Journal of Sea Research 48 (2002) 143 156 www.elsevier.

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Impact of biotic and abiotic processes on sediment dynamics and the consequences to the structure and functioning of the intertidal zone
John Widdows *, Mary Brinsley
Plymouth Marine Laboratory, Centre for Coastal and Marine Sciences, Prospect Place, West Hoe, Plymouth, PL1 3DH, England, UK Received 30 November 2001; accepted 30 May 2002

Abstract This paper reviews field and laboratory studies using flumes to quantify the erodability of undisturbed intertidal sediments as a function of changes in (1) the natural benthic community structure and sediment properties, and (2) the abundance of key intertidal species. Sediment erodability, which varies spatially and temporally, is dependent on the interactions between physical processes, sediment properties and biological processes, particularly the balance between two functional groups of biota, the stabilisers and the destabilisers. Bio-stabilisers can influence the hydrodynamics and provide some physical protection to the bed (e.g. mussel beds, macroalgae, salt marsh macrophytes), or can enhance cohesiveness and alter the critical erosion threshold (e.g. microphytobenthos). In contrast, bio-destabilisers (e.g. bioturbators such as Macoma balthica, Hydrobia ulvae) increase surface roughness, reduce the critical erosion threshold and enhance the erosion rate. Field studies in the Humber (England) and Westerschelde (Netherlands) have shown that interannual changes in sediment erodability were a result of a shift from a stabilised sediment dominated by microphytobenthos to a destabilised sediment dominated by M. balthica. Interannual changes in key biota, their influence on sediment erosion, and the consequences for intertidal ecology and morphology, appear to be driven in part by climatic factors (primarily a shift from mild to cold winters). Quantification and understanding of these benthic processes has been used to parameterise mathematical models of intertidal sediment dynamics, and this has provided insight into the relative importance of biological and physical factors in determining sediment erosion/accretion in the intertidal zone. D 2002 Elsevier Science B.V. All rights reserved.
Keywords: Intertidal sediments; Erosion; Deposition; Biota; Climatic factors; Modelling; NW Europe

1. Introduction When examining the structure and functioning of intertidal sediment shores and associated benthic communities, it is necessary to consider: (1) the

Corresponding author. E-mail address: j.widdows@pml.ac.uk (J. Widdows).

impact of environmental factors on biota (i.e. ecophysiology, eco-toxicology and biogeography), (2) the influence of biota on the composition of benthic communities (i.e. trophic interactions), and (3) the impact of biota on their environment (i.e. ecological engineering, eco-morphology). All ecosystems are controlled and organised by a small set of key plant, animal and abiotic processes that engineer and structure the landscape or substrate

1385-1101/02/$ - see front matter D 2002 Elsevier Science B.V. All rights reserved. PII: S 1 3 8 5 - 11 0 1 ( 0 2 ) 0 0 1 4 8 - X

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at different scales (Lawton and Jones, 1995). In the field of marine benthic ecology the concept of ecological or bio-engineering is well established, but there is a requirement for improved quantification, understanding and modelling of the role of biological as well as physical processes in hydro- and sediment dynamics. Such information is essential for the development and improved parameterisation of models for forecasting changes in sediment dynamics and estuarine morphology, particularly in response to sea-level rise and climate change. The aim of this paper is to review recent field and laboratory studies using flumes to quantify the erodability of intertidal sediments as a function of changes in (1) the natural benthic community structure and sediment properties, and (2) the density of a range of key intertidal organisms. The interaction between the biological and physical processes, and the consequences for the structuring and functioning of an intertidal sediment shore have also been examined. Clearly, it is necessary to consider not only how biota structure and modify their intertidal sediment environment (biostabilisers and -destabilisers), but also some of the key drivers that influence community structure and functioning (grazing/predation, climate). Such research is highly relevant to the improved quantification, understanding and modelling of sediment dynamics and estuarine morphology, which in turn is relevant to important environmental issues such as assessment of: a) impact of sea-level rise and increased storminess on natural coastal defences (tidal flats, salt marshes, biogenic reefs); b) sustainability of intertidal habitats for birds, fisheries, aquaculture; c) potential restoration of valuable ecological areas (e.g. wetlands); d) potential for mobilisation of historically polluted sediments; e) dredging and disposal of sediment from shipping channels and harbours/marinas, including the beneficial use of this material on the intertidal. It is clear that cohesive sediment dynamics cannot be modelled based on physical processes alone because biota can affect sediment erosion/deposition by several orders of magnitude (Paterson and Black,

