than other algae and
bare substrates, in part dependent upon the production of
mucilage (Salant, 2011). In particular, diatoms reduce
the quantity and depth of particle inltration, presumably
by blocking pore spaces (Salant, 2011). As a consequence of increased settlement, layers of periphyton are
often predominantly inorganic (Figure 1). Although the
silica cell wall of diatoms contributes to the inorganic
content of periphyton layers where they dominate, this
alone cannot explain the high proportion of inorganic
material in diatom dominated periphyton (up to 99%
inorganic; Figure 1). The impact of the retention of ne
sediment by diatoms mats on other organisms is
unknown, but it may have severe implications for
organisms living below the river bed surface (e.g.
salmonid sh eggs).
In addition to increased retention of settled sediments, the
presence of a layer of diatoms (or other algae) on the surface
of anoxic sediments reduces the re-release of phosphate
from sediments. Under anoxic conditions, insoluble phosphate salts become soluble (Mortimer, 1941) and are
released to the water if the sediment surface becomes
anoxic (usually restricted to slow owing/still waters):
photosynthesis by benthic algae oxidizes the sediment
surface leading to precipitation of phosphate salts (Bostrm
et al., 1988; Carlton and Wetzel, 1988). Diatoms (and other
periphyton) also reduce the release of nutrients (e.g.
phosphate and ammonia) from the bed by biological uptake.
Being at the watersediment interface puts diatoms in an
advantageous position to access nutrients released by
microbial activity in the hyperheos, particularly where
regions of up-welling bring nutrients to the surface once
more (Dent et al., 2001; Pretty et al., 2006).
Hydrol. Process. (2012)
DOI: 10.1002/hyp

INTERACTIONS BETWEEN DIATOMS AND FINE SEDIMENT

THE IMPACTS OF FINE SEDIMENT ON DIATOMS

Shading

As diatoms are photosynthetic, they are dependent

upon light; any increase in the turbidity of the water
column caused by suspended ne sediment will reduce
light availability and hence photosynthesis and diatom
biomass. Although phytoplankton are suspended in the
mixed water column and experience an improved light
climate compared with phytobenthos, turbidity affects the
amount of time that they spend in photosynthetic
darkness, with a subsequent impact on average chlorophyll concentrations (Diehl, 2002). Phytoplankton can be
effectively excluded where turbidity is high and the
mixing depth (driven by wind, ow and the morphology
of the water body) is substantially greater than the depth
to which light can penetrate (Diehl et al., 2002). Turbidity
from ne sediments also reduces the light available for
benthic diatoms, both in lakes and in rivers. Clay
additions to experimental streams caused a reduction in
periphyton biomass (Parkhill and Gulliver, 2002). At its
most extreme, constant high turbidity from ne sediment
and other particulates suspended in the water column can
attenuate light to such an extent that benthic diatoms are
excluded from large areas of deeper water (Vermaat and
De Bruyne, 1993). Because of the impact of turbidity on
light availability, gross primary production (measured as
g O2 m 2 day 1) is lower in reaches affected by sustained
loads of ne sediment, for example below placer gold
mines (LaPerriere et al., 1983; Van Nieuwenhuyse and
LaPerriere, 1986; Lloyd et al., 1987; Pain, 1987). A
similar reduction in primary production, benthic chlorophyll and diatom density was reported downstream of
gravel extraction in a French river (Rivier and Seguier,
1985). In a model relating gross primary production to
turbidity (Lloyd et al., 1987), an increase from 1 to 5
NTU caused gross primary production to decline by 313
%, and an increase from 1 to 25 NTU caused a decline of
1350 %. It should be noted that the high turbidity found
below mine workings is generally sustained for much
longer periods than is typical for normal streams, where
high turbidity is associated with high ow events.
Furthermore, it is difcult to separate the effects of light
availability from other effects of ne sediments.
However, shading effects are not restricted to suspended
ne sediment because deposited ne sediment has a
profound shading effect on benthic algae where it settles.
Because of a reduced likelihood of downward reection and
scattering, particles attenuate more light when settled than
when in suspension (Sand-Jensen and Borum, 1984; SandJensen, 1990; Vermaat and Hootsmans, 1991). Although
translucent inorganic particles (e.g. quartz) can provide a
conduit for light through more opaque layers (Losee and
Wetzel, 1983), a layer of deposited ne sediment will
restrict the light available to the algae underneath. If shading
from deposited ne sediment results in death of the lower
strata of the diatom layer, the strength of attachment to the
substrate can be reduced and cause sloughing of the layer.
For motile diatoms, shading from deposited ne sediment
Copyright 2012 John Wiley & Sons, Ltd.

