Stefan G. Bolam

Dense aggregations of Pygospio elegans (Claparède): effect on macrofaunal community structure and sediments

Abstract Epibenthic biogenic structures such as polychaete tubes are conspicuous features of many marine soft-bottom habitats. This paper compares the benthic macrofauna in patches with high and low densities of the tube-dweller Pygospio elegans on intertidal sandflats in eastern Scotland (UK). The main aim of this study was to determine potential differences in the macrofaunal community structure, the size distribution of individual species and sediment properties. Multivariate data analyses revealed that the macrofaunal community composition (excluding P. elegans ) within patches was always significantly different from outside patches, mainly due to variability in the abundances of Cerastoderma edule and Corophium volutator . In addition to P. elegans , 5 taxa were sufficiently abundant for univariate analyses, 4 of these ( Capitella capitata , C. edule , Macoma balthica and C. volutator ) being significantly more abundant within P. elegans patches than in surrounding, non-patch sediments. The size distribution of P. elegans was significantly different between patches (bimodal distribution) and non-patches (skewed distribution). Similarly, there was a greater proportion of larger C. capitata individuals within patches compared to non-patch sediments. Sediment organic content and silt/clay fraction were always significantly higher in patch sediments while redox profiles showed no differences except at the end of the study period when the top 2 cm within patches were more positive and more negative at 4 cm. These results imply that even relatively small (1–1.5 m 2 ) P. elegans patches can have large effects on the spatial variability of macrofaunal community structure on intertidal sandflats. Towards the end of the study there were marked visual changes in the P. elegans patches, such as wave-ripple marks on the surface, which signified their demise. This coincided with dramatic changes in the invertebrate community structure within patches. Along with the decline in P. elegans numbers, dramatic increases in the densities of the 2 bivalve species C. edule and M. balthica occurred, while remaining comparatively stable outside patches. This suggested that the conditions within P. elegans patches were particularly favourable to successful bivalve spat settlement. We postulate that P. elegans patches have limited longevity and propose that enhanced bivalve competition within them leads to rapid decreases in P. elegans numbers.

Dense aggregations of tube-building polychaetes: response to small-scale disturbances

Many species of tube-building polychaetes form dense aggregations or ‘patches’ within marine soft-bottom habitats. Although the ecological importance of such patches has been well studied, the effect of small-scale disturbances within patches has not been investigated. On the intertidal sandflat Drum Sands, Firth of Forth, Scotland, the tube-building spionid Pygospio elegans Claparede forms patches approximately 1–1.5 m2, wherein its density is significantly greater than in surrounding sediments. This study explicitly investigates whether the initial communities colonising small-scale disturbances within P. elegans patches differ in terms of community structure and colonisation mode from those of surrounding sediments. Defaunated sediments (1/125 m2) were implanted into the sediments within and outside patches during April, August and December 1997, and sampled after 3 weeks. The results indicated that some species colonised patches and non-patches in significantly different numbers, at least at certain times of the year. Following these controlled disturbances, P. elegans larvae colonised patches in significantly higher numbers compared to non-patch areas during April 1997, but showed the opposite trend in August 1997 when larval recruitment was relatively low. Adult P. elegans colonised patch sediments in higher numbers than non-patch ones during August and December. Capitella capitata (Fabricius) consistently colonised all defaunated areas in relatively high numbers while Cerastoderma edule (L.) and Macoma balthica (L.) colonised non-patch azoic sediments in significantly higher numbers than patch azoics during April 1997. Multivariate analyses, together with nonparametric analysis of similarities (ANOSIM) tests, revealed that the community composition of the disturbed sediments was significantly different between patches and non-patches during April 1997 only. These analyses also indicated that there was a strong temporal effect on community composition since the April community was markedly different from those of August and December 1997. Sediment measurements taken during December suggested that the observed faunal differences were unlikely to have been a response to differences in bulk sediment characteristics or redox potential. It is proposed that the initial recolonisation differences were likely to have been due to the near-bed hydrodynamic effects of P. elegans tubes creating local sediment stabilisation. The ecological implications of differential recruitment following small-scale disturbances within patches compared to outside patches are discussed. The ability of P. elegans patches to enhance successful conspecific colonisation, as demonstrated here, may explain the longevity of such patches. Their persistence promotes alpha diversity and small-scale patchiness, together with providing a local source of colonists for the initial recovery of large-scale disturbances.