1999; Whitehouse et al., 2000; Widdows, 2001). Some of the concepts and relationships between water flow, sediment dynamics and benthic organisms have been reviewed by Paterson and Black (1999). Sediment erodability, which varies spatially and temporally, is dependent on the interactions between physical processes, sediment properties and biological processes, particularly the balance between two functional groups of biota, the bio-stabilisers and the biodestabilisers (or bioturbators). Bio-stabilisers (e.g. mussel beds/oyster reefs, seagrass beds, macroalgal mats, microphytobenthos and salt marsh macrophytes) can modify their immediate physical environment by reducing tidal currents, wave action and sediment resuspension, thus reducing turbidity and increasing light penetration, as well as enhancing sediment cohesiveness and sedimentation processes. In contrast, the bio-destabilisers increase sediment erosion/ resuspension, turbidity and modify properties of surficial sediments by increasing bed roughness, sediment water content, grazing on biostabilisers and producing faecal pellets.

2. Description of intertidal gradient and measurement of sediment erodability Biota living in the intertidal zone of meso- and macrotidal estuaries and coastal embayments are adapted to a dynamic environment dominated by the tidal flooding and ebbing of water with periods of aerial exposure. For example, in the Humber estuary (tidal range of 7.7 m) with extensive intertidal sand and mudflats up to 4 km wide, spring tidal currents can reach maximum speeds of up to 0.6 m s 1 at the mid-shore level, and declining shorewards to 0.4 m s 1 on the upper shore (Christie and Dyer, 1998; Black, 1998). Maximum tidal currents during neap tides are generally below the critical erosion threshold (e.g. only 0.1 to 0.2 m s 1 in the Humber estuary; Black, 1998; Widdows et al., 1998a). Near-bed current speeds (i.e. z = 0.1 m) and bed shear stresses will ultimately be dependent on tidal amplitude, shore height, shore gradient (shallow gradients increasing tidal currents) and bed roughness. Biota and the biological processes relevant to sediment dynamics have to be considered in relation to an underlying physical gradient of increasing air exposure shorewards (from

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subtidal (0 h) to 11 h air exposure in a 12.5 h tidal cycle on the upper shore), declining tidal currents (from >0.6 to 0.03 m s 1), declining wave action (depending on fetch and water depth), and a general reduction of grain size (from sand to cohesive fine muds). Sediment erosion potential or erodability is defined in terms of critical erosion threshold, erosion rate (g m 2 s 1) and mass of sediment eroded (g m 2) in relation to measured current velocities and estimated bed shear stresses. The erosion potential of marine cohesive sediments has been measured using various devices described by Black and Paterson (1997). However, the studies reviewed here are mainly restricted to annular flumes because they can measure sediment erosion and deposition in relation to natural sediment with its benthic community (i.e. undisturbed during sampling) and can accommodate a wide range of biota from microphytobenthos to mussel beds and salt marsh plants (for detailed descriptions see Amos et al., 1992; Widdows et al., 1998b). In general, the results are expressed in relation to free-stream current velocity at 0.1 m above the bed in preference to bed shear stress, because the former parameter can be quantified directly and with greater accuracy both in

flumes and the field (Widdows et al., 2000a). In contrast, bed shear stress is estimated indirectly, assuming a simple log velocity profile, and there is considerable variation in the estimated bed shear stress in the literature. In addition, the boundary layer flow over tidal flats is usually complex due to accelerating and decelerating tidal flow in shallow water, and the influence of biota on the hydrodynamics (Paterson and Black, 1999). Processes of sediment erosion and deposition are central to the behaviour and ecology of intertidal and subtidal organisms. This review focuses on recent studies that have quantified the impact of key intertidal biota on sediment dynamics and benthic pelagic exchanges, beginning with biota on the lower shore and progressing to the upper shore (conceptual model; Fig. 1). Mussels on the lower shore experience and utilise strong tidal currents, enhancing sediment stability by orders of magnitude through the physical protection of the bed and impact on hydrodynamics via increased friction drag. On the middle to upper shore the structure and functioning of the benthic community is probably more changeable, with the overall balance shifting spatially and temporally between bio-stabilisers such as microphytobenthos and

Fig. 1. Schematic diagram summarising some of the major biological and physical factors influencing sediment stability in the intertidal zone. The dotted line represents the general shoreward increase in sediment stability with increasing cohesiveness and consolidation due to increasing air exposure and declining currents/wave action. The solid lines represent the long-term increase in sediment stability as a result of biota that persist for many years to >100 years (e.g. mussel beds and salt marsh). The dashed oscillating line represents the spatial (and temporal) changes in sediment erodability due to short-term shifts (0.1 to >1 year) in the balance between the biostabilisers (microphytobenthos) and biodestabilisers (Macoma balthica Hydrobia ulvae).

