The Impact of Fine Sediment on Macroinvertebrates

RIVER RESEARCH AND APPLICATIONS River Res. Applic. (2011) Published online in Wiley Online Library (wileyonlinelibrary.com) DOI: 10.1002/rra.1516

THE IMPACT OF FINE SEDIMENT ON MACRO‐INVERTEBRATES

J. I. JONES,a* J. F. MURPHY,a A. L. COLLINS,b,c D. A. SEAR,c P. S. NADENd and P. D. ARMITAGEe a School of Biological and Chemical Sciences, Queen Mary University of London, London, UK b Soils Crops and Water, ADAS, Wolverhampton, West Midlands, UK c School of Geography, University of Southampton, Southampton, UK d Centre for Ecology and Hydrology, Wallingford, Oxfordshire, UK e Freshwater Biological Association, The River Laboratory, Wareham, Dorset, UK

ABSTRACT The sustainable use of water resources requires clear guidelines for the management of diffuse pollution inputs to rivers. Without informed guidelines, management decisions are unlikely to deliver cost‐effective improvements in the quality of rivers as required by current water policy. Here, we review the evidence available for deriving improved guidelines on the loading of fine sediment to rivers based on the impact on macro‐invertebrates. The relationship between macro‐invertebrates and fine sediments is poorly defined. Studies of the impacts of fine sediment on macro‐invertebrates have been undertaken at various scales, which has an influence on the range of responses displayed and the reliability of the results obtained; results obtained from investigations at smaller scales may not manifest at the scale required to manage rivers and vice versa. Many of the identified effects of increased loading of fine sediment on macro‐invertebrates occur as a consequence of deposition on the river bed, yet many current management guidelines are based on suspended sediment targets. On this basis, existing water quality guidelines for sediment management are unlikely to be appropriate. Copyright © 2011 John Wiley & Sons, Ltd. key words: macro‐invertebrates; benthos; deposition; suspended solids; turbidity; bioassessment; thresholds

Received 1 July 2010; Revised 22 October 2010; Accepted 10 February 2011

INTRODUCTION treatment sources) would, therefore, help resolve any confounding influence and guide appropriate management Because of their ubiquity, high diversity and range of decisions. sensitivities, macro‐invertebrates are used extensively to The erosion and deposition of fine sediment are intrinsic and assess pollution in freshwaters. Much of the focus of the natural components of the hydro‐geomorphic processes of science of biomonitoring (using biological samples to fluvial systems. Indeed, aquatic biota contribute significantly determine the extent of stress) has been driven by concerns to the production [e.g. faecal particles, phytoplankton (Wotton about the impact of sewage pollution on potable water and Malmqvist, 2001; Wharton et al., 2006; Wotton and resources (Jones et al., 2010). Nevertheless, it has long Warren, 2007)], processing [e.g. breakdown of detritus, been noted that fine sediment (defined here as inorganic and nutrient cycling (Trimmer et al.,2009)]anddownstream organic particles of less than 2 mm in diameter) can have a conveyance [e.g. bioturbation, retention by macrophytes detrimental effect on aquatic invertebrate communities (e.g. (Barko et al., 1991; Vaughn and Hakenkamp, 2001; Nogaro Ellis, 1936). Such observations have lead to efforts to et al.,2006;Nogaroet al.,2009)]offine sediments. However, describe and quantify the impact of fine sediments on human activities in catchments have resulted in increased invertebrates. Largely, these investigations have been driven delivery of fine sediment to watercourses, such that the loading by an interest in the effects of fine sediment per se. of fine sediment to many rivers now far exceeds background However, any impact of fine sediment (from sources other (pre‐industrial) conditions (Walling and Fang, 2003; Foster than sewage effluent) on macro‐invertebrate communities is et al., 2011). Concerns about the impact that such increased likely to confound interpretation of these communities when loads are having on the ecology and geomorphology of they are viewed in terms of organic pollution alone. An freshwaters have led to suggestions that fine sediment is one of improved understanding of the effects of fine sediment from the most widespread and detrimental forms of aquatic all key catchment sources (including that from water pollution (Ritchie, 1972; Lemly, 1982). The impacts on biota are wide ranging and can be profound (e.g. Waters, 1995; *Correspondence to: J. I. Jones, School of Biological and Chemical Wood and Armitage, 1997; Collins et al., 2011; Kemp et al., Sciences, Queen Mary University of London, Mile End Road, London, fi E1 4NS, UK. 2011; Jones et al., 2011); here we review the effects of ne E‐mail: [email protected] sediment on macro‐invertebrates.

An important aspect of the management of fine sediment as a behavioural response to increased sediment load, see loads in rivers is determination of an acceptable level of succeeding sections) or cause damage to their body parts (as input from key catchment sources (Collins and Anthony, for fish; Newcombe and MacDonald, 1991). Particles need 2008; Cooper et al., 2008; Collins et al., 2009). Unlike not be truly in suspension, and it is likely that larger many other pollutants, a certain amount of fine sediment is particles, particularly sand, moving by saltation or as necessary for rivers to function normally, and it is only where bedload cause the most damage. In experimental channels, excess loading occurs that the negative impacts of fine saltating sand caused catastrophic drift, which was not seen sediment are expressed. Here, we review the mechanisms by in patches where sand deposited (Culp et al., 1986). which fine sediment can impact macro‐invertebrates and the Nevertheless, smaller particles may be equally damaging if difficulties in translating the available evidence of impacts moving at high velocity. Unprotected, fine and fleshy body into practical targets for fine sediment management in river parts, such as gills and filter‐feeding apparatus, are catchments (Figure 1). particularly prone to damage, with obvious consequences for the individuals affected. Behavioural responses to protect such sensitive structures from abrasion can ensue PHYSICAL EFFECTS OF INCREASED [such as retraction of filter combs; (Kurtak, 1978)], although FINE SEDIMENT LOADING such avoidance behaviour inevitably disrupts normal functioning. Brachycentrus (Trichoptera) switch from Abrasion filtering with limbs extended into the water to trap particles Increased loads of fine sediment result in both increased (Gallepp, 1974) to grazing when suspended sediment loads concentrations of suspended particles and increased rates of are high (Voelz and Ward, 1992), presumably because of deposition (Walling, 1995; Sear et al., 2008). Although abrasion from particles (although possibly because of difficult to quantify, it is likely that invertebrates are prone reduced quality of food; see Food availability and quality to abrasion from particles in the flow, which in extreme section). Where retraction is not possible, individuals may circumstances, may cause individuals to become dislodged retreat to areas of low velocity or spend time cleaning and (although invertebrates will also release from the substrate repairing damaged structures (Edington and Hildrew, 1995).

Figure 1. Summary diagram illustrating the direct and indirect mechanisms by which ﬁne sediments impact upon macro‐invertebrates. Impacts are caused by both suspended and deposited particles. Arrows show interacting effects and impacts on macro‐invertebrates at the individual, species and community levels (collectively represented here as a mayﬂy larvae). The strength and direction (+/−) of effects are not given as they are dependent upon the taxa affected; some taxa and communities respond positively to changes, others negatively.

Clogging are adapted to live in fluvial depositional zones and benefit from the rate of influx of organic (food) particles. In such As well as physical damage by abrasion, the transport of habitats, invertebrates are usually motile, albeit very slow in fine particles, particularly clays and silts, can result in a some circumstances (e.g. Unionidae), and move to keep pace build up on the organs, disrupting the normal functioning of with accreting sediment. Issues arise when the rate of gills and filter‐feeding apparatus, making respiration and accretion exceeds the ability of individuals to excavate feeding difficult. Many species of invertebrates collect themselves, which is highly dependent upon individual taxa particles from the water by filter feeding and have a variety and the particle size of deposited materials (Wood et al., of special structures for doing so. Bivalve molluscs are 2005). Sedentary invertebrates typical of habitats where the capable of expelling unwanted particles from their gills but intrinsic rates of accretion are low are particularly vulnerable, can expend considerable energy and mucus in producing generally not having the ability to keep pace with increased pseudofaeces (unwanted uningested particles) in order to accretion [e.g. Margaritifera margaritifera (L.) (Geist and expel sediment particles from their gills (MacIsaac and Auerswald, 2007; Osterling et al., 2008)]. Smaller individ- Rocha, 1995; Iglesias et al., 1996), thus considerably uals (Wood et al., 2001) and certain life stages, such as reducing the efficiency of feeding. Where loads are immotile eggs, can be more vulnerable to burial than others. sufficiently high, more energy can be expended in removing Although direct effects of burial may not affect such unwanted particles than is gained from feeding, leading to vulnerable stages (e.g. through lack of access to food), the cessation of feeding where loads of suspended silt are high increase in the mortality of caddis fly pupae after burial was (Ellis, 1936). Like molluscs, Cladocera also reject unwanted attributed to a reduced oxygen supply because of reduced particles from their filter combs and spend an increased water movement around the pupal cases (Rutherford and proportion of their time (and energy) cleaning these Mackay, 1986). However, prediction of which taxa are structures where concentrations of inorganic suspended sensitive to burial is not straight forward. Nymphs of the solids are high, but these negative effects can be balanced mayfly Baetis rhodani (Pictet) are highly sensitive to burial, where Cladocera utilize increased loads of fine particulate whereas nymphs of the stonefly Nemoura cambrica organic matter (Arruda, 1983; Arruda et al., 1983; Hart, (Stephens) and larvae of the caddis fly Melampophylax 1992). Blackfly (Simuliidae) larvae are less selective in mucoreus (Hagen) are capable of excavating themselves what they consume, ingesting most of the material even after substantial (1 cm) burial (Wood et al., 2001; Wood collected. Hence, the ingestion rate of blackfly larvae et al., 2005). Larvae of the caddis fly Potamophylax declines with increasing inorganic suspended sediment as cingulatus (Stephens) abandon their cases to escape burial their guts fill with the inert particles (Gaugler and Molloy, (Dobson et al., 2000), whereas the larvae of the caddis flies in 1980). Feeding rate is impaired at concentrations of − the families Molannidae and Goeridae prefer a thin covering particles <125 µm greater than 50 mg L 1 (Kurtak, 1978). of fine sediment (Wallace et al., 1990). Substantial burial by Similarly, many species of caddis fly larvae produce nets of large particles (>2 mm) is required to trap the water slater silk to trap passing particles. These nets can become Asellus aquaticus (L.), whereas larvae of the caddis fly clogged with fine sediments, leading to an increase in Hydropsyche pellucidula (Curtis) become trapped by smaller cleaning activity and eventual abandonment (Edington and particles (<500 µm) if deposited in a thick layer (Wood et al., Hildrew, 1995; Strand and Merritt, 1997). Filter‐feeding 2005; Table I). Although physical entrapment of inverte- blackfly and caddis fly larvae were notably absent from a brates by burial can cause mortality by reducing access to stream receiving high inputs of fine sediment from a food, it is the changes in the chemical environment that ensue military training area when compared to an adjacent that have the most profound effects (e.g. reduced supply of undisturbed catchment (Armitage et al., 1999; Armitage oxygen, see Oxygen concentration section). and Blackburn, 2001).