may not present a substantial problem, as they can move

through the deposited sediment to higher light intensities at
the river bed surface (Harper, 1976; Hay et al., 1993). As a
consequence (and for other reasons, see below), there is a
tendency for the diatom assemblage to become dominated
by motile taxa where rates of deposition of ne sediments
are high (Yamada and Nakamura, 2002; Dickman et al.,
2005), and there is the potential for total diatom biomass to
compensate for losses due to the shading effects of
deposited ne sediment. In indoor-controlled experimental
channels, the initial response to a short-term (3 days)
addition of sediment was a reduction in benthic chlorophyll-a and photosynthetic efciency, followed by a
recovery to almost full compensation, but with different
assemblage composition, a month after the deposition event
(Izagirre et al., 2009). Where frequent pulses of surface
deposition of ne sediment occur, the potential for recovery
by periphyton will be compromised, which in turn affects
diatom assemblage composition and important ecosystem
processes such as primary production.
Burial and erosion

In addition to reducing the light available to benthic

diatoms, excessive deposition of ne sediments will result
in burial. In turn, because of the altered ow patterns and
biological activity, this will alter the physical and
chemical environment in which the buried diatoms nd
themselves. Concentrations of reactive and potentially
toxic reduced ions (e.g. Fe2+ and Mn2+) can be high in
anoxic sediments (Wetzel, 1983). Furthermore, deposited
sediments may contain other toxic substances such as
heavy metals, which can have a negative impact on
diatom assemblages (Ivorra et al., 2000, 2002). Whereas
motile forms can migrate to the surface, non-motile forms
will have to endure these potentially toxic conditions until
erosive ow events bring them to the river bed surface
once more.
However, the most profound effect of depositing ne
sediment is the smothering of substrata to which diatoms
attach. The relatively unstable deposited ne sediments
(compared with larger particles) are not suitable for the
attachment of long-lived sedentary species. Hence, nonmotile and particularly chain-forming diatoms cannot
establish easily, further pushing the assemblage towards
single celled and motile taxa. A shift in assemblage
composition towards motile taxa can be seen even where
larger particles are covered with a layer of nes (Dickman
et al., 2005). The lack of stability in patches where easily
erodible ne sediments accumulate, either accreting or
eroding dependent upon ow conditions, tends to result in
reduced taxon richness and biomass compared with more
stable patches (Biggs et al., 1998a; Biggs and Smith, 2002;
Matthaei et al., 2003). When comparing across streams,
those with stable bed sediments support a higher biomass of
diatoms than those that have unstable beds, for example
because of high amounts of deposited ne sediment (Iversen
et al., 1991; Biggs, 1995; Jowett and Biggs, 1997; Biggs
et al., 1999a; Biggs and Smith, 2002).
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J. I. JONES ET AL.

However, the dynamics of benthic diatom assemblages

within a river may not be quite as straight forward; river beds
are a mosaic of eroding and depositing patches, with
disturbance history having both long-term and short-term
effects on benthic diatoms. In the river Isar, Germany,
Matthaei et al. (2003) found that 3 months after a ood, algal
biomass, total diatom density and diatom taxon richness
were highest in depositional patches (where chains of
diatoms were found buried in the substratum), immediately
(6 days) after a second ood biomass was highest in stable
patches, and 4 weeks later, diatoms were most abundant in
the erosive patches. They concluded that assemblage
succession and disturbance history interacted to determine
how patches developed (Matthaei et al., 2003). The
assemblage that develops will depend on rate of deposition
of ne sediments and the frequency with which these
deposits are disturbed and remobilized by high ow events.
Scouring