Impact and recovery associated with the deposition of capital dredgings at UK disposal sites: Lessons for future licensing and monitoring

The majority of studies relating to impacts and recovery at dredgings disposal sites have concentrated on areas subject to regular and frequent disposals of maintenance dredgings over relatively long time periods. In comparison less is known regarding the significance of impacts and the recovery processes associated with the disposal of capital dredgings that commonly involves the infrequent deposition of heterogenous material over relatively restricted time periods. Impacts and recovery processes are likely to be different to those associated with the disposal of maintenance dredgings. For example, findings suggest that capital dredgings deposited at both the Roughs Tower and Barrow-in-Furness result in the occurrence of persistent changes to seafloor substrata within the license area and this subsequently effects the composition of associated faunal communities present. Moreover, whilst the two disposal sites are geographically distinct similar species are identified as being particularly sensitive to capital disposal activities in both areas.

Spatial distribution of macrofaunal assemblages along the English Channel

The present study investigates the species distribution patterns and macrofaunal assemblages along the English Channel and the environmental factors contributing to observed patterns. Seven distinct macrofaunal assemblages were identified based on Hamon grab samples. In the western Channel, an Echinocyamus/Nemertea assemblage dominated, giving way to an Abra/ Scalibregma assemblage in inshore waters. A Verruca/Sabellaria assemblage was identified to occupy large regions of the mid-Channel while those of the east Channel were more spatially variable. While variations in depth, sediment particle size and wave and tidal stress were significantly related to variations in assemblage structure, temperature did not appear to be greatly influential. In addition, acoustic data obtained for each station sampled allowed us to determine how representative the observed biological communities are over spatial scales somewhat larger than the sampling points. In contrast to earlier comparable studies using anchor dredges, the biological communities sampled in this study did not exhibit a large east-west distinction; the most abundant species and key assemblages were found to be present along the length of the Channel. The possible reasons for these differences with earlier studies are discussed.

Impacts of dredged material disposal on macrobenthic invertebrate communities: A comparison of structural and functional (secondary production) changes at disposal sites around England and Wales

Although the impacts of dredged material disposal in the marine environment have been well studied, there is currently a limited understanding of the associated impacts on benthic function. This study compares macrofaunal structural and functional (based on secondary production estimates) responses to dredged material disposal at 14 sites across the coast of England and Wales. Disposal resulted in significant reductions of total secondary production at seven sites; no sites exhibited significant increases in production estimates. These seven sites were generally those which displayed significant structural impacts. There was no clear relationship between multivariate structural changes and taxonomic contribution to total production, indicating that a lack of change in the former (regarded as a sensitive indicator of change) does not always signify a lack of a significant functional impact. The need to evaluate functional changes, in addition to structural impacts, with respect to dredged material disposal site monitoring is discussed.

An approach for the identification of exemplar sites for scaling up targeted field observations of benthic biogeochemistry in heterogeneous environments

Continental shelf sediments are globally important for biogeochemical activity. Quantification of shelf-scale stocks and fluxes of carbon and nutrients requires the extrapolation of observations made at limited points in space and time. The procedure for selecting exemplar sites to form the basis of this up-scaling is discussed in relation to a UK-funded research programme investigating biogeochemistry in shelf seas. A three-step selection process is proposed in which (1) a target area representative of UK shelf sediment heterogeneity is selected, (2) the target area is assessed for spatial heterogeneity in sediment and habitat type, bed and water column structure and hydrodynamic forcing, and (3) study sites are selected within this target area encompassing the range of spatial heterogeneity required to address key scientific questions regarding shelf scale biogeochemistry, and minimise confounding variables. This led to the selection of four sites within the Celtic Sea that are significantly different in terms of their sediment, bed structure, and macrofaunal, meiofaunal and microbial community structures and diversity, but have minimal variations in water depth, tidal and wave magnitudes and directions, temperature and salinity. They form the basis of a research cruise programme of observation, sampling and experimentation encompassing the spring bloom cycle. Typical variation in key biogeochemical, sediment, biological and hydrodynamic parameters over a pre to post bloom period are presented, with a discussion of anthropogenic influences in the region. This methodology ensures the best likelihood of site-specific work being useful for up-scaling activities, increasing our understanding of benthic biogeochemistry at the UK-shelf scale.