Burial Substrate composition Burial can present difficulties for sedentary animals and, Together with increased accretion, fine sediment deposition where rates of deposition are high, even motile animals can results in changes to the composition of the bed of rivers. be affected. This was one of the earliest noted effects of Where inputs of fine sediment to catchments are increased, increased sediment inputs; increased mortality in certain the average size of particles becomes smaller, interstices species of bivalve molluscs was noted as a consequence of between larger particles become filled and, where a surface rapid burial with most of the common species tested not drape of deposited sediment occurs, the stability of the bed being able to survive deposition of a layer of silt 0.6–2.5 cm may be reduced (Kaufmann et al., 2009). Although these thick over the bed but capable of persisting on raised changes can alter the flow of water through the river bed, with platforms (Ellis, 1936). Yet, many species of invertebrates consequent impacts on the hyporheic chemical environment

Table I. Effects of ﬁne sediment on stream and river macro‐invertebrates

Species Effect Sediment load Sediment type Author

LABORATORY ASSESSMENTS

Baetis rhodani Ephemeroptera Burial 5 mm depth 125 µm to >4 mm (Wood et al., 2005) Asellus aquaticus Isopoda Burial 5 mm depth >4 mm (Wood et al., 2005) 10 mm depth >2 mm Hydropsyche pellucidula Trichoptera Burial 10 mm depth <500 µm (Wood et al., 2005) Ephemerella subvaria Ephemeroptera Increased drift 2.68 g L−1 Potter’s clay (Ciborowski et al.,1977) Simuliidae Diptera Feeding >50mgL−1 <125 µm (Kurtak, 1978; Gaugler Inhibition and Molloy, 1980) Pteronarcys californica Plecoptera Feeding 1.5 mg L−1 (Hornig and Brusven, inhibition 1986) Hesperophylax occidentalis Trichoptera Feeding 1.5 mg L−1 (Hornig and Brusven, inhibition 1986) Bivalvia Feeding 600 mg L−1 Diatomaceous (Aldridge et al., 1987) inhibition earth Bivalvia Reduced 600 mg L−1 Diatomaceous (Aldridge et al., 1987) metabolism earth Cladocera Feeding 25 000 mg L−1 (Alabaster and Lloyd, Copepoda inhibition 1982) Daphnia spp. Cladocera Positive and ≈200 NTU Lake sediment (Hart, 1992) negative (≈200 mg L−1) effects on growth Potamopyrgus antipodarum Gasteropoda Reduced >0.3 g cm−2 <63 µm (Broekhuizen et al., growth 2001) Physa integra Gasteropoda Reduced 0.06 g cm−2 day−1 <63 µm (Kent and Stelzer, 2008) growth Deleatidium Ephemeroptera No effect 20 000 NTU <0.05 mm (Suren et al., 2005) Zephlebia 22 400 mg L−1 Polyplectropus Trichoptera Triplectides Xanthocnemis zealandica Odonata Paranephrops planiﬁrons Decapoda Rhithrogena semicolorata Ephemeroptera Increased drift 250–2000 mg L−1 <63 µm (Molinos and Donohue, Baetis rhodani No effect on 2009) survival Asellus aquaticus Amphipoda No effect >2000 mg L−1 <63 µm (Molinos and Donohue, 2009) Glossosoma boltoni Trichoptera No effect on survival Hydropsyche betteni Trichoptera Reduced 23 NTU <0.6 mm (Strand and Merritt, survival 773 mg L−1 1997) No effect on growth Reymondia horei Gasteropoda Reduced survival 100 cm3 L−1 63–212 µm (Donohue and Irvine Ostracoda 2003) Ceratopsyche sparna Trichoptera No effect on 23 NTU <0.6 mm (Strand and Merritt, growth or 773 mg L−1 1997) survival Ischnura heterostricta Odonata Reduced feeding 1000‐1500 NTU Kaolin (Kefford et al., 2010) Ischnura aurora Odonata Increased survival 1000 NTU Kaolin (Kefford et al., 2010) Physa acuta Gasteropoda Reduced egg 5 mm depth Kaolin (Kefford et al., 2010) Gyraulus tasmanica hatching Chironomus cloacalis Diptera (Continues)

Table I. (Continued)

Species Effect Sediment load Sediment type Author

FIELD SCALE EXPERIMENTAL MANIPULATIONS

Paraleptophlebia Ephemeroptera Increased drift Depositing (Culp et al., 1986) sediment Increased drift Saltating (Culp et al., 1986) sediment Baetis rhodani Ephemeroptera Increased drift 4–5kgm−2 Sand (Larsen and Ormerod, Baetis muticus 2010) Ecdyonurus spp. Simulidae Diptera Increased drift 133 mg L−1 (Doeg and Milledge, Chironomidae 1991) Helodidae Coleoptera Diptera Increased drift 2.95–14 mg L−1 (Rosenberg and Wiens, Plecoptera 1978) Ephemeroptera Increased drift 550–700 kg 2 mm (mean) (Matthaei et al., 2006) Paracalliope ﬂuviatilis Amphipoda Increased drift (Suren and Jowett, Oxyethira albiceps Trichoptera 2001) Hydrobiosis sp. Trichoptera Chironomidae Diptera EPT Reduced 5 mm depth Volcanic ash (Brusven and Hornig, colonization 1984) Baetis rhodani Ephemeroptera Reduced 4–5kgm−2 Sand (Larsen and Ormerod, Ecdyonurus spp. density 2010) Leuctra hippopus Plecoptera Leuctra moselyi EPT Reduced 40% <2 mm (Kaller et al., 2001) richness All taxa Reduced 550–700 kg 2 mm (mean) (Matthaei et al., 2006) EPT richness All taxa Reduced 0–30% <2 mm (Angradi, 1999) EPT density Paraleptophlebia Ephemeroptera Chironominae Diptera Baetidae Ephemeroptera Increased 0–30% <2 mm (Angradi, 1999) Orthocladinae Diptera relative abundance Physa integra Gasteropoda Reduced 0.37 g cm−2 day−1 <63 µm (Kent and Stelzer, 2008) growth Bivalvia Mortality 6–25 mm depth Silt (Ellis, 1936)

CASE STUDIES

Oligochaeta No effect 45% 0.2–0.02 (Hamilton, 1961) Plecoptera mm, 45% 0.02– Ephemeroptera 0.002 mm, 10% Trichoptera <0.002 mm Diptera Mollusca All Taxa Reduced 8–177 mg L−1 <1 µm (Quinn et al., 1992) density 1.3–8.2 NTU All Taxa Reduced 62 mg L−1 (Wagener and LaPerriere, density 1985) All Taxa Reduced <500 mg L−1 (Cline et al., 1982) richness and density

(Continues)

Table I. (Continued)

Species Effect Sediment load Sediment type Author

All Taxa Reduced richness 4610 mg L−1 (Doeg and Koehn, 1994) and density All Taxa Reduced richness (Gray, 2004) and density All Taxa Reduced richness (Fossati et al., 2001) and density All Taxa Reduced richness (Milner and Piorkowski, and density 2004) All Taxa Reduced richness (Blettler and Marchese, and density 2005) All Taxa Reduced richness (Ciutti et al., 2000) and density All Taxa Reduced richness 4.7 g L−1 (Crosa et al., 2010) and density All Taxa Reduced richness (Ehrhart et al., 2002) Glossiphonia complanata Hirudinea Reduced density 22–336 mg L−1 (Extence, 1978) Erpobdella octoculata Ancylus ﬂuviatilis Gasteropoda EPT Reduced richness 10% more cover <2 mm (Fritz et al., 1999) Chironomidae Diptera Reduced density 300 mg L−1 (Gray and Ward, 1982) Cladocera Reduced survival 82–392 mg L−1 (Robertson, 1957) Reduced index Mean increase (Chen et al., 2009) score 0.36 NTU 0.41 mg L−1 Reduced density, 3.52–131.4 NTU Gold mine (Yule et al., 2010) richness, and 7–214 mg L−1 tailings combined food web with sewage complexity Polycelis felina Tricladida Loss or decline 10 000 m3 0.300–0.599 mm (Nuttall, 1972) Gammarus pulex Crustacea deposited in Leuctra nigra Plecoptera catchment Leuctra hippopus Leuctra geniculata Amphinemura sulcicollis Ephemera danica Ephemeroptera Beatis (Alainites) muticus Polycentropus kingi Trichoptera Sericostoma personatum Simuliidae Diptera Tubiﬁcidae Oligochaeta Increase 10 000 m3 0.300–0.599 mm (Nuttall, 1972) Rithrogena semicolorata Ephemeroptera Deposited Beatis rhodani in catchment

CORRELATION

EPT Reduced richness >0.8–0.9% <0.25 mm (Kaller and Hartman, of riffle substrate 2004) composition Ephemeroptera Reduced richness 30% cover Patch scale (Larsen et al., 2009) Plecoptera Trichoptera Oligochaeta Increased relative 30% cover Reach scale (Larsen et al., 2009) abundance

Antocha spp. Diptera Reduced density To 16% riffle Reach scale (Cover et al., 2008) Chironominae Diptera surface <4 mm Ecclisomyia spp. Trichoptera composition

(Continues)

Table I. (Continued)

Species Effect Sediment load Sediment type Author

Attenella delantala Ephemoptera Increased density To 16% riffle Reach scale (Cover et al., 2008) Zapada columbiana Plecoptera surface <4 mm composition Chironomidae (certain spp.) Diptera Increased density 0% to 100% % bed (Zweig and Rabeni, Oligochaeta composition 2001) Patch scale Dicranota spp., Diptera Reduced density % riffle surface Patch scale (Cover et al., 2008) Drunella doddsi Ephemeroptera composition <4 mm