Even when attached to hard substrates, physical abrasion

by suspended particles can damage or dislodge diatoms
(Francoeur and Biggs, 2006). This is particularly true of
saltating sand particles, which can cause substantial damage
to cells (Delgado et al., 1991). Hence, diatom assemblages
do not develop well on introduced, articial substrata in
streams with highly mobile sand substrates (SkytteJohannsen, 2008). The susceptibility to scouring and
sloughing depends on the morphological structure of both
the individual and the community (Francoeur et al., 1998),
their physiological condition (Biggs and Close, 1989; Biggs
et al., 1999b; Biggs and Smith, 2002), and local ow and
substrate conditions. Similarly, the diatoms that grow
attached to substrate particles can be damaged if the bed is
disturbed. For diatoms attached to larger particles (e.g.
boulders), burial is likely to be more important than physical
damage caused by the particle moving. Diatoms attached to
smaller particles are more likely to be damaged when the
substrate they are attached to moves. Unconsolidated sand
grains typically have infrequent small, adpressed taxa (e.g.
Staurosira species) growing on them, which because of
their low stature and robust cell walls are more resistant to
damage (Jewson et al., 2006): even these taxa are lost if
disturbance is frequent (Vilbaste and Truu, 2003). As a
consequence of this scouring by ne particles, even suitable
substrates do not develop a rich diatom ora where the river
bed is unstable and comprises ne inorganic particles
(particularly sand).
Nutrient availability

Not all impacts of ne sediments on diatoms are

detrimental. Deposited ne sediments are often rich in
nutrients, and the anoxic conditions that can develop
(particularly where the organic content is high) result in
re-mobilization of these nutrients. As benthic diatoms sit on
the interface between the sediment and the water, they are in
the ideal position to exploit these nutrients. Hence, nutrientrich deposits of ne sediment can result in increased growths
of diatoms and a shift in the ora towards species
Copyright 2012 John Wiley & Sons, Ltd.

characteristic of nutrient-rich conditions. The outcome for

the diatom ora depends on the balance of negative and
positive effects of ne sediments on diatom growth, in part
inuenced by the rate of deposition and stability of
deposited material. Hence, the response to increasing
deposited sediment may not be linear. In a survey of New
Zealand rivers, benthic algal chlorophyll-a showed a stress
subsidy response to sediment, i.e. an initial increase
followed by a decline at higher bed sediment loads, with
little response to dissolved nutrients in the water (Wagenhoff
et al., 2011). Compound effects can also occur where
multiple stressors interact, e.g. increased sediment and low
ows (Matthaei et al., 2010). The effect of enrichment by
ne sediment is particularly apparent in permeable gravels,
where regions of down-welling result in recycling of
nutrients from ne sediments entrained within the river
bed; diatoms attached to hard substrates can exploit these
recycled nutrients when they return to the surface in regions
of up-welling hyporheic water (Dent et al., 2001; Pretty
et al., 2006). Where ne sediments are both unstable and
infertile, i.e. inorganic sands, increased growth offers little
compensation for the negative effects described previously.
On the other hand, nutrient-rich ne sediments, if relatively
stable, can be colonized by lamentous green algae such as
Cladophora glomerata agg. and Enteromorpha intestinalis;
although presenting opportunities for epiphytic diatoms,
these large structural species encourage further deposition of
ne sediments in turn and reduce the benthic diatom
assemblage that develops underneath them through shading
and burial (Sand-Jensen et al., 1989b; Dodds, 1991).
Indirect effects

Diatoms are not the only biological component of

freshwaters to be affected by increased loads of ne
sediment. Invertebrates (Jones et al., 2012a), sh (Kemp
et al., 2011), macrophytes (Jones et al., 2012b) and other
algae are all affected by increased loads of ne sediment,
which in turn has consequences for the diatom assemblages
that they interact with. Of particular note are macrophytes,
which inuence ow and the rate of retention of nes (Jones
et al., 2012b) and have a profound effect on the light climate
that benthic algae experience with consequent impacts on
their growth and standing stock (Sand-Jensen et al., 1989b).
Whereas direct shading by macrophytes will reduce benthic
algal growth underneath stands (Sand-Jensen, 1997), the
macrophytes themselves will be colonized by growths of
diatoms (Sand-Jensen et al., 1989a). In itself, this should not
inuence assessments of ecological quality, as most WFD
compliant protocols stipulate that the diatom assemblage on
macrophytes should be sampled where hard substrates are
lacking (e.g. WFD-UKTAG, 2008a,2008b), but the different conditions experienced by epiphytic diatoms compared
with their benthic counterparts will result in differences in
assemblage composition, adding to the uncertainty of
assessments.
Changes in the animal community as a consequence of
increased loads of ne sediment will have an effect on
diatoms either directly, where herbivores are affected, or
Hydrol. Process. (2012)
DOI: 10.1002/hyp