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the macroalgae (Enteromorpha) and the de-stabilisers (Macoma balthica, Hydrobia ulvae). Salt marsh plants dominate the upper part of the intertidal zone and their mode of impact is via hydrodynamics (increased friction drag) and sediment trapping.

3. Impact of intertidal biota on sediment erodability 3.1. Mussel beds on lower to mid shore Mussel beds (Mytilus edulis), if they are not disturbed or over-fished, can form extensive biogenic reefs on lower sediment shores, particularly at the mouths of meso- and macro-tidal estuaries where there is good tidal flow. However, sessile suspension feeders such as mussels, when inhabiting sediment shores with high tidal currents need to achieve a balance between maximizing food supply and minimizing erosion of the bed/substrate. Mussels benefit from high tidal flows by maximizing replenishment of food supply while simultaneously minimizing seston depletion immediately above extensive mussel beds. In these same environments they are potentially vulnerable to erosion and instability if the sediment is not protected/stabilised. High density mussel beds not only physically protect the sediment surface from resuspension and erosion at times of maximum current speeds (spring tides), but also enhance the flux of suspended particulate matter (SPM) from the water column to the bed (via suspension feeding and biodeposition). Biodeposition rates (g m 2 h 1) are ultimately dependent on the product of clearance rates of individual mussels (typically c. 4 l g 1 dry tissue mass h 1 at estuarine sites), the SPM concentration (g l 1) and the mussel density (up to 3500 individuals m 2). Hence, biodeposition rates in silty estuaries can be high, up to 60 g m 2 h 1, equivalent to 40 times the natural sedimentation rate (Widdows et al., 1998c). However, the biodeposits have a critical threshold for resuspension at 0.2 m s 1 and therefore the majority of biodeposits will be resuspended by tidal currents and not consolidated or incorporated into the bed (Widdows et al., 1998c). Below current speeds of 0.2 m s 1 the net biodeposition is greater than net erosion, whereas above this threshold the balance changes to net erosion (Widdows et al.,

1998c). The accumulation of a small proportion of the biodeposits in the sandy sediment will gradually increase the silt content of the sediment, particularly during the spring and summer, thus modifying the sediment properties and environment for other benthic organisms (Widdows et al., 2000a). The silt content will then be reduced by resuspension during winter storms. Mussels (Mytilus edulis) from SW England have been shown to maintain their suspension feeding activity or clearance rates independent of current speeds between at least 0.05 and 0.8 m s 1 (Widdows et al., 2002). These observations are consistent with the occurrence of mussels in environments with high current speeds and turbulence (e.g. coasts of Western Europe and cooling water outflows at power stations) but are in contrast with a marked decline in clearance rate recorded in North American mussels (Mytilus edulis) exposed to currents between 0.05 to 0.25 m s 1 (Wildish and Miyares, 1990; Newell, 1999). This may reflect geographical differences (genotypic or phenotypic) between mussel populations (Widdows et al., 2002). Flume studies have quantified the erodability of cohesive sediments (46% silt) associated with mussel beds in the mouth of the Humber estuary (Widdows et al., 1998c). Sediment resuspension was measured in relation to changes in mussel bed density (% cover) and was found to decline in an exponential manner with increasing mussel bed density. At the highest densities, mussels reduce sediment erosion by 10-fold. In contrast, mussel beds overlying sandy substrates at Exmouth (England) showed evidence of a marked non-linear relationship between sediment erodability and mussel bed density (Fig. 2; Widdows et al., 2002). Maximum erosion occurred at intermediate densities (25 and 50% cover) compared to bare sand (0% cover). This was primarily due to increased scouring erosion around clumps of mussels (low density condition) and the resuspension of silty biodeposits accumulated in the sediment associated with mussels. Flume studies have also shown that the friction drag induced by mussel beds, regardless of mussel densities between 25 100%, reduces the water flow and bed shear stress by 25% relative to a bare sand (Widdows et al., 2002). This will be a contributory factor to the reduction in sediment erosion.