Epeorus spp. All taxa Reduced density % riffle surface <2 mm (Angradi, 1999) EPT composition Attenella delantala Ephemeroptera Increased relative % riffle surface Patch scale (Cover et al., 2008) Oligochaeta abundance composition <4 mm Orthocladinae Diptera Increased relative % riffle surface <2 mm (Angradi, 1999) abundance composition Baetidae Ephemeroptera Reduced relative % riffle surface <2 mm (Angradi, 1999) abundance composition EPT Reduced richness ≥100 mL Pumped (Herbst and Kane, sediment volume 2006) Oligochaeta Increased density ≥100 mL Pumped (Herbst and Kane, Chironomidae Diptera sediment volume 2006) EPT Reduced density −6.4 to −0.3 phi Bed particle size (Roy et al., 2003b) All taxa and richness Taxa on riffles Reduced richness −6.4 to −0.3 phi Bed particle size (Roy et al., 2003a) Taxa on banks No effect on −6.4 to −0.3 phi Bed particle size (Roy et al., 2003a) density or richness All taxa Reduced density % bed (Quinn and Hickey, and richness composition 1990) All taxa Reduced density 0% to 100% % bed (Zweig and Rabeni, EPT and richness composition 2001) Patch scale EPT Reduced density % bed (Niyogi et al., 2007) and richness composition Ecclisomyia sp. Trichoptera Reduced relative 7.3% % bed surface (Bryce et al. 2010) Pteronarcys sp. Plecoptera abundance 8.2% composition Oligophlebodes sp. Trichoptera 8.8% Epeorus grandis Ephemeroptera 9.1% Arctopsyche grandis Trichoptera 10.2% Epeorus longimanus Ephemeroptera 11.4% Megarcys sp. Plecoptera 11.4% Caudatella hysterix Ephemeroptera 12.3% − Polycelis felina Tricladida Reduced relative 6–35 231 mg L 1 China clay works (Nuttall and Bielby, Potamopyrgus jenkinsi Gasteropoda abundance 1973) Leuctra fusca Plecoptera Ephemerella ignata Ephemeroptera Rhithrogena semicolorata Sericostoma personatum Trichoptera Limnephilidae Hydropsyche instabilis Dicranota sp. Diptera Simuliidae Naididae Oligochaeta Increased relative 6–35 231 mg L−1 China clay works (Nuttall and Bielby, Tubiﬁcidae abundance 1973) Perlodes microcephala Plecoptera Beatis rhodani Ephemeroptera (Continues)

Table I. (Continued)

Species Effect Sediment load Sediment type Author

Chironomidae Diptera Oligochaeta Increased density Strongly (Quinn et al., 1997b) Chironomidae Diptera associated with Potamopyrgus Gasteropoda differences in Oxyethira Trichoptera land use

EPT, Ephemeroptera, Plecoptera and Trichoptera; the number of taxa in these families are commonly used to assess impacts on invertebrate community. NTU, nephelometric turbidity units.

(see chemical effects section), the changes in the physical conditions, the invertebrate community becomes more structure of the bed also have a more direct impact on the vulnerable to physical disturbance of the bed during flood animals that live there. Most invertebrate species have specific events [or any event that depletes the fauna such as pollution or requirements of the substrate they live in and tend to avoid low flow (Dunbar et al., 2010)]: recolonization of denuded patches that fail to meet these requirements (Culp et al.,1983; patches takes longer when motile species are lacking from the Peckarsky, 1991; Williams and Smith, 1996; Sarriquet et al., community (Gjerløv et al., 2003). 2007). Blackfly larvae tether themselves by means of a posterior circlet of hooks onto strands of silk which they attach to comparatively clean substrate, and they avoid substrates covered by a surface drape of loose sediments (Bass, 1998). CHEMICAL EFFECTS OF INCREASED Several species of crawling mayflylarvaewillavoidfiner, less FINE SEDIMENT LOADING stable substrates as they cannot grip them effectively Oxygen concentration (Ciborowski et al., 1977; Corkum et al.,1977).Otherspecies, for example, certain Chironomidae and Ephemeridae, select The deposition of fine sediments on river beds is usually finer sediments into which they build tunnels. However, few associated with profound changes to the chemical environment. taxa are found on unstable deposits of sand where erosion and Reduced percolation of water through the substrate results in abrasion make it difficult for invertebrates to hold station the establishment of more pronounced gradients of oxygen and (Culp et al., 1986; Armitage and Cannan, 2000). other dissolved substances (Pretty et al., 2006). Where However, invertebrates do not just live on the surface of deposited material has a high organic content, microbial the river bed; both macrofauna (>500 µm) and meiofauna activity can lead to oxygen depletion and a build up of (42–500 µm) penetrate the substrate to some depth (Swan and potentially toxic substances such as ammonium, ferrous and Palmer, 2000; Stead et al., 2004) by either burrowing or manganous ions. Although the increased supply of particulate working through the interstices between larger particles. In organic matter may benefit some species of invertebrates part, invertebrates use the deeper layers of the substrate as (Welton and Clarke, 1980; Lemly, 1982; Arruda et al., 1983; refugia to hide from risks more apparent on the substrate Hart, 1992; Jackson et al., 2007), many species are sensitive to surface, such as high flow and predators (Lancaster et al., 1991; the oxygen depletion, and associated chemical changes, that Lancaster, 1996, Swan and Palmer, 2000). Fine sediments accompany such increases [e.g. M. margaritifera (Geist and may increase the embeddedness of stones, thus reducing Auerswald, 2007)]. Dependent upon their specific oxygen erosion during high‐flow events and creating areas of low requirements, this chemical change restricts the depth to which shear stress. However, as interstices fill with fine sediments, invertebrates can penetrate into the deposited sediment (Swan the ability of invertebrates to penetrate the river bed by and Palmer, 2000; Stead et al., 2003). For example, the depth to crawling between larger particles becomes reduced, affecting which hexagenid mayflies borrow into soft sediments is invertebrate distribution (McClelland and Brusven, 1980) and strongly correlated with oxygen depletion and the deposition of the availability of refugia (Lancaster and Hildrew, 1993): these organic matter (Rasmussen, 1988; Krieger et al., 2007), interstices (and other areas of low shear stress) are used by indicating that depth penetration is affected by the chemical invertebrates as refugia to avoid ‘wash‐out’ during high‐flow environment as well as the physical changes detailed above. events (Lancaster and Hildrew, 1993). The availability of flow Nevertheless, invertebrates can encourage remobilization and refugia on the river bed has a significant influence on oxygenation of sediments by bioturbation and burrowing community composition, with motile species lacking where (Nogaro et al., 2006; Zimmerman and de Szalay, 2007; Nogaro refugia are scarce (Gjerløv et al., 2003). Under such et al., 2009). The changes in invertebrate community that

Copyright © 2011 John Wiley & Sons, Ltd. River Res. Applic. (2011) DOI: 10.1002/rra FINE SEDIMENT AND MACRO‐INVERTEBRATES accompany sewage pollution, primarily acting through oxygen food available to filter‐feeding invertebrates. Where con- depletion, are well documented (Hynes, 1960) and result in a centrations of particulate organic matter are increased above predictable loss of taxa from the community (Jones et al., intrinsic levels, there is the potential for populations of filter 2010): it is likely that similar changes will occur with feeders to expand. However, the overall quality of increasing loads of fine particulate organic matter from other particulate food resources may decline if the concentration key catchment sources releasing organic material including of particulate inorganic matter increases disproportionately, farm steadings, damaged road verges and riparian areas influencing ingestion rate (as previously discussed; Gaugler (Collins et al., 2009). and Molloy, 1980). Deposit feeders can also benefitfrom increased retention of particulate organic material on the river bed, although again increases in quantity can be counteracted by declines in nutritional quality where INDIRECT EFFECTS OF INCREASED inorganic matter comprises a large proportion of the FINE SEDIMENT LOADING available resource (Nuttall and Bielby, 1973). Similarly, the nutritional quality of periphyton (the film of attached Habitat availability algae, fungi, bacteria, organic matter and sedimented As well as the direct effects of fine sediment upon the material found on the surface of stones) can decline as invertebrates, there are a number of consequences of increased the proportion of inorganic material in the layer increases. fine sediment loading that impact upon the invertebrate Further declines in periphyton quality can occur because of community via indirect pathways. Many aspects of river the effect of turbidity on algal growth (Parkhill and ecosystems are altered as a consequence of increased loading Gulliver, 2002); suspended sediment reduces light penetra- of fine sediment: of paramount importance are changes to the tion to the river bed, resulting in less algal growth and habitats available to invertebrates, particularly, changes in therefore poorer nutritional quality of periphyton (Quinn substrate particle size and changes in the macrophyte et al., 1992, 1997a). The decline in periphyton food quality community. There is a strong relationship between substrate may have severe implications for scraping invertebrates that composition and invertebrate distribution at the patch scale depend upon this resource, such as snails. The egestion rate (Culp et al., 1983), and invertebrate community is strongly of the gasteropod Physa integra Haldeman increased with correlated with mesoscale habitat patches (Pardo and experimental additions of silt, suggesting increased con- Armitage, 1997; Collier et al., 1998; Armitage and Cannan, sumption to compensate for reduced food quality (Kent and 2000; Buffagni et al.,2000)defined by water depth, substrate Stelzer 2008). Nevertheless, longer‐term experiments with composition and the presence and type of macrophytes (Kemp another gasteropod, Potamopyrgus antipodarum (J.E. Gray), et al., 2000). Furthermore, changes in habitat patches are often and the mayfly Deleatidium sp. indicate that high levels of the best descriptor of change in invertebrate community (Petts deposition impact the growth rate of grazers (Broekhuizen et al., 1993). Dependent upon the quantity and quality of the et al., 2001). Furthermore, changes occur in the availability of fine sediment entering the river channel, increased loadings periphyton because of fine sediment loading. The reduced result in either an increase or decrease in the abundance of light penetration with increased turbidity results in a reduction macrophytes, which in turn affect conveyance of sediment, in the proportion of the bed where algae can grow (Vermaat respectively increasing or reducing retention of fine particulate and De Bruyne, 1993; Köhler et al., 2010). Macrophytes shade material on the river bed (see Jones et al., in review). Species the river bed, restricting the growth of benthic algae beneath composition of macrophytes is likely to change with increased them (Sand‐Jensen et al.,1989);fine sediments interact loading, tending towards more rapidly growing and/or strongly with macrophytes (Jones et al., in review), and any emergent species (Wood and Armitage, 1997). Areas of low change in the abundance of macrophytes as a consequence of flow are created by plant stands and fine organic rich fine sediment loading will affect the availability of benthic sediments accrue, altering conditions for sessile organisms algae. However, macrophytes present new substrate for such as bivalves (Fritz et al., 2004). All these changes in periphyton high in the water column, and any losses of habitat availability will have important consequences for the benthic algal food resources can be counteracted by an invertebrate community present at a site. increase in the availability of periphyton growing on plant surfaces, which can be exploited by grazing invertebrates, both generalist and specialized for feeding on this resource (Diehl Food availability and quality and Kornijów, 1998). Furthermore, the macrophytes them- Dependent upon the key sources of fine sediment, selves are exploited by invertebrates (Sand‐Jensen and loadings to rivers can contain substantial quantities of Madsen, 1989) and can offer nutritionally better quality food organic matter, influencing the concentration of particulate than the terrestrial leaf detritus that supports many river organic matter suspended in the water and therefore the communities (Sand‐Jensen, 1997). These changes to the