INTERACTIONS BETWEEN DIATOMS AND FINE SEDIMENT

indirectly where effects on predators (e.g. sh) cascade

down to diatoms (Bronmark and Vermaat, 1998; Diehl and
Kornijw, 1998; Jones et al., 1998). Herbivory can have a
marked effect on benthic algal communities, potentially
reducing standing stock to such an extent that responses to
nutrient enrichment are masked (Jones et al., 1999;
Hillebrand, 2002; Jones et al., 2002). Hence, the humped
response of benthic chlorophyll-a to ne sediment bed load
observed in New Zealand streams by Wagenhoff et al.
(2011) may have been a consequence of grazing by
increased densities of the mud snail, Potamopyrgus
antipodarium (Gray), where bed loads were highest, rather
than a direct response of algae to bed sediment. Most
importantly, grazing by invertebrates has a profound effect
on community composition (Jones et al., 2000b), potentially
confounding interpretation of diatom assemblages. Often,
small rapidly growing species dominate where grazing is
intense (Jones et al., 2000b; Hillebrand, 2002): many
species with these characteristics are indicative of high
nutrient conditions (Kelly et al., 2001). Nevertheless,
predicting the consequences of such interactions between
ne sediment, diatoms and fauna is difcult, as direct and
indirect pathways are likely to have contrasting effects.
Similar effects of herbivory on planktonic algal
communities are apparent, with grazing by zooplankton
being a major determinant of community composition and
standing stock (Jeppesen et al., 1999; Reynolds, 2006).
Effects of ne sediment on zooplankton, for example
through the clogging of lter combs or supply of organic
particles (Arruda, 1983; Arruda et al., 1983; Hart, 1992),
could potentially inuence the rate of herbivory, and thus
diatom assemblages in standing waters. Further indirect
effects of ne sediments on diatoms may be apparent
where sh populations are affected. The cascade from sh
through zooplankton to phytoplankton is well described
(Carpenter and Kitchell, 1993): any pronounced effect on
sh is likely to propagate through the community.

DIATOM SENSITIVITY TO SEDIMENT AND THE

CLASSIFICATION OF WATER BODY STATUS
Because of the preservation of their frustules in some
lake deposits, diatoms have been used to reconstruct past
environmental conditions. Hence, relationships between
diatom assemblage composition and a variety of environmental parameters have been derived. In the early 20th
century relationships with trophic status (Naumann, 1921),
salinity (Kolbe, 1927) and pH (Hustedt, 1937-9) were
developed where species were classied to categories of
sensitivity. This approach has proven useful in a number of
ways: the relationships with trophic status have been rened
and now form the basis of the various indices used
to classify water bodies for the WFD using diatoms, e.g.
Diatom Assessment for River Ecological Status and Diatom
Assessment for Lake Ecological Quality (based on Trophic
Diatom Index (Kelly and Whitton, 1995; Kelly et al., 2009):
WFD-UKTAG, 2008a,2008b), Indice de Polluosensibilit
(CEMAGREF, 1982; Kahlert et al., 2009), Biological
Copyright 2012 John Wiley & Sons, Ltd.