Development of indicators of ecosystem functioning in a temperate shelf sea: a combined fieldwork and modelling approach

A conceptual model of the main carbon and nitrogen flows through pelagic and benthic food webs was used to identify the key biogeochemical processes representing ecosystem functioning, and to select indicators of each of these processes. A combined fieldwork and modelling approach was used to provide the data required to evaluate the indicators in terms of their suitability for assessing and managing the impacts of climate change and demersal trawling. Four of our 16 proposed indicators (phytoplankton production and productivity, near-bed oxygen concentrations and oxygen penetration of the seabed) met the majority of criteria we used for evaluating indicators. Five indicators (depth of anoxic sediment, zoobenthos biomass, production, productivity and bioturbation potential) did not comply with sufficient criteria to be considered as good indicators. Six of our proposed indicators (zooplankton biomass, size structure, production and productivity; ecosystem productivity; ecosystem balance) could not be assessed for sensitivity and specificity using our models, and therefore need to be addressed in future work aimed at improving both the models and the fieldwork. Our results indicate that evaluation of indicators is difficult, because of the number and variety of human pressures which need to be considered in reality, and the interactions between these pressures and the ecosystem components which they affect. The challenge will be to establish if there are indeed any indicators which are able to meet the majority of criteria for good indicators in holistic ecosystem-based assessments.

Impacts of maintenance dredged material disposal on macrobenthic structure and secondary productivity

The results of a monitoring programme to assess the spatial impacts associated with ongoing dredged material disposal activity at a dispersive, coastal disposal site (southwest UK) are described. Benthic impacts were assessed using benthic community structure and secondary productivity estimates. Analyses of univariate indices (including secondary production) and multivariate community structure revealed differences between stations inside and those outside the disposal site were minimal. Generally, stations within and outside the disposal site were characterised by the same species. Regression models indicated that the variability in biological structure and secondary production was predominantly accounted for by natural variables (e.g., depth, sediment granulometry) with only a small amount of residual variability being due to contaminant variables. Thus, the elevated levels of certain contaminants in the vicinity of the disposal area were not sufficient to result in significant ecological or ecotoxicological changes. We ascribe such findings partly to the dispersive nature of the disposal site.

Application of biological traits to further our understanding of the impacts of dredged material disposal on benthic assemblages

While the effects of coastal disposal of dredged material on benthic assemblage structure have been well studied, our understanding of the mechanism of such responses, and their potential ecological implications, remain relatively unknown. Data from a licenced disposal site off the northeast coast of England are analysed to address this and improve our ability to make informed licencing decisions for this activity. Assemblages within the disposal site displayed reduced number of species and total invertebrate density, an altered assemblage taxonomic structure, and a shift towards a greater numerical dominance of less-productive individuals. Following separate analyses of biological response and effect traits, a novel approach for marine benthic trait analysis, we identify the traits responsible (i.e. response traits) for the observed structural alterations. Furthermore, analysis of the effect traits revealed that the assemblages characterising the disposal site possess a greater bioturbative capability compared to those not directly impacted by disposal.

Towards an integrated approach to marine benthic monitoring.

In the UK, most marine benthic monitoring is carried out in a piecemeal fashion, funded by different sectors of industry that utilise the marine environment under licence. Monitoring requirements are imposed by licence conditions, which can vary considerably between licences. The UK Government also conducts marine environmental surveys in support of its legislative commitments. The present investigation reviews these different monitoring approaches to highlight whether synergies between them could be developed into an integrated approach to marine benthic monitoring. An integrated approach would have ecological benefits, as greater consistency in sampling and analytical protocols would reduce uncertainty in the predictions of impact, and facilitate the assessment of Good Environmental Status under the Marine Strategy Framework Directive. The same approach would also be of financial benefit, as spatio-temporal duplication in sampling would be reduced, and the value of acquired data would be maximised, resulting in a more efficient and cost-effective approach.