Copyright © 2011 John Wiley & Sons, Ltd. River Res. Applic. (2011) DOI: 10.1002/rra J. I. JONES ET AL. quality, availability and source of food resources will SCALES OF RESPONSE OF INVERTEBRATES TO inevitably have implications for the invertebrate community INCREASED FINE SEDIMENT LOADING present at a site. Invertebrates display different scales of response to increased fine sediment loading. The most immediate Food web changes response to an increase in the concentration of suspended Fine sediments can have a profound impact upon fish fine sediment is an increase in the number of animals populations, particularly a direct impact on egg survival and entering the drift (Ciborowski et al., 1977; White and subsequent recruitment (Waters, 1995). By restricting the Gammon, 1977; Rosenberg and Wiens, 1978; Doeg and supply of oxygen to eggs, either buried in the river bed or Milledge, 1991; Suren and Jowett, 2001; Matthaei et al., broadcast, fine sediments can result in the death of fish eggs 2006; Larsen and Ormerod, 2010; see Table I), although (Wood and Armitage, 1997; Sear et al., 2008). Further, direct some authors have suggested that the effects of increased (e.g. clogging of gills) and indirect (e.g. changes in habitat) discharge, often associated with increased sediment loads, impacts on fish populations occur as a result of increased have a greater impact on drift densities than sediment load loading of fine sediment (for a full review, see Kemp et al., per se (Bond and Downes, 2003). Increased drift may in 2011). Most species of Unionid bivalve are parasitic on fish in part be due to individuals becoming dislodged by moving their early life stages (often with high specificity), and fine particles (Culp et al., 1986), but there appears to be an sediments may impact their populations because of the active behavioural component also with individuals actively importance of fish for recruitment (Box and Mossa, 1999). As using drift to avoid the negative impacts (e.g. burial) of invertebrates comprise the main food resource of many increased concentrations of suspended sediment (Ciborowski species of fish, invertebrates are released from predation et al., 1977). The sudden increase in densities of drifting where fish populations decline as a consequence of increased invertebrates can result in a depletion of the standing stock in fine sediment loading. Even where fish populations are not the benthos, particularly of motile taxa, resulting in a change in impacted, increased turbidity can reduce the visibility of abundance and composition of the remaining community invertebrate prey to fish, thus reducing predation risk (Doeg and Milledge, 1991; Suren and Jowett, 2001; Matthaei (Gardener, 1981; Berg and Northcote, 1985; Zamor and et al., 2006; Larsen and Ormerod, 2010). Notwithstanding the Grossman, 2007). The implications for the invertebrate apparent effects on the benthos, such a behavioural response community will depend on the extent to which predation can lead to increased survival of individuals and rapid controls population growth; in unproductive or frequently recolonization of a site after a sediment pulse has passed. disturbed conditions, change in fish density has little impact Direct and indirect impacts of fine sediment can manifest on invertebrates (Peckarsky et al., 2008), whereas fish can on the growth rate (e.g. Broekhuizen et al., 2001; Kent and exert significant control on invertebrate populations in more Stelzer, 2008) and mortality (e.g. Ellis, 1936) of individuals, productive environments (Jones and Jeppesen, 2007). Many which can be detected through reduced population growth fish species are visual predators and select larger prey items, rate (e.g. Broekhuizen et al., 2001). As individuals succumb thus having a more intense impact on larger, typically to the direct negative impacts of increased sediment loads, predatory invertebrates, altering invertebrate community changes in the abundance of individual species will composition and in turn impacting upon the taxa these manifest, which in turn alter community composition (e.g. invertebrates feed upon. Correspondingly, if the impacts of Fossati et al., 2001; Ehrhart et al., 2002; Freeman and fine sediment lead to the loss of prey species or basal Schorr, 2004). Similarly, changes in community compo- resources from the community, those invertebrates that are sition can occur as changes in the available habitat favour reliant on these resources will have to alter their diet or in turn different species (Armitage and Cannan, 2000; Hutchens be lost from the community. In the one study that has et al., 2009). Nevertheless, the scaling up of impacts from investigated the impact of fine sediments on food web the patch to the whole river reach is challenging and fraught structure, inputs from an alluvial gold mine (combined with with difficulties. Relationships between deposited fine sewage from the mining village) in Indonesia were associated sediment and invertebrate community that are observed at with a reduction in the number of elements, links, linkage the patch/mesohabitat scale are often not obvious when density and complexity of the food web of polluted sites samples are collected at the whole reach scale (Larsen (Yule et al., 2010). At the most impacted sites, several et al., 2009), where the community can be composed of functional types were lost, indicating that certain resources animals from both depositional and erosional patches. In were no longer contributing to the food web (Yule et al., part, this may be due to difficulties of quantifying sediment 2010). Although such community‐level changes may be stress at the larger scale, but there are clear issues regarding profound, discriminating these indirect effects from the other the most appropriate scale to use to detect impacts on impacts of fine sediment is extremely difficult. macro‐invertebrates.

SCALES OF DETECTION OF EFFECTS OF INCREASED Field‐scale experimental manipulations FINE SEDIMENT LOADING Experimental channels that approximate to the field scale, Current targets for fine sediments in rivers are commonly particularly flow‐through channels fed by river water where based on concentrations of suspended solids (Collins and natural colonization by a range of organisms is possible, have Anthony, 2008), yet the impacts on invertebrates are largely been used to emulate natural conditions and provide more driven by deposition of material (as discussed earlier) which is, realistic tests of the impact of fine sediment on macro‐ in part, a consequence of in‐stream conditions (Hutchens et al., invertebrates (Connolly and Pearson, 2007) and meiofauna 2009) and hence the experimental conditions that are used to (Radwell and Brown, 2006). Such flow‐through artificial determine the impact. Hence, the design of investigations used channels enable some behavioural responses to fine sediment to assess the impact of fine sediment on macro‐invertebrates is loading to be expressed and quantified, such as drift and paramount, not only in terms of the response of the animals but recolonization by invertebrates (Suren and Jowett, 2001). of the manifestation of impacts. Objective quantitative There are logistic constraints on the number of treatment information on the sensitivity of river invertebrates to concentrations that can be tested; large‐scale, typically outdoor increased fine sediment loading can be derived using different channels require considerable manpower to maintain, and the approaches. The constraints on the general applicability of the number of replicate channels available constrains experimen- information are dependent upon the approach used. Typically, tal designs (Lamberti and Steinman, 1993; Belanger, 1997). A the approaches used to assess the response of biota to further constraint arises from the scarcity of such large‐scale pollutants, including sediment, are the following: experimental facilities; it is only possible to use such facilities (1) laboratory assessments, to test the impact on the few river types (and hence water (2) field‐scale experimental manipulations (experimental chemistry and community types) where the facilities are sited. channels and simulated events), Hence, the impacts predicted using such channels may not be (3) case studies of pollution events and applicable to wider, more distant sites and their species. (4) correlation of field survey data. Nevertheless, studies using such facilities can provide valuable information at the population or community level The approaches differ in the degree of experimental (see Table I). Experiments detailing the impact of loadings of fi control possible (decreasing from 1 to 4) and in type of ne sediment have illustrated the effect of added sediment on response, scale and general applicability to the natural envi- growth and survival (e.g. Ellis, 1936; Kent and Stelzer, 2008; ronment (increasing from 1 to 4). Any information derived Molinos and Donohue, 2009), drift (Doeg and Milledge, 1991; from these different sources should be interpreted with these Suren and Jowett, 2001; Molinos and Donohue, 2009), constraints in mind. Details of the response of invertebrates to burrowing (Doeg and Milledge, 1991; Zimmerman and fine sediment investigated at various scales are given in de Szalay, 2007), invertebrate community (McClelland and Table I. Brusven, 1980; Bond and Downes, 2003; Connolly and Pearson, 2007) and growth and production of basal resources (Parkhill and Gulliver, 2002). Experiments have Laboratory assessments been conducted in lakes (Donohue et al., 2003; Donohue and The sensitivity of animals to pollutants is typically derived Irvine, 2004), where experimental enclosures can be estab- through assessments of chronic, or incipient, toxicity in lished relatively easily, but extending results to riverine continuous exposure tests that establish the concentration invertebrate communities and conditions is not valid. that causes mortality of 50% of the test organisms (lethal An alternative approach is to simulate pulsed events concentration 50 or LC50) conducted under controlled by experimental dosing of real rivers with sediment. conditions. Although clearly of some use in predicting the Here, sites, chosen carefully so that any damage does not biological impact of continuous discharges of toxic pollut- influence substantial lengths of river downstream, are ants, the relevance of such techniques to the impact of fine dosed with sediment, and the consequence followed. Such sediment on invertebrates, where effects are rarely through controlled dosing experiments provide the most appropriate direct toxicity, is limited. Nevertheless, pertinent experiments information for assessing the impact of sediment pollution have been conducted on the ability of invertebrate taxa to on rivers. However, the logistics involved in undertaking withstand burial by fine sediments of differing particle size such large‐scale manipulations are such that they have not (Wood et al., 2001; Wood et al., 2005) and on the impact of been undertaken that often. Hence, only a few river types fine sediment on feeding rate (Kurtak, 1978; Gaugler and and certain types of sediment pollution have been Molloy, 1980; Hart, 1992; Broekhuizen et al., 2001; Kent investigated (see Table I). Experiments in the upper Usk and Stelzer, 2008) and survival (Donohue and Irvine, 2003) (Wales, UK) have shown that experimental addition of (see Table I). sand resulted in increased drift, particularly of mayflies