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Fig. 2. Effect of Mytilus edulis density (% bed cover) and depth-averaged current velocity (z = 0 to 0.1 m) on erodability of sandy substrate at Exmouth, England (redrawn from Widdows et al., 2002).

Results of erosion studies indicate that a reduction in mussel coverage (density) from 100 to 50% or less will enhance sediment erosion at current velocities >0.25 m s 1, re-suspend biodeposits and therefore make the mussel bed more vulnerable to erosion during maximum spring tides and storm events (Widdows et al., 2002). The more open coastal areas with sandy substrates and a long fetch (e.g. Wash and Wadden Sea) will be particularly vulnerable if mussel beds have been opened up and physically disturbed, either by fishing activities or storms. Well-established beds, together with accumulated sediments, can be 1 2 m above the natural sediment level and thus have a very visible impact on estuarine morphology and are recorded as mussel scalps on geological maps. However, once these beds have been severely eroded then they are difficult to re-establish (e.g. mussel beds in the Wash and Wadden Sea compared to beds at Exmouth which are not fished). 3.2. Biostabilisers and bioturbators on mid to upper shore On the middle and upper fine sediment shores of estuaries and embayments in NW Europe there is evidence of marked spatial and temporal variations in sediment stability (Widdows et al., 1998a; Widdows et al., 2000a,b; Andersen, 2001), primarily due to

the seasonal and interannual changes in the balance between bio-stabilisers (such as microphytobenthos, Enteromorpha spp., Cladophora rupestris) and biodestabilisers (such as Macoma balthica, Hydrobia ulvae). 3.2.1. Algal biofilm - microphytobenthos The role of the microphytobenthos as sediment stabilisers has been well documented and quantified (Paterson, 1989; Underwood and Paterson, 1993; Sutherland et al., 1998; Paterson and Black, 1999). Recently, Sutherland et al. (1998) demonstrated strong relationships between critical erosion thresholds and microphytobenthos density, measured in terms of sediment concentrations of colloidal carbohydrate and chlorophyll-a. These studies showed a significant increase in critical shear stress for erosion (U * crit) and a reduction in erosion rate with increasing mucus-like carbohydrate concentrations and sediment chlorophyll-a (Fig. 3). These cellular exudates, also referred to as extra-polymeric substances (EPS), are used in the locomotion of diatoms, bacteria and cyanobacteria, and may also protect cells against desiccation during periods of air exposure. Multi-disciplinary field campaigns in the Humber estuary, England (Amos et al., 1998; Widdows et al., 2000b), the Westerschelde, Netherlands (Widdows et al., 2000a; De Brouwer et al., 2000) and Wadden Sea,

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Fig. 3. Relationship between sediment erodability (erosion rate and U * crit) and microphytobenthos biomass (chlorophyll-a) (redrawn from Sutherland et al., 1998). The units of chlorophyll-a concentration have been expressed as Ag g 1 dry sediment and as mg m 2.

Germany (Riethmu ller et al., 2000; Andersen, 2001) have recorded considerable spatial and temporal variation in sediment erodability. Enhanced sediment stability, characterised by higher critical erosion thresholds and lower erosion rates, was due primarily to the establishment of microphytobenthos or algal biofilms on the sediment surface. 3.2.2. Bioturbators/grazers The clam, Macoma balthica, can be present at very high but variable densities in intertidal cohesive sediments of northern temperate environments. Sediment erosion studies using annular flumes in both the Humber and the Westerschelde have recorded a significant correlation between sediment erodability and the activity and density of M. balthica (Widdows et al., 1998a, 2000a,b). Associated laboratory studies quantified the impact of M. balthica on sediment erosion with increasing current velocity/bed shear stress (Fig. 4; Widdows et al., 1998c). The erosion rate and sediment mass eroded was significantly enhanced by the activities of M. balthica and was density dependent. Recorded spatial and temporal variation in sediment erodability on the mid to upper shores of both the Humber and Westerschelde estuaries (Widdows et al., 2000a,b) was mainly due to changes in the relative balance between the biomass of bio-stabilisers (microphytobenthos) and bio-destabilisers (M. balthica). In