Diatom Index (Coste et al., 2009), the German Trophic

Diatom Index (Coring et al., 1999), and the Trophieindex,
Saprobienindex and Referenzartenindex (Rott et al., 1997,
1999). The response to salinity has been used to describe
climate change using the sediments from endorheic (closed
basin) lakes (Fritz et al., 1991), and the relationships with
pH have been further developed (e.g. Battarbee, 1986;
Battarbee et al., 1999; Smol, 2008) and used to detect the
acidication of lakes as a consequence of atmospheric
pollution (Charles et al., 1989; Battarbee et al., 1990; Dixit
et al., 1992). Recent environmental construction works have
moved away from the categorization of species responses to
more numerical approaches such as transfer functions,
where individual optima are determined from training sets
(see Juggins et al., 1995), although the ability of either
approach to discern changes in a single environmental
variable against a background of variation in many
parameters has been criticized (Sayer, 2001; Sayer et al.,
2010); it has been suggested that the use of multivariate
regression tree models, which can describe the effects of
multiple drivers, are far superior for environmental
reconstruction (e.g. Davidson et al., 2010). Such confounding effects of multiple drivers have been noted before: the
TDI is unable to distinguish between eutrophication and
organic pollution (Kelly and Whitton, 1995), such that the
proportion of the sample that is tolerant of organic pollution
is used to aid interpretation.
As ne sediments inuence diatom assemblages in many
ways, it is not surprising that indices based on benthic
diatom assemblage structure have been proposed. These
comprise simply of the relative abundance of motile species
(e.g. Bahls, 1993). This measure is based on the fact that
many raphid species are capable of migrating through
deposited sediment to avoid its negative impacts. However,
the exact list of species that are motile varies amongst
authors (Table II). Furthermore, the use of such an index is
prone to complication as species respond to ne sediments
in ways that are correlated with other pressures. Kelly et al.
(2007) note that the gradient of TDI ecological quality ratio,
indicating a shift from nutrient sensitive to nutrient tolerant
taxa, is also correlated with a shift to motile taxa; although
this may indicate a functional change in the assemblage,
such change confounds interpretation when motility
responds to sediment load rather than nutrient availability
(and vice versa). Porter et al. (2008) reported that the
abundance of taxa with continuously high dissolved oxygen
requirements decreased signicantly with increases in
nutrient and suspended sediment concentrations, confounding interpretation of indices developed for the detection of
organic pollution (Lowe, 1974; van Dam et al., 1994).
Similarly, impacts of suspended sediment that do not entail
burial (e.g. scouring) are not reected in indices of motility,
and their effects may be interpreted as evidence of other
stressors. Such issues are not exclusive to diatoms but arise
whenever BQEs are interpreted as change across a single
gradient of pressure (Jones et al., 2010). More complex
models of diatom assemblage response, such as classication and regression trees (CART) and RIVPACS (River
Invertebrate Prediction and Classication System) models,
Hydrol. Process. (2012)
DOI: 10.1002/hyp

J. I. JONES ET AL.

Table II. Motile diatom taxa as dened by various workers and assessment methods. Where taxonomic revisions have occurred, taxa
included as previous classication
Molloy
(1992)a

Bahls
(1993)

Kutka and
Richards (1996)

Stevenson and
Bahls (1999)b

Kelly et al.
(2001)

Dickman et al.
(2005)

Amphipleura
Anomoeoneis
Bacillaria
Brachysira
Caloneis

a
a
a

Kelly et al.
(2005)

Spaulding et al.
(2010)i

Bacillaria

Bacillaria
Brachysira
Caloneis
Campylodiscus
Capartogramma

g
g

Cavinula
Craticula

Craticula

Craticula
Cylindrotheca
Cymbellak
Denticula
Diadesmisl
Diploneis

Cylindrotheca
Diadesmis

a
a
a

Frustulia

Gyrosigma
Hantzschia

Navicula
a

Navicula
Nitzschia

Navicula
Nitzschia

Navicula
Nitzschia

Navicula
Nitzschia
Pinnularia

Diploneis
Eunotiac

Frustulia

Gyrosigma
Hantzschiah
Navicula
Nitzschiad
Pinnularia

Navicula
Nitzschia
g

Placoneis
a

Surirella

Surirella

Stauroneis
Surirella

Epithemia
Frustulia
Gomphoneis
Gomphonema

Sellaphora

Sellaphora

Surirella

Synedra acusf

Mastogloia
Navicula
Nitzschia
Pinnularia
Placoneis
Rhopalodia
Sellaphora
Stauroneis
Surirella