(B. rhodani, B. muticus and Ecdyonurus spp.), blackflies, Correlation chironomids and helodid beetles, with a consequent reduction in the benthic densities (Larsen and Ormerod, Wide applicability across river types can be achieved 2010). Reductions in invertebrate taxon richness and using correlation techniques to assess relationships between richness of Ephemeroptera, Plecoptera and Trichoptera large‐scale biological and chemical sampling data. Here, were observed in New Zealand streams in a more data on the occurrence of biota and the sediment conditions comprehensive study, where naturally deposited river of those sites are analysed together. Although any analyses sediment was experimentally added to replicated streams are constrained by the quality of the data, by using a large of different land‐use type (Matthaei et al., 2006). number of samples collected at a scale where realistic responses are possible, the likelihood of an accurate description of the sensitivity of biota to fine sediment can Case studies be increased. Such techniques have shown a relationship Most case studies report the impact of increased sediment between small (patch)‐scale measures of invertebrate loading from episodic events, usually construction or mining community and deposited material, indicating that the local works (Nuttall and Bielby, 1973; Extence, 1978; Cline et al., distribution of Ephemeroptera, Plecoptera and Trichoptera 1982; Stout and Coburn, 1989; Ehrhart et al., 2002; Lane and is influenced by substrate composition, but have failed to Sheridan, 2002; Milner and Piorkowski, 2004; Blettler and indicate more than weak relationships, particularly with the Marchese, 2005; Kreutzweiser et al., 2005; Hedrick et al., abundance of Oligochaeta and Chironomidae, at the more 2007; Levesque and Dube, 2007; Chen et al., 2009), or realistic reach scale (Angradi, 1999; Herbst and Kane, 2006; continuous activities, such as the impact of livestock (Fritz Cover et al., 2008; Larsen et al., 2009; see Table I). Some et al., 1999; Braccia and Voshell, 2006; Braccia and Voshell, contradictory results have been found between experiments 2007), military practices (Quist et al., 2003; Maloney et al., and correlation with field data (increased/reduced relative 2005; Williams et al.,2005;Bhatet al., 2006; Maloney and abundance of Baetidae; Angradi, 1999, see Table I). It has Feminella, 2006) and the confluence of rivers (Svendsen et al., long been known that macro‐invertebrate community 2009). Because of the often diffuse nature of sediment composition is strongly correlated with river‐bed surface pollution events, there are relatively few case studies reporting particle size composition (e.g. Moss et al. 1987) and such their impact (for summary, see Table I). Unless there is correlations have been used to identify taxa associated with continuous monitoring (e.g. using turbidity sensors), peaks a low proportion of fine sediment in the bed (Carlisle et al., of suspended sediment concentration are often missed 2007; Cormier et al. 2008; Bryce et al., 2010). However, at (Bilotta and Brazier, 2008) since peaks of fine sediment the reach scale, separating the effects of covariables associated concentration often occur as a consequence of rainfall events with natural variability between river reaches (topography, flushing material into neighbouring water courses (Smith geology, etc.) from those of fine sediment per se is difficult et al., 2003). The difficulties of accurate quantification of (Hutchens et al., 2009), and the physical characteristics of the consequences of pulsed events are compounded by a individual sites are likely to have a large bearing on results, frequent lack of biological information from both before particularly where the number of sites investigated is low. and after an event. As a consequence, it is rarely possible to Hence, such correlation approaches are not capable of draw accurate conclusions from case studies (Angermeier determining whether taxa are responding to increased loads et al., 2004). Situations where continuously elevated con- of fine sediment per se or some other feature of river reach type centrations of sediment occur, often associated with mining associated with the deposition of fine sediment (e.g. slope, activity, can provide some information regarding chronic velocity), and use of correlations between bed surface impacts on the invertebrate community (e.g. Fritz et al., 1999; composition and macro‐invertebrate taxa to set targets will Glozier et al., 2002; Williams et al., 2005; Svendsen et al., tend to produce targets for all river types based on conditions 2009) but may not reflect the impact of pulsed events where found at erosional sites. Although not used to investigate the there is the potential for recovery between episodes. impact of fine sediment on macro‐invertebrates, weighted Nevertheless, certain taxa do seem to be negatively impacted averaging provides a technique where the influence of the by increased loads of fine sediment, particularly certain characteristics of the site are partialled out (i.e. their influence Ephemeroptera, Plecoptera and Trichoptera, whereas others, removed), leaving only the effect of the fine sediment. particularly certain Diptera and Oligochaeta, are positively Although the use of any technique that relies on field‐derived impacted. An increase in Chironomidae and decrease in data is inherently problematic, when combined with pertinent the proportion of scrapers was noted over 3 years when measures of fine sediment loading, the weighted averaging benthic inorganic sediment increased from 800 g m−2 to over approach is likely to provide the best description of the 5000gm−2 as a result of modification of a road crossing association between fine sediment and macro‐invertebrates. (Kreutzweiser et al.,2005). As correlation does not provide proof of cause–effect, any

Copyright © 2011 John Wiley & Sons, Ltd. River Res. Applic. (2011) DOI: 10.1002/rra FINE SEDIMENT AND MACRO‐INVERTEBRATES relationships derived using such techniques will require types, before appropriate revised targets can be defined. In verification by experimental manipulation. addition, the development of revised targets for catchment sediment control should be based on the linkages between fi TARGETS ne sediment pressures and other forms of freshwater biota such as fish. There is considerable evidence to suggest that increased fi loading of ne sediment has both direct and indirect impacts ACKNOWLEDGEMENT on macro‐invertebrates (Figure 1). However, quantifying these impacts is fraught with difficulty; both a realistic The authors gratefully acknowledge the funding provided measure of the extent of the stress (i.e. the additional fine by the Department for Environment, Food and Rural sediment loading above the intrinsic rate) and an appropriate Affairs (Defra, contract WQ0128; Extending the evidence scale of response (i.e. at a scale that allows a community‐level base on the ecological impacts of fine sediment and response) are required. Furthermore, the extent of impact developing a framework for targeting mitigation of appears to be dependent upon context (Hutchens et al.,2009); agricultural sediment losses). the sensitivity of biological communities is largely dependent upon the unimpacted condition, and a blanket, single target for all river types is unlikely to provide adequate protection REFERENCES to vulnerable taxa in sensitive river types. Yet, targets for fine sediment loading are required as part of cost‐effective Alabaster JS, Lloyd DS. 1982. Finely divided solids. In Water Quality ‐ Criteria for Freshwater Fish, Alabaster JS, Lloyd DS (eds). Butterworth; management of river basins: key components of the macro London; 1–20. invertebrate community of rivers appear likely to be impacted, Aldridge DW, Payne BS, Miller AC. 1987. The effects of intermittent resulting in many rivers failing to achieve ‘good’ ecological exposure to suspended solids and turbulence on three species of status under current conditions (Collins and Anthony, 2008; freshwater mussels. Environmental Pollution 45:17–28. Collins et al., 2009). Management of other stressors in Angermeier PL, Wheeler AP, Rosenberger AE. 2004. A conceptual framework for assessing impacts of roads on aquatic biota. Fisheries 29: isolation is unlikely to result in the required improvement 19–29. (Mainstone and Parr, 2002; Mainstone et al., 2008; Collins Angradi TR. 1999. Fine sediment and macroinvertebrate assemblages in et al., 2009). In order to be defensible, any targets set must be Appalachian streams: a field experiment with biomonitoring applica- related to a demonstrable impact and be defined in tions. Journal of the North American Benthological Society 18:49–66. determinands that are both quantifiable and relevant. Many Armitage PD, Blackburn JH. 2001. The macroinvertebrate fauna of fi ‐ the Holy Stream, a small tributary of the River Frome, Dorset. of the impacts of ne sediment on macro invertebrates appear Proceedings of the Dorset Natural History and Archeological Society related to the deposition of material, which is in part dependent 123:95– 100. on local hydraulic and hydrological conditions, yet current Armitage PD, Blackburn JH, Wiggers R, Harris I (1999) The environmental targets are frequently defined in terms of suspended sediment quality of the Bovington Stream and its effects on the adjacent River Frome (Dorset), assessed with macroinvertebrate data. Proceedings of the Dorset concentration. Translating concentrations of total suspended 121 – fi Natural History and Archeological Society :123 128. sediments into quanti able biological impact is a necessary Armitage PD, Cannan CE. 2000. Annual changes in summer patterns of step if such regularly monitored water‐quality measures are to mesohabitat distribution and associated macroinvertebrate assemblages. be used to help inform the management of river basins. The Hydrological Processes 14: 3161–3179. evidence available describing the impact of fine sediment on Arruda JA. 1983. Daphnid filtering comb area and the control of filtering macro‐invertebrate communities, although demonstrating rate. Journal of Freshwater Ecology 2: 219–224. fi fi Arruda JA, Marzolf GR, Faulk RT. 1983. The role of suspended sediments clear cause for concern, is insuf cient to de ne quantitative in the nutrition of zooplankton in turbid reservoirs. Ecology 64:1225–1235. targets that are appropriate across a range of river types. Most Barko JW, Gunnison D, Carpenter SR. 1991. Sediment interactions with studies have shown impacts at the patch (mesohabitat) scale or submersed macrophyte growth and community dynamics. Aquatic Botany smaller, and scaling these up to an appropriate scale for the 41:41–65. management of river basins is challenging (Larsen et al., Bass JAB. 1998. Last‐Instar Larvae and Pupae of the Simuliidae of Britain 2009), yet it is at this larger scale that managers need to work, and Ireland: A Key with Brief Ecological Notes. FBA: Ambleside. Belanger SE. 1997. Literature review and analysis of biological complexity given the focus of current water policy including the European in model stream ecosystems: influence of size and experimental design. Union Water Framework Directive (European Parliament, Ecotoxicology and Environmental Safety 36:1–16. 2000). Setting targets based on current information, predom- Berg L, Northcote TG. 1985. Changes in territorial, gill‐flaring, and feeding inantly from small‐scale studies, is likely to lead to overly behaviour in juvenile coho salmon (Oncorhynchus kisutch) following ‐ prescriptive definitions and reduce the cost effectiveness of short term pulses of suspended sediment. Canadian Journal of Fisheries and Aquatic Sciences 42: 1410–1417. river basin management options focusing upon sediment Bhat S, Jacobs JM, Hatfield K, Prenger J. 2006. Relationships between control. New analysis is clearly required, using large‐scale stream water chemistry and military land use in forested watersheds in data covering community‐level response over a range of river Fort Benning, Georgia. Ecological Indicators 6: 458–466.