both estuaries during the spring of 1996, there was evidence of a well-developed microphytobenthos at a time of low M. balthica abundance on the tidal flats. This was accompanied by high sediment stability, with a critical erosion velocity (Ucrit) of 0.35 m s 1 and low erosion rates. In the spring and autumn of 1997, there was a contrasting situation with a less dominant microphytobenthos and significantly higher densities of M. balthica. As a result of the burrowing, deposit feeding and grazing activity of M. balthica, the surficial sediment had a lower Ucrit ( < 0.15 m s 1) and a higher erosion rate and the sediment was more easily eroded (10 to 100-fold increase). Field studies also showed that interannual variation in biotically induced sediment erodability, between 1995 and 1997, was greater than seasonal or spatial variation (Widdows et al., 2000b). A comparison of spring 1996 and 1997 showed a marked year to year shift in erodability in both the Humber and the Westerschelde estuaries (Widdows et al., 2000a,b). Long term studies by Beukema et al. (1998) in the Wadden Sea (1973 1996) have demonstrated a significant negative correlation between winter temperatures and the subsequent Macoma recruitment. M. balthica densities are generally found to be higher in the year following a cold winter and it is hypothesised that this is due to an increase in fecundity and a reduction in predators (Beukema et al., 1998). This relationship

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Fig. 4. Influence of Macoma balthica density (individuals m 2) and depth-averaged current velocity (z = 0 to 0.1 m) on mass of sediment eroded m 2 (redrawn from Widdows et al., 1998c). Estimated bed shear stress (Pa) in relation to measured depth-averaged current velocities (U = cm s 1) in the PML annular flume is described by the equation: Bed shear stress (Pa) = 0.0008U2 0.0006U + 0.0052 (Widdows et al., 1998a).

also implies that population changes in M. balthica will occur over a large spatial scale (i.e. NW Europe). Therefore recorded interannual changes in erodability of intertidal sediments are probably driven by climate factors mediated through interannual biological cycles, such as those of the bioturbator M. balthica. Clearly there is also an interaction between the M. balthica density, its grazing activity and the microphytobenthos biomass. At times of high M. balthica abundance the microphytobenthos will be heavily grazed and its biomass lowered. There is also evidence of predation by birds influencing sediment stability indirectly and through a cascade effect (Dabourn et al., 1993). Corophium volutator is an important grazer on the microphytobenthos and the removal of C. volutator by large flocks of intertidal waders promoted the development of the microphytobenthos, which in turn enhanced sediment stability. A pattern of decreasing sediment stability with distance from the upper shore salt marsh has been recorded by Austen et al. (1999) in the Danish Wadden Sea, with areas close to the salt marsh

dominated by microphytobenthos and areas offshore being increasingly dominated by H. ulvae. Austen et al. (1999) concluded that a variety of factors can influence sediment erodability, acting together in a complex manner, but identifying causal relationships required further experiments. Subsequent field studies in the Danish Wadden Sea (using the EROMES erosion device) have shown that high densities H. ulvae can have a significant effect on sediment erodability by decreasing the erosion threshold and increasing erosion rate (Andersen, 2001). Erosion rate was found to be dependent on the faecal pellet content of the sediment and there was a strong seasonal variation with higher production of faecal pellets and higher sediment erosion rates during the warmer months. At a contrasting site in the Wadden Sea, the sediment was dominated by diatom biofilms and had low densities of H. ulvae, and this resulted in higher erosion thresholds and lower erosion rates during the warmer months (Andersen, 2001). Flume experiments by Blanchard et al. (1997) also indicated that the presence of H. ulvae can enhance sediment resuspension.

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3.2.3. Macroalgal mats of enteromorpha Recent flume studies have also quantified the impact of the macroalga Entermorpha sp. on hydroand sediment dynamics (Romano et al., submitted). Enteromorpha has become a nuisance macroalgae in many European estuaries, particularly in sheltered areas receiving nutrient enrichment. It is of growing concern because when covering extensive intertidal areas at high densities, Enteromorpha can have adverse effects on macrofauna, fish and birds. These effects include: (1) physical smothering of sediment surface, (2) reducing water flow/exchange for suspension feeders, and (3) causing anoxic conditions within the sediment following burial during storm events. The UK Environment Agency regards 25% cover by Enteromorpha as a maximum critical threshold (Environment Agency, 1999). Although Enteromorpha has a seasonal growth cycle it has become more persistent and significantly increased its percent cover over the last few decades (Raffaelli et al., 1999). Its persistence may be due in part to the positive feedback that it induces. Enteromorpha lowers near-bed current velocities, thus reducing sediment resuspension/turbidity which in turn increases light penetration, and this will enhance