Specically stipulates that Amphipleura, Anomoeoneis, Bacillaria paradoxa Gmelin, Caloneis, Diploneis, Eunotia, Frustulia, Gyrosigma, Hantzschia
amphioxys (Ehr.) Grun., Nitzschia and Stauroneis are non-motile.
b
Also includes an unspecied and other motile taxa.
c
Eunotia exigua described as motile, Eunotia minor as attached.
d
Nitzchia heueriana described as attached, other members of genus as motile.
e
Surirella minuta and Surirella roba described as attached, other members of genus not described.
f
Araphid: other members of this genus described as attached.
g
Taxa indicated as being Attachment: Not attached (oating or freely motile) at genus level are not included as motile if all constituent species are
described as Motile: No. Named non-motile taxa: Amphipleura pellucida, Anomoeoneis sphaerophora, Brachysira vitrea, Caloneis amphisbaena,
Caloneis bacillum, Caloneis silicula, Caloneis subsalina, Diploneis elliptica, Diploneis oblongella, Eunotia exigua, Eunotia formica, Eunotia pectinalis,
Frustulia rhomboides, Frustulia vulgaris, Pinnularia appendiculata, Pinnularia borealis, Stauroneis anceps, Stauroneis kriegeri, Stauroneis
phoenicenteron, Stauroneis smithii.
h
Included as motile here because motility mentioned in ecological notes despite the only constituent species, Hantzschia amphioxys, being described as
non-motile.
i
Five categories of motility given: 1 non-motile; 2 weakly motile, e.g. Actinella; 3 slightly motile, e.g. Cocconeis; 4 moderately motile. e.g. Navicula
tripunctata; 5 highly motile, e.g. a large cell-like Surirella; and extensive raphe. For consistency, categories 4 and 5 are considered motile here, with
highly motile taxa indicated by italics.
j
Details of motility not given.
k
Cymbella mexicana and Enyconema reimeri described as highly motile; Cymbella janischii, C.proxima, Cymbellonitzschia and Encyonopsis described
as moderately motile. Other constituent taxa described as slightly motile.
l
Diadesmis confervacea moderately motile, D. pantropica slightly motile.
m
Taxon not included either as Hantzschia or Nitzschia spp.

appear more capable of interpreting the impact of stressors

on diatom assemblages than simple indices (Cao et al.,
2007; Feio et al., 2007).
Existing targets for sediment management in river
catchments are largely based on relationships between
sediment stress and impacts on sh, especially salmonids,
but these thresholds have been widely criticized. Given
the requirement to revise targets for informing river
Copyright 2012 John Wiley & Sons, Ltd.

catchment sediment management (Collins et al., 2011),

there is a need to continue exploring the scope for
coupling sediment pressure models with diatom indices.
The pressure models should have the capability for
predicting past, present or future projected sediment loadings
as well as the expected benets of targeted mitigation options
such as those on-farm measures supported by existing agrienvironment schemes. Coupling the sediment stress models
Hydrol. Process. (2012)
DOI: 10.1002/hyp

INTERACTIONS BETWEEN DIATOMS AND FINE SEDIMENT

with diatom indices should sit alongside the corresponding

linking up of predicted sediment pressures and impacts on a
range of alternative BQEs, including sh and macroinvertebrates. In combination, this approach would support a
weight-of-evidence methodology for supporting improved
sediment management for freshwater ecological status.
CONCLUDING REMARKS
To conclude, it is apparent that excessive ne sediment
pressures have various impacts on planktonic, and particularly benthic, diatom assemblages. To date, detection of
these impacts has relied solely on the proportion of motile
taxa (variously dened: see Table II), an index that is likely
to detect only some of the effects of increased loading of ne
sediment. Furthermore, the impacts of ne sediments are
likely to confound the interpretation of indices designed to
determine the impact of other stressors. The majority of
diatom indices used for bioassessment have been designed
to predict dissolved nutrient (particularly phosphorus)
availability; the full extent of impacts of ne sediment
other than through burial are not known but may result in an
imprecise diagnosis of nutrient stress. Nevertheless, the
extent of impact of increased ne sediment loading is likely
to be dependent upon context; the sensitivity of biological
communities is largely dependent upon the unimpacted
condition, and a blanket prediction for all river types is
unlikely to be adequate. Currently, we lack anything more
than cursory investigations into the effects of increased
sediment loads on diatoms, and the tools available for
routine monitoring are insufciently sophisticated to detect
where increased loading of ne sediment is having a
detrimental impact (Feio et al., 2007). Detailed investigations are required that take into account, and separately
identify, the impacts of both nutrient and sediment pollution
across a range of river types. A potential approach might be
to link the archives of sediment accumulation recorded in
river slackwater deposits, or in lake sediments, with changes
in fossil diatom assemblages during periods prior to major
intensive human disturbance, thus reducing the obscuration
resulting from multiple pressures. Without further renements of diatom-based indices to take into account the
impacts of ne sediment, it is likely that misdiagnosis of
pollution pressures on water resources will occur.
ACKNOWLEDGEMENTS

The authors gratefully acknowledge the funding provided

by the Department for Environment, Food and Rural Affairs
(Defra, contract WQ0128; Extending the evidence base on
the ecological impacts of ne sediment and developing a
framework for targeting mitigation of agricultural sediment
losses). Thanks are extended to Martin Kelly and an
anonymous reviewer for helpful suggestions.
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