Bilotta GS, Brazier RE. 2008. Understanding the influence of suspended artificial stream channels and recirculating mesocosms. Hydrobiologia solids on water quality and aquatic biota. Water Research 42: 2849– 2861. 592:423–438. Blettler MCM, Marchese MR. 2005. Effects of bridge construction on Cooper DM, Naden P, Old G, Laize C. 2008. Development of guideline the benthic invertebrates structure in the Parana River Delta. Interciencia sediment targets to support management of sediment inputs into aquatic 30:60–66. systems. Natural England Research Report NERR008. Natural England: Bond NR, Downes BJ. 2003. The independent and interactive effects of Sheffield. fine sediment and flow on benthic invertebrate communities character- Corkum LD, Pointing PJ, Ciborowski JJH. 1977. Influence of current istic of small upland streams. Freshwater Biology 48: 455–465. velocity and substrate on distribution and drift of 2 species of mayflies Box JB, Mossa J. 1999. Sediment, land use, and freshwater mussels: (Ephemeroptera). Canadian Journal of Zoology 55: 1970–1977. prospects and problems. Journal of the North American Benthological Cormier SM, Paul JF, Spehar RL, Shaw‐Allen P, Berry WJ, Suter GW. 2008. Society 18:99–117. Using field data and weight of evidence to develop water quality criteria. Braccia A, Voshell JR. 2006. Benthic macroinvertebrate fauna in small streams Integrated Environmental Assessment and Management 4: 490–504. used by cattle in the Blue Ridge Mountains, Virginia. Northeastern Cover MR, May CL, Dietrich WE, Resh VH. 2008. Quantitative linkages Naturalist 13:269–286. among sediment supply, streambed fine sediment, and benthic macro- Braccia A, Voshell JR. 2007. Benthic macroinvertebrate responses to invertebrates in northern California streams. Journal of the North increasing levels of cattle grazing in Blue Ridge Mountain streams, American Benthological Society 27: 135–149. Virginia, USA. Environmental Monitoring and Assessment 131:185– 200. Crosa G, Castelli E, Gentili G, Espa P. 2010. Effects of suspended Broekhuizen N, Parkyn S, Miller D. 2001. Fine sediment effects on feeding sediments from reservoir flushing on fish and macroinvertebrates in an and growth in the invertebrate grazers Potamopyrgus antipodarum alpine stream. Aquatic Sciences 72:85–95. (Gastropoda, Hydrobiidae) and Deleatidium sp (Ephemeroptera, Ciutti F, Cappelletti C, Monauni C, Pozzi S. 2000. Effetti dello svaso di un Leptophlebiidae). Hydrobiologia 457: 125–132. bacino idroelettrico sulla comunità di macroinvertebrati. (Effects of Brusven MA, Hornig CE. 1984. Effects of suspended and deposited controlled emptying of a hydroelectric reservoir on the macroinverte- volcanic ash on survival and behavior of stream insects. Journal of the brate community: in Italian). Rivista di Idrobiologia 39: 165–184. Kansas Entomological Society 57:55–62. Culp JM, Walde SJ, Davies RW. 1983. Relative importance of substrate Bryce SA, Lomnicky GA, Kaufmann PR. 2010. Protecting sediment‐ particle‐size and detritus to stream benthic macroinvertebrate microdistribu- sensitive aquatic species in mountain streams through the application of tion. Canadian Journal of Fisheries and Aquatic Sciences 40:1568–1574. biologically based streambed sediment criteria. Journal of the North Culp JM, Wrona FJ, Davies RW. 1986. Response of stream benthos and American Benthological Society 29: 657–672. drift to fine sediment deposition versus transport. Canadian Journal of Buffagni A, Crosa GA, Harper DM, Kemp JL. 2000. Using Zoology 64: 1345–1351. macroinvertebrate species assemblages to identify river channel habitat Diehl S, Kornijów R. 1998. Influence of submerged macrophytes on trophic units: an application of the functional habitats concept to a large, interactions among fish and macroinvertebrates. In The Structuring Role of unpolluted Italian river (River Ticino, northern Italy). Hydrobiologia Submerged Macrophytes in Lakes. Jeppesen E, Søndergaard M, 435: 213–225. Søndergaard M, Christoffersen K (eds). Springer‐Verlag: New York; Carlisle DM, Meador MR, Mouton SR, Ruhl PM. 2007. Estimation and 24– 46. application of indicator values for common macroinvertebrate genera Dobson M, Poynter K, Cariss H. 2000. Case abandonment as a response to and families of the United States. Ecological Indicators 7:22–33. burial by Potamophylax cingulatus (Trichoptera: Limnephilidae) larvae. Chen YS, Viadero RC, Wei XC, Fortney R, Hedrick LB, Welsh SA, Aquatic Insects 22:99–107. Anderson JT, Lin LS. 2009. Effects of highway construction on stream Doeg TJ, Koehn JD. 1994. Effects of draining and desilting a small weir on water quality and macroinvertebrate condition in a Mid‐Atlantic Highlands downstream fish and macroinvertebrates. Regulated Rivers: Research & watershed, USA. Journal of Environmental Quality 38:1672–1682. Management 9: 263–277. Ciborowski JJH, Pointing PJ, Corkum LD. 1977. Effect of current velocity Doeg TJ, Milledge GA. 1991. Effect of experimentally increasing and sediment on drift of mayfly Ephemerella subvaria McDunnough. concentrations of suspended sediment on macroinvertebrate drift. Freshwater Biology 7: 567–572. Australian Journal of Marine and Freshwater Research 42:519–526. Cline LD, Short RA, Ward JV. 1982. The influence of highway construction Donohue I, Irvine K. 2003. Effects of sediment particle size composition on on the macroinvertebrates and epilithic algae of a high mountain stream. survivorship of benthic invertebrates from Lake Tanganyika, Africa. Hydrobiologia 96:149–159. Archiv für Hydrobiologie 157: 131–144. Collier KJ, Wilcock RJ, Meredith AS. 1998. Influence of substrate type and Donohue I, Irvine K. 2004. Size‐specific effects of increased sediment loads on physico‐chemical conditions on macroinvertebrate faunas and biotic gastropod communities in Lake Tanganyika, Africa. Hydrobiologia 522: indices of some lowland Waikato, New Zealand, streams. New Zealand 337–342. Journal of Marine and Freshwater Research 32:1–19. Donohue I, Verheyen E, Irvine K. 2003. In situ experiments on the effects Collins AL, Anthony SG. 2008. Assessing the likelihood of catchments of increased sediment loads on littoral rocky shore communities in Lake across England and Wales meeting ‘good ecological status’ due to Tanganyika, East Africa. Freshwater Biology 48: 1603–1616. sediment contributions from agricultural sources. Environmental Science Dunbar MJ, Pedersen ML, Cadman D, Extence C, Waddingham J, & Policy 11: 163–170. Chadd R, Larsen SE. 2010. River discharge and local‐scale physical Collins AL, Naden PS, Sear DA, Jones JI, Foster IDL, Morrow K. 2011. Water habitat influence macroinvertebrate LIFE scores. Freshwater Biology quality targets for catchment sediment management: international experi- 55: 226–242. ence and prospects. Hydological Processes. DOI: 10.1002/hyp.7965 Edington JM, Hildrew AG. 1995. Caseless Caddis Larvae of the British Collins AL, Mcgonigle DF, Evans R, Zhang Y, Duethmann D, Gooday R. Isles. FBA Scientific Publication No. 53: Ambleside. 2009. Emerging priorities in the management of diffuse pollution at Ehrhart BJ, Shannon RD, Jarrett AR. 2002. Effects of construction site catchment scale. International Journal of River Basin Management 7: sedimentation basins on receiving stream ecosystems. Transactions of 179–185. the ASAE 45: 675–680. Connolly NM, Pearson RG. 2007. The effect of fine sedimentation on tropical Ellis MM. 1936. Erosion silt as a factor in aquatic environments. Ecology stream macroinvertebrate assemblages: a comparison using flowthrough 17:29–42.