its growth potential. A number of these aspects were investigated and quantified in recent flume experiments (Romano et al., submitted). The main conclusions of these studies were that there is: (1) a significant increase in friction drag with increasing Enteromorpha biomass or % cover (ranging from a 15% reduction in current speed at 10% cover to 42% reduction at 60% cover); (2) a significant reduction in sediment erosion (60 to 90% reduction) with an increase in Enteromorpha from 10 to 60% cover (Fig. 5); and (3) sediment deposition is enhanced in the presence of Enteromorpha (60% cover), particularly during the first 30 min of slack water (i.e. < 0.05 m s 1) when deposition rate was 48% higher than for bare sediment. Once trapped this fine material was not readily resuspended following an increase in current speed. Raffaelli et al. (1998) have assessed the various ecological consequences of Enteromorpha growth in the intertidal zone and experimental flume studies (Romano et al., submitted) have investigated some of the sedimentological effects. However, the possible causes of extensive macroalgal growth remains a contentious issue and Raffaelli et al. (1999) were not able to reach a conclusion. Sfriso and Marcomini

Fig. 5. Impact of different densities of Enteromorpha on sediment erosion relative to bare sediment (100%), expressed both at a standardised depth averaged current velocity (z = 0 to 0.1 m) of 0.25 m s 1 [A] and at the observed lower depth averaged current velocities induced by each density of Enteromorpha [B] (redrawn from Romano et al., submitted).

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(1996) found that unfavourable weather conditions during consecutive years, extending winter-like conditions until May June, prompted the nuisance macroalga Ulva to progressively reduce its spatial expansion in the Venice lagoon. In NW Europe, the North Atlantic Oscillation index (NAO) is one important indicator of climate change (Hurrell, 1995) and has been shown to correlate with biological cycles in biota abundance, including benthic macrofauna (Hagberg and Tunberg, 2000). When the NAO index is plotted against the long-term data set of macroalgal coverage in the Ythan estuary (data from Raffaelli et al., 1999), there is a significant positive correlation (Fig. 6). This suggests that the extensive growth of Enteromorpha recorded post-1990 in the UK estuaries is associated with the positive NAO values, which reflect the milder winters dominated by warmer westerly winds from the Atlantic. Consequently, there is increasing evidence that the development and persistence of macroalgal blooms may by partially driven by large-scale climatic factors. 3.3. Stabilisation of the upper shore 3.3.1. Effect of prolonged air exposure and exclusion of Macoma above high water neap tide level On the upper intertidal, above high water neap tide level (HWNT), there is a period of up to 8 days of continuous air exposure during the shift from spring

to neap tidal cycles. Flume erosion measurements (after sediment collection and re-immersion for 3 h) have shown a marked increase in sediment stability on the upper shore in the Humber estuary (Widdows et al., 2000b). Furthermore, experiments have demonstrated that this enhanced sediment stability is due to the combination of two factors, namely prolonged air exposure and de-watering as well as the absence of bioturbation by the clam M. balthica. The HWNT level represents the upper limit of the intertidal range and activity of M. balthica at high densities. Over the course of 8 days (May 1997), erosion measurements showed a significant increase in Ucrit from 0.18 to 0.30 m s 1 and this was accompanied by a 7-fold reduction in mass of sediment eroded at a current speed of 0.3 m s 1. This increased sediment stability during the neap tide cycle is likely to be enhanced further in response to elevated temperatures during the summer period. These sediment erodability measurements were in the absence of any bioturbation by Macoma. However, three hours after M. balthica individuals were added and allowed to bioturbate the surface sediment, the Ucrit was lowered from 0.30 to 0.19 m s 1 and sediment erodability was significantly increased by a factor of 4. Therefore approximately 50% of the recorded increase in sediment stability above the HWNT level is due to air exposure/desiccation and 50% is due to the absence or low density of the M. balthica. The increased flooding

Fig. 6. Relationship between the North Atlantic Oscillation Index (NAO) and Enteromorpha cover in intertidal areas of Ythan estuary from 1969 1997 (Enteromorpha data from Raffaelli et al., 1999; NAO data from J. Hurrell).

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of the upper shore during the neap to spring cycle will result in a gradual reduction in sediment stability (i.e. reduction in Ucrit from 0.30 to 0.18 m s 1). This rate of change has yet to be quantified, but the re-hydration of sediment and increase in erodability will occur over many tidal cycles and does not occur within a single cycle (i.e. after 3 h immersion). In contrast to M. balthica, the bioturbator Hydrobia ulvae can occur at high densities on the upper shore above the HWNT level, extending into the lower salt marsh. Therefore at sites where H. ulvae is abundant the sediment erodability may be greater than that recorded for the Humber upper shore during the neap tide cycle. 3.3.2. Salt marsh macrophytes Above the HWNT level, salt marsh plants represent the main biota dominating and influencing the structure and functioning of the high shore. The lower limits of pioneering salt marsh vegetation (e.g. Spartina anglica and Salicornia europaea) may be partially determined by grazing and bioturbatory activity (i.e. trophic interactions). For example, the presence of increasing densities of the amphipod (Corophium volutator) and the polychaete (Nereis diversicolor)

appears to induce negative effects on these salt marsh macrophytes (Gerdol and Hughes, 1993; Emmerson, 2000). Recent annular flume studies (Widdows et al., unpublished data) have quantified the effects of Spartina anglica stem densities on the hydrodynamics and sediment dynamics. There was a non-linear densitydependent relationship between depth-averaged current velocity and stem density. At densities >400 stems m 2 water flow was reduced by 75% in comparison with bare sediment, whereas at densities < 400 stems m 2 there was a marked increase in water flow with declining stem density (Fig. 7). This suggests that the surficial sediments are more vulnerable to erosion by currents when Spartina stem densities are below 400 stems m 2. These observations are consistent with the lowest recorded density of 400 stems m 2 found at the front margins of the salt marsh (Fig. 7) and the recorded scouring around small clumps of S. anglica in the Humber estuary (Brown, 1998). Erosion experiments (Fig. 8) showed that the critical erosion velocities were low in the presence and absence of Spartina stems (0.05 and 0.07 m s 1, respectively; the slightly lower threshold with Spartina stems was probably due to scouring

Fig. 7. Effect of varying Spartina anglica stem density (stems m 2; artificially planted) on depth-averaged current velocity (z = 0 to 0.1 m) in annular flume over a range of drive plate speeds (zero stem density represented by mud cored from in front of salt marsh). Horizontal dashed line represents range of stem densities from front edge to inner Spartina salt marsh at Blaxton creek, Tavy estuary, SW England. (Widdows et al., unpubl. data).

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Fig. 8. Comparison of sediment mass eroded and depth averaged current velocities (z = 0 to 0.1 m) in the presence of Spartina anglica stems (588 stems m 2; filled circles) and absence of stems (open circles) in relation to drive plate speed in the annular flumes. Spartina anglica stems were cut and removed without disturbing the sediment. Mass eroded (g m 2) expressed in relation to flume drive plate speed to demonstrate five-fold reduction in measured current velocity (via friction drag) caused by Spartina stems (mean F semi-range). Sediment with Spartina collected using flume quadrant cores from Blaxton salt marsh, Tavy estuary, SW England. (Widdows et al., unpubl. data).

effects). The major impact of Spartina stems was to reduce currents via friction drag and at a density of 588 stems m 2 there was a 5-fold reduction in nearbed current speeds (Fig. 8). Such quantification indicates the nature and scale of the reduction in sediment erodability provided by salt marsh plants, which form an important part of the natural coastal defence.

4. Overall impact of biota on intertidal sediment dynamics: causes and consequences The biological and physical processes occurring at the different intertidal levels, from lower shore to the upper shore and salt marsh, are not independent but are inter-related. Earlier studies have rarely measured and coupled processes on the mudflat with those on the salt marsh, but rather considered them as isolated and separate habitats. As part of the UK Land Ocean Interaction Study (LOIS) programme, multi-disciplinary studies carried out simultaneous measurements of biota, sediment erodability and bed-level changes on the Skeffling mudflat on the Humber estuary. This provided evidence of important coupling between events on the mudflat and the salt marsh, as well as

insight into the causes and consequences of temporal changes in sediment erodability in the intertidal zone (Widdows et al., 2000b). The key biological physical interactions and the mechanisms involved are summarised as follows: Climate is a major driver of interannual variation in biota, with cold winters resulting in an increase in M. balthica abundance on mudflats. This causes an increase in sediment erodability on the tidal flats, which in turn increases the SPM in the water column and this is transported onto the salt marsh on the high spring tides. As a result there is an increase in sediment accretion (values of 2.8 mm month 1 on the Skeffling (Humber) salt marsh in spring 1997; Widdows et al., 2000b). The converse situation occurs in years following mild winters when the mudflats tend to have welldeveloped diatom bio-films and low M. balthica abundance. This results in enhanced sediment stability due to bio-stabilisation and causes a reduction in shoreward sediment transport and accretion on the salt marsh (values of 0.3 mm month 1 in spring 1996). Intermittent storms and wave action may also be important physical processes that interact with the biotically modified sediment erodability, enhancing sediment transport onto the salt marsh. However, there