European Parliament. 2000. Parliament and Council Directive 2000/60/ Hutchens JJ, Schuldt JA, Richards C, Johnson LB, Host GE, Breneman EC. Establishing a framework for community action in the field of water DH. 2009. Multi‐scale mechanistic indicators of Midwestern USA policy.Official Journal PE‐CONS 3639/1/00 REV 1: European Union, stream macroinvertebrates. Ecological Indicators 9: 1138–1150. Brussels, Belgium. Hynes HBN. 1960. The Biology of Polluted Waters. Liverpool University Extence CA. 1978. Effects of motorway construction on an urban stream. Press: Liverpool. Environmental Pollution 17: 245–252. Iglesias JIP, Urrutia MB, Navarro E, Alvarezjorna P, Larretxea X, Bougrier S, Fossati O, Wasson JG, Hery C, Salinas G, Marin R. 2001. Impact of sediment Heral M. 1996. Variability of feeding processes in the cockle Cerastoderma releases on water chemistry and macroinvertebrate communities in clear edule (L) in response to changes in seston concentration and composition. water Andean streams (Bolivia). Archiv für Hydrobiologie 151:33–50. Journal of Experimental Marine Biology and Ecology 197:121–143. Foster IDL, Collins AL, Naden PS, Sear DA, Jones JI. 2011. Jackson CR, Batzer DP, Cross SS, Haggerty SM, Sturm CA. 2007. Paleolimnology, policy and practice; the potential for lake‐sediment Headwater streams and timber harvest: channel, macroinvertebrate, and based reconstructions to determine modern background sediment amphibian response and recovery. Forest Science 53: 356–370. delivery to rivers. Journal of Paleolimnology 45: 287–360. Jones JI, Davy‐Bowker J, Murphy JF, Pretty JL. 2010. Ecological monitoring Freeman PL, Schorr MS. 2004. Influence of watershed urbanization on fine and assessment of pollution in rivers. In Ecology of Industrial Pollution, sediment and macroinvertebrate assemblage characteristics in Tennessee Batty LC, Hallberg KB (eds). Cambridge University Press: Cambridge; Ridge and Valley Streams. Journal of Freshwater Ecology 19: 353–362. 126–146. Fritz KM, Dodds WK. Pontius J. 1999. The effects of bison crossings on Jones JI, Jeppesen E. 2007. Body size and trophic cascades in lakes. In the macroinvertebrate community in a tallgrass prairie stream. American Body Size: The Structure and Function of Aquatic Ecosystems, Hildrew Midland Naturalist 141: 253–265. AG, Raffaelli DG, Edmonds‐Brown R (eds). Cambridge University Fritz KM, Gangloff MM, Feminella JW. 2004. Habitat modification by the Press: Cambridge; 118–139. stream macrophyte Justicia americana and its effects on biota. Oecologia Jones JI, Collins AL, Naden PS, Sear DA. 2011. The relationship between 140:388–397. fine sediment and macrophytes in rivers. River Research and Applica- Gallepp G. 1974. Diel periodicity in behaviour of the caddisfly, Brachycentrus tions. DOI: 10.1002/rra.1486 americanus (Banks). Freshwater Biology 4:193–204. Kaller MD, Hartman KJ. 2004. Evidence of a threshold level of fine Gardener MB. 1981. Effects of turbidity on feeding rates and selectivity sediment accumulation for altering benthic macroinvertebrate commu- of bluegills. Transactions of the American Fisheries Society 110: nities. Hydrobiologia 518:95–104. 446–450. Kaller MD, Hartman KJ, Angradi TR. 2001. Experimental determination Gaugler R, Molloy D. 1980. Feeding inhibition in black fly larvae (Diptera, of benthic macro invertebrate metric sensitivity to fine sediment in Simuliidae) and its effects on the pathogenicity of Bacillus thuringiensis Appalachian streams. Proceedings of the Fifty‐Fifth Annual Confer- var israelensis. Environmental Entomology 9: 704–708. ence of the Southeastern Association of Fish and Wildlife Agencies Geist J, Auerswald K. 2007. Physicochemical stream bed characteristics and 55: 105–115. recruitment of the freshwater pearl mussel (Margaritifera margaritifera). Kaufmann PR, Larsen DP, Faustini JM. 2009. Bed stability and sedi- Freshwater Biology 52:2299–2316. mentation associated with human disturbances in Pacific Northwest Gjerløv C, Hildrew AG, Jones JI. 2003. Mobility of stream invertebrates in streams. Journal of the American Water Resources Association 45: relation to disturbance and refugia: a test of habitat templet theory. 434– 459. Journal of the North American Benthological Society 22:207–223. Kefford BJ, Zalizniak L, Dunlop JE, Nugegoda D, Choy SC. 2010. How Glozier NE, Culp JM, Reynoldson TB, Bailey RC, Lowell RB, Trudel L. 2002. are macrophytes of slow flowing lotic systems directly affected by Assessing metal mine effects using benthic invertebrates for Canada’s suspended and deposited sediments? Environmental Pollution 158: environmental effects program. Water Quality Research Journal of Canada 543–550. 37:251–278. Kemp JL, Harper DM, Crosa GA. 2000. The habitat‐scale ecohydraulics of Gray L. 2004. Changes in water quality and macroinvertebrate commu- rivers. Ecological Engineering 16:17–29. nities resulting from urban stormflows in the Provo River, Utah, USA. Kemp PS, Sear DA, Collins AL, Naden P, Jones JI. 2011. The impacts of fine Hydrobiologia 518:33–46. sediment on riverine fish. Hydrological Processes. DOI: 10.1002/hyp.7940 Gray LJ, Ward JV. 1982. Effects of sediment releases from a reservoir on Kent TR, Stelzer RS. 2008. Effects of deposited fine sediment on life stream macroinvertebrates. Hydrobiologia 96: 177–184. history traits of Physa integra snails. Hydrobiologia 596: 329–340. Hamilton JD. 1961. The effect of sand‐pit washings on stream fauna. Köhler J, Hachoł J, Hilt S. 2010. Regulation of submersed macrophyte Verhandlungen der Internationale Vereinigung fur Theoretische und biomass in a temperate lowland river: Interactions between shading by Angewandte Limnologie 14:435–439. bank vegetation, epiphyton and water turbidity. Aquatic Botany 92: Hart RC. 1992. Experimental studies of food and suspended sediment 129–136. effects on growth and reproduction of 6 planktonic cladocerans. Journal Kreutzweiser DP, Capell SS, Good KP. 2005. Effects of fine sediment of Plankton Research 14: 1425–1448. inputs from a logging road on stream insect communities: a large‐scale Hedrick LB, Welsh SA, Anderson JT. 2007. Effects of highway experimental approach in a Canadian headwater stream. Aquatic Ecology construction on sediment and benthic macroinvertebrates in two 39:55–66. tributaries of the lost river, West Virginia. Journal of Freshwater Ecology Krieger KA, Bur MT, Ciborowski JJH, Barton DR, Schloesser DW. 2007. 22: 561–569. Distribution and abundance of burrowing mayflies (Hexagenia spp.) in Herbst DB, Kane JM. 2006. Fine Sediment Deposition and Invertebrate Lake Erie, 1997–2005. Journal of Great Lakes Research 33:20–33. Communities in the middle Truckee River, California: Development of Kurtak DC. 1978. Efficiency of filter feeding of black fly larvae (Diptera Criteria for Establishing TMDLs. Lahontan Regional Water Quality Simuliidae). Canadian Journal of Zoology 56: 1608–1623. Control Board Report. Lamberti GA, Steinman AD. 1993. Research in artificial streams— Hornig CE, Brusven MA. 1986 Effects of suspended sediment on leaf applications, uses, and abuses. Journal of the North American processing by Hesperophylax occidentalis (Trichoptera, Limnephilidae) Benthological Society 12: 313–384. and Pteronarcys californica (Plecoptera, Pteronarcidae). Great Basin Lancaster J. 1996. Scaling the effects of predation and disturbance in a Naturalist 46:33–38. patchy environment. Oecologia 107: 321–331.

Lancaster J, Hildrew AG. 1993. Characterizing in‐stream flow refugia. interface: bioturbation and consumer‐substrate interaction. Oecologia 161: Canadian Journal of Fisheries and Aquatic Sciences 50: 1663–1675. 125–138. Lancaster J, Hildrew AG, Townsend CR. 1991. Invertebrate predation on Nuttall PM. 1972. The effect of sand deposition upon the macro- patchy and mobile prey in streams. The Journal of Animal Ecology 60: invertebrate fauna of the River Camel, Cornwall. Freshwater Biology 625–641. 2: 181–186. Lane PNJ, Sheridan GJ. 2002. Impact of an unsealed forest road stream Nuttall PM, Bielby GH. 1973. The effects of china‐clay wastes on stream crossing: water quality and sediment sources. Hydrological Processes invertebrates. Environmental Pollution 5:77–86. 16: 2599–2612. Osterling EM, Greenberg LA, Arvidsson BL. 2008. Relationship of biotic and Larsen S, Ormerod SJ. 2010. Low‐level effects of inert sediments on abiotic factors to recruitment patterns in Margaritifera margaritifera. temperate stream invertebrates. Freshwater Biology 55: 476–486. Biological Conservation 141:1365–1370. Larsen S, Vaughan IP, Ormerod SJ. 2009. Scale‐dependent effects of fine Pardo I, Armitage PD. 1997. Species assemblages as descriptors of sediments on temperate headwater invertebrates. Freshwater Biology 54: mesohabitats. Hydrobiologia 344: 111–128. 203–219. Parkhill KL, Gulliver JS. 2002. Effect of inorganic sediment on whole‐ Lemly AD. (1982). Modification of benthic insect communities in polluted stream productivity. Hydrobiologia 472:5–17. streams—combined effects of sedimentation and nutrient enrichment. Peckarsky BL. 1991. habitat selection by stream‐dwelling predatory Hydrobiologia 87: 229–245. stoneflies. Canadian Journal of Fisheries and Aquatic Sciences 48: Levesque LM, Dube MG. 2007. Review of the effects of in‐stream pipeline 1069–1076. crossing construction on aquatic ecosystems and examination of Canadian Peckarsky BL, Kerans BL, Taylor BW, Mcintosh AR. 2008. Predator methodologies for impact assessment. Environmental Monitoring and effects on prey population dynamics in open systems. Oecologia 156: Assessment 132:395–409. 431–440. MacIsaac HJ, Rocha R. 1995. Effects of suspended clay on zebra mussel Petts G, Armitage P, Castella E. 1993. Physical habitat changes and (Dreissena polymorpha) faeces and pseudofaeces production. Archiv für macroinvertebrate response to river regulation—the River Rede, UK. Hydrobiologie 135:53–64. Regulated Rivers: Research & Management 8: 167–178. Mainstone CP, Dils RM, Withers PJA. 2008. Controlling sediment and Pretty JL, Hildrew AG, Trimmer M. 2006. Nutrient dynamics in relation to phosphorus transfer to receiving waters—a strategic management perspec- surface‐subsurface hydrological exchange in a groundwater fed chalk tive for England and Wales. Journal of Hydrology 350: 131–143. stream. Journal of Hydrology 330:84–100. Mainstone CP, Parr W. 2002. Phosphorus in rivers—ecology and Quinn JM, Cooper AB, Davies‐Colley RJ, Rutherford JC, Williamson RB. management. The Science of the Total Environment 282:25–47. 1997b. Land use effects on habitat, water quality, periphyton, and Maloney KO, Feminella JW. 2006. Evaluation of single‐ and multi‐metric benthic invertebrates in Waikato, New Zealand, hill‐country streams. benthic macroinvertebrate indicators of catchment disturbance over time New Zealand Journal of Marine and Freshwater Research 31: 579–597. at the Fort Benning Military Installation, Georgia, USA. Ecological Quinn JM, Cooper AB, Stroud MJ, Burrell GP. 1997a. Shade effects on Indicators 6: 469–484. stream periphyton and invertebrates: an experiment in streamside Maloney KO, Mulholland PJ, Feminella JW. 2005. Influence of catchment‐ channels. New Zealand Journal of Marine and Freshwater Research scale military land use on stream physical and organic matter variables in 31: 665–683. small southeastern plains catchments (USA). Environmental Management Quinn JM, Davies‐Colley RJ, Hickey CW, Vickers ML, Ryan PA. 1992. 35:677–691. Effects of clay discharges on streams 2. Benthic invertebrates. Matthaei CD, Weller F, Kelly DW, Townsend CR. 2006. Impacts of fine Hydrobiologia 248: 235–247. sediment addition to tussock, pasture, dairy and deer farming streams in Quinn JM, Hickey CW. 1990. Characterization and classification of benthic New Zealand. Freshwater Biology 51: 2154–2172. invertebrate communities in 88 New Zealand rivers in relation to Mcclelland WT, Brusven MA. 1980. Effects of sedimentation on the environmental factors. New Zealand Journal of Marine and Freshwater behavior and distribution of riffle insects in a laboratory stream. Aquatic Research 24: 387–409. Insects 2: 161–169. Quist MC, Fay PA, Guy CS, Knapp AK, Rubenstein BN. 2003. Military Milner AM, Piorkowski RJ. 2004. Macroinvertebrate assemblages in streams training effects on terrestrial and aquatic communities on a grassland of interior Alaska following alluvial gold mining. River Research and military installation. Ecological Applications 13: 432–442. Applications 20:719–731. Radwell AJ, Brown AV. 2006. Influence of fine sediments on meiofauna Molinos JG, Donohue I. 2009. Differential contribution of concentration colonization densities in artificial stream channels. Archiv für Hydrobiologie and exposure time to sediment dose effects on stream biota. Journal of 165:63–75. the North American Benthological Society 28: 110–121. Rasmussen JB. 1988. Habitat requirements of burrowing mayflies Moss D, Furse MT, Wright JF, Armitage PD. 1987. The prediction of the Ephemeridae Hexagenia in lakes with special reference to the effects macroinvertebrate fauna of unpolluted running water sites in Great of eutrophication. Journal of the North American Benthological Society Britain using environmental data. Freshwater Biology 17:41–52. 7:51–64. Newcombe CP, MacDonald DD. 1991. Effects of suspended sediments on Ritchie JC. 1972. Sediment, fish and fish habitat. Journal of Soil and Water aquatic ecosystems. North American Journal of Fisheries Management Conservation 27: 124–125. 11:72–82. Robertson M. 1957. The effects of suspended materials on the reproductive Niyogi DK, Koren M, Arbuckle CJ, Townsend CR. 2007. Stream communities rate of Daphnia magna. Publications of the Institute of Marine Sciences, along a catchment land‐use gradient: subsidy‐stress responses to pastoral University of Texas 4: 265–277. development. Environmental Management 39:213–225. Rosenberg DM, Wiens AP. 1978. Effects of sediment addition on Nogaro G, Mermillod‐Blondin F, Francois‐Carcaillet F, Gaudet JP, Lafont macrobenthic invertebrates in a northern Canadian river. Water Research M, Gibert J. 2006. Invertebrate bioturbation can reduce the clogging of 12: 753–763. sediment: an experimental study using infiltration sediment columns. Roy AH, Rosemond AD, Leigh DS, Paul MJ, Wallace JB. 2003a. Habitat‐ Freshwater Biology 51: 1458–1473. specific responses of stream insects to land cover disturbance: biological Nogaro G, Mermillod‐Blondin F, Valett MH, Francois‐Carcaillet F, Gaudet consequences and monitoring implications. Journal of the North JP, Lafont M, Gibert J. 2009. Ecosystem engineering at the sediment‐water American Benthological Society 22: 292–307.