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is increasing evidence that storm events may be less important than expected. For example, recent field studies indicate that storms may only contribute 27% of the sediment deposited on salt marshes in Virginia, USA, during high spring tides (Christiansen, 1998). In addition, there was relatively little impact of a storm event on sediment properties and erodability on a tidal flat in the Westerschelde in late August 1996 (Widdows et al., 2000a). More recently, studies of sediment dynamics in open coastal areas such as Morecambe Bay (England) concluded that asymmetry of tidal currents was the dominant agent of sediment transport in the intertidal zone, with waves being of secondary importance (Mason and Garg, 2001).

5. Modelling of intertidal sediment dynamics As a result of flume studies quantifying the density-dependent relationships of key biota on hydrodynamics and sediment dynamics, it is now possible to improve the parameterisation of models of intertidal sediment dynamics and begin to incorporate biotic influences (e.g. Willows et al., 1998; Wood and Widdows, 2002). The BIOSED model (Wood and Widdows, 2002) is a simple 1D onshore offshore

model of water movement with a semi-empirical model of cohesive sediment erosion and deposition. To date, it has examined the effects of various tidal amplitudes and associated tidal currents, the impact of biota densities (M. balthica and microphytobenthic chlorophyll-a) and offshore sediment concentrations (external supply) on sediment erosion, transport and deposition along a shore-normal transect (Fig. 9). Model outputs show that changes in erosion and deposition caused by natural variation in biota densities were as large as those caused by changes in tidal currents over the spring-neap cycle, or by doubling external sediment supply. Seasonal variation in microphytobenthos alters deposition on the upper shore by a factor of two, and interannual variation in the bioturbator (M. balthica) changed net deposition by a factor of five. These initial modelling studies have been validated for the Skeffling mudflat (Humber estuary) and show that biotic influences on intertidal sediment dynamics are significant and play a role in determining sediment budgets over tidal to monthly timescales. Improved modelling of the biological physical processes determining sediment accretion/erosion on the intertidal shore will advance our understanding and forecasting the potential changes in natural coastal

Fig. 9. Model results of net erosion and deposition of sediment after one 3.5 m spring tide for four different biota scenarios: L,L - Low Macoma balthica (100 individuals m 2) and Low microphytobenthic chlorophyll-a (10 Ag g 1 dry wt); L,H - Low Macoma (100 m 2) and High chlorophyll-a (60 Ag g 1); H,L - High Macoma (1000 m 2) and Low chlorophyll-a (10 Ag g 1); H,H - High Macoma (1000 m 2) and High chlorophyll-a (30 Ag g 1). (From Wood and Widdows, 2002).

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defences, particularly in response to climate changes and sea-level rise. Key questions include: What are the consequences of global warming and a northerly shift in geographical distribution of key species? For example, will milder winters and warmer summers result in the reduction in abundance of M. balthica and the dominance of bio-stabilisers such as micro- and macroalgae, thus increasing sediment stability on the tidal flats and reduced accretion on salt marshes? How will the estuarine morphology/bathymetry adapt to sea-level rise and will important intertidal habitats be reduced in area? These questions concern large spatial and temporal scale problems and are logistically difficult to address by direct experiment or field studies. They will require the use of appropriate models incorporating relevant biological and physical processes to enable forecasting of changes in estuarine morphology.

and by NERC Core Research Programme (Estuarine and Coastal Function and Health) at PML.

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6. Conclusions The interactions between the biotic and abiotic factors and their influence on the behaviour of intertidal sediments and the resultant morphology are complex and poorly characterised. To date, the role of a few key species has been identified and quantified, but many other intertidal biota still remain to be studied. It is unknown what proportion of species in various benthic communities are bio-engineers and have a measurable impact on sediment dynamics. Lawton and Jones (1995) have suggested that all ecosystems are controlled and organised by a relatively small set of key species and abiotic processes; this hypothesis needs to be tested in intertidal environments. Lack of knowledge of the precise nature of biological sediment interaction coupled with a poor understanding of how to parameterise biological effects so that they can be incorporated into numerical sediment transport and geomorphological models is a major impediment to progress.

Acknowledgements This research was supported by the NERC Land Ocean Interaction Study programme, by the DEFRA funded BIOSED programme (contract no. AE0259),

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