Roy AH, Rosemond AD, Paul MJ, Leigh DS, Wallace JB. 2003b. Stream Voelz NJ, Ward JV. 1992. Feeding habits and food resources of filter‐ macroinvertebrate response to catchment urbanisation (Georgia, USA). feeding Trichoptera in a regulated mountain stream. Hydrobiologia 231: Freshwater Biology 48: 329–346. 187–196. Rutherford JE, Mackay RJ. 1986. Patterns of pupal mortality in field Wagener SM, Laperriere JD. 1985. Effects of placer mining on the populations of Hydropsyche and Cheumatopsyche (Trichoptera, invertebrate communities of interior Alaska. Freshwater Invertebrate Hydropsychidae). Freshwater Biology 16: 337–350. Biology 4: 208–214. Sand‐Jensen K. 1997. Macrophytes as biological engineers in the ecology Wallace ID, Wallace B, Philipson GN. 1990. A Key to the Case‐bearing of Danish streams. In Freshwater Biology. Priorities and Development Caddis Larvae of Britain and Ireland. Freshwater Biological Association: in Danish Research, Sand‐Jensen K, Pedersen O (eds). G.E.C. Gad: Ambleside. Copenhagen; 74–101. Walling DE. 1995. Suspended sediment yields in a changing environment. Sand‐Jensen K, Jeppesen E, Nielsen K, Van Der Bijl L, Hjermind L, In Changing River Channels, Gurnell AM, Petts GE (eds). John Wiley & Nielsen LW, Iversen TM. 1989. Growth of macrophytes and ecosystem Sons: Chichester; 149–176. consequences in a lowland Danish stream. Freshwater Biology 22:15–32. Walling DE, Fang D. 2003. Recent trends in the suspended sediment loads Sand‐Jensen K, Madsen TV. 1989. Invertebrates graze submerged rooted of the world’s rivers. Global and Planetary Change 39: 111–126. macrophytes in lowland steams. Oikos 55: 420–423. Waters TF. 1995. Sediment in Streams: Sources, Biological Effects, and Sarriquet PE, Bordenave P, Marmonier P. 2007. Effects of bottom sediment Control. American Fisheries Society Monograph 7: Bethesda, MD. restoration on interstitial habitat characteristics and benthic macro- Welton JS, Clarke RT. 1980. Laboratory studies on the reproduction and invertebrate assemblages in a headwater stream. River Research and growth of the amphipod, Gammarus pulex (L). The Journal of Animal Applications 23: 815–828. Ecology 49: 581–592. Sear DA, Frostick LB, Rollinson G, Lisle TE. 2008. The significance Wharton G, Cotton JA, Wotton RS, Bass JAB, Heppell CM, Trimmer M, and mechanics of fine sediment infiltration and accumulation in gravel Sanders IA, Warren LL. 2006. Macrophytes and suspension‐feeding spawning beds. In Salmonid Spawning Habitat in Rivers; Physical invertebrates modify flows and fine sediments in the Frome and Piddle Controls, Biological Responses and Approaches to Remediation, catchments, Dorset (UK). Journal of Hydrology 330: 171–184. Sear DA, Devries P (eds). AFS: Bethesda, MA; 149–174. White DS, Gammon JR. 1977. The effect of suspended solids on Smith BPG, Naden PS, Leeks GJL, Wass PD. 2003. Characterising the fine macroinvertebrate drift in an Indiana creek. Proceedings of the Indiana sediment budget of a reach of the River Swale, Yorkshire, UK during the Academy of Science 86:182–188. 1994 to 1995 winter season. Hydrobiologia 494: 135–143. Williams DD, Smith MR. 1996. Colonization dynamics of river benthos in Stead TK, Schmid‐Araya JM, Hildrew AG. 2003. All creatures great and response to local changes in bed characteristics. Freshwater Biology 36: small: patterns in the stream benthos across a wide range of metazoan 237–248. body size. Freshwater Biology 48: 532–547. Williams LR, Bonner TH, Hudson JD, Williams MG, Leavy TR, Williams Stead TK, Schmid‐Araya JM, Hildrew AG. 2004. The contribution of CS. 2005. Interactive effects of environmental variability and military subsurface invertebrates to benthic density and biomass in a gravel training on stream biota of three headwater drainages in western stream. Archiv für Hydrobiologie 160: 171–191. Louisiana. Transactions of the American Fisheries Society 134: 192–206. Stout BM, Coburn CB. 1989. Impact of highway construction on leaf processing Wood PJ, Armitage PD. 1997. Biological effects of fine sediment in the in aquatic habitats of eastern Tennessee. Hydrobiologia 178: 233–242. lotic environment. Environmental Management 21: 203–217. Strand RM, Merritt RW. 1997. Effects of episodic sedimentation on the Wood PJ, Toone J, Greenwood MT, Armitage PD. 2005. The response of net‐spinning caddisflies Hydropsyche betteni and Ceratopsyche sparna four lotic macroinvertebrate taxa to burial by sediments. Archiv für (Trichoptera: Hydropsychidae). Environmental Pollution 98: 129–134. Hydrobiologie 163: 145–162. Suren AM, Jowett IG. 2001. Effects of deposited sediment on invertebrate Wood PJ, Vann AR, Wanless PJ. 2001. The response of Melampophylax drift: an experimental study. New Zealand Journal of Marine and mucoreus (Hagen) (Trichoptera: Limnephilidae) to rapid sedimentation. Freshwater Research 35: 725–737. Hydrobiologia 455: 183–188. Suren AM, Martin ML, Smith BJ. 2005. Short‐term effects of high Wotton RS, Malmqvist B. 2001. Feces in aquatic ecosystems. Bioscience suspended sediments on six common New Zealand stream invertebrates. 51: 537–544. Hydrobiologia 548:67–74. Wotton RS, Warren LL. 2007. Impacts of suspension feeders on the Svendsen KM, Renshaw CE, Magilligan FJ, Nislow KH, Kaste JM. 2009. modification and transport of stream seston. Fundamental and Applied Flow and sediment regimes at tributary junctions on a regulated river: Limnology 169: 231–236. impact on sediment residence time and benthic macroinvertebrate Yule CM, Boyero L, Marchant R. 2010. Effects of sediment pollution on food communities. Hydrological Processes 23: 284–296. webs in a tropical river (Borneo, Indonesia). Marine and Freshwater Swan CM, Palmer MA. 2000. What drives small‐scale spatial patterns in Research 61:204–213. lotic meiofauna communities? Freshwater Biology 44: 109–121. Zamor RM, Grossman GD. 2007. Turbidity affects foraging success of drift‐ Trimmer M, Sanders IA, Heppell CM. 2009. Carbon and nitrogen cycling feeding rosyside dace. Transactions of the American Fisheries Society 136: in a vegetated lowland chalk river impacted by sediment. Hydrological 167–176. Processes 23: 2225–2238. Zimmerman GF, De Szalay FA. 2007. Influence of unionid mussels Vaughn CC, Hakenkamp CC. 2001. The functional role of burrowing (Mollusca: Unionidae) on sediment stability: an artificial stream study. bivalves in freshwater ecosystems. Freshwater Biology 46: 1431–1446. Fundamental and Applied Limnology 168: 299–306. Vermaat JE, De Bruyne RJ. 1993. Factors limiting the distribution of Zweig LD, Rabeni CF. 2001. Biomonitoring for deposited sediment using submerged waterplants in the lowland River Vecht (the N.L.). Freshwater benthic invertebrates: a test on 4 Missouri streams. Journal of the North Biology 30:147–157. American Benthological Society 20: 643–657.