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JNCC ReportNo: 557

Sublittoral Mud CEM literature review and ancillary info v 1.0

Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection

Coates, D.A., Alexander, D., Stafford, R. & Herbert, R.J.H.

Jun-15

© JNCC, Peterborough 2015

ISSN 0963 8901

Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection

1. Overview

2. Habitat Characterisation

3. Species Selection

4. Trait Selection

5. Faunal Traits Matrix

6. Faunal Traits Summary

This spreadsheet forms part of the project "Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0860) accompanies the report of the same title. Ancillary information which supports and informs the project is contained within the worksheets presented, in addition to the results of the literature review. A description of the information presented in each worksheet is described in the section headings below.

This worksheet contains physical and chemical characterising information for each biotope type using information from the EUNIS classification and Marine Habitat Classification for Britain and Ireland (both based on Connor et al. 2004).

This worksheet is split into two sections, 3a and 3b, and indicates how species were selected from the biotopes and refined for the purposes of the project. The species list presented in worksheet 3a is a complete list of taxa taken from the biotope information pages made available through the EUNIS classification and the Marine Habitat Classification for Britain and Ireland (both based on Connor et al. 2004). A total of 155 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed. Worksheet 3b indicates the process used to refine the full list of 155 species to a manageable number for inclusion in the project. Species were selected for each named biotope using an iterative approach. Initially those species listed in the biotope names were included (Columns B to D), followed by any important species listed in the biotope descriptions (Columns E to L). The resulting revised list of 53 species to be included in the project is presented below the table.

Worksheet 4 presents the species traits that were included within the literature review. Species traits were used in order to inform links within the models and to classify functional groups. A list of 41 common species traits was extracted from the MarLIN BIOTIC Database (MarLIN, 2006), which was supplemented by 6 additional traits added by the project team. This list was refined to an appropriate number of traits for inclusion in the project using expert judgement. An indication of whether the trait was used in the project and a short rationale for each is presented in Columns B and C. Definitions of each trait and a list of trait categories (for those which have standard responses) are presented in Worksheet 10.

The Faunal Traits Matrix presents the results of the literature review for the species and traits selected in Worksheets 3 and 4. Data are entered so that one row represents one reference (identified by the reference code in Column Z). As such, there are multiple rows per species listed, although if one source yielded several pieces of information there may be several entries per row. Confidence in the source is included in Column AA, see Worksheets 8 and 9 for details on how this was assigned.

A summary of the information collected as part of the species traits literature review is presented in Worksheet 6. This Worksheet informs the gap analysis and identifies data gaps in the project, as well as indicating where expert judgement has been used to inform species traits. Entries in the matrix are colour coded according to the legend underneath the table to facilitate interpretation. A list of biotopes relevant to each species is also presented (note that the biotopes listed are those which fauna have been assessed to represent for the purposes of this project, as outlined in Worksheet 3b. It is likely that species are found in other biotopes not identified by this project due to the high degree of species overlaps between biotopes).

7. Interactions Matrix

8. Reference Summary

9. Confidence Assessment

10. Definitions

For More Information

Version Control

BUILD STATUS:

Worksheet 7 contains the Interactions Matrix. This forms the main part of the literature review results and provides information on the relevant environmental drivers, ecosystem functions and ecosystem processes which occur in shallow sublittoral mud habitats around the UK. Information is arranged so that one row represents one piece of information/one link within a model. Several columns provide metadata used for filtering the table when building the output models. The reference code for each row is provided in Column A (see Worksheet 8 for details). Column B provides information on the model level the information informs (see Worksheet 10 for descriptions of model levels and associated report and output models). The model parameter assessed (Column C) refers to the specific model component the information refers to (see Worksheet 10 for definitions). Column D indicates which model component is influenced by the information, and Column E to any specific species named in the information (if applicable). Column F contains metadata on the direction of interaction (input/output or feedback). Column G indicates whether the interaction is positive or negative (if applicable) and Column H shows the magnitude of Interaction. Definitions and criteria for the direction and magnitude of interactions are presented in Worksheet 10. Any limitations of the information are presented in Column I. A summary of the information taken from the source is shown in Column J, along with the source scale or location in Column K. Any additional reference which may be relevant to the source are shown in Column N. The source confidence score is included as the last column, please see Worksheet 9 and the accompanying report for details of how this was assigned.

A full reference list of all sources is presented in Worksheet 8. A unique reference code is assigned to each source which is used to identify the source in all worksheets. The full source reference is presented in Column B, along with the unedited source abstract in Column C. A summary of the information taken from the source is included in Column D (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns E and F in accordance with the criteria laid out in Worksheet 9 and the accompanying report.

Worksheet 9 presents the confidence assessment used for assigning confidence to individual sources. Confidence is taken as the lowest common denominator for both quality and applicability. Please see accompanying report for full confidence methodology.

Definitions of key project features are presented in Worksheet 10. For more information regarding how these features relate to the project, please see accompanying report.

This report should be cited as:

Coates, D.A., Alexander, D., Stafford, R. and Herbert, R.J.H. 2015. Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection. Marine Ecological Surveys Ltd - A report for the Joint Nature Conservation Committee, JNCC Report No: [Report No.].

For further information please contact:

Joint Nature Conservation CommitteeMonkstone HouseCity RoadPeterborough PE1 1JYwww.jncc.defra.gov.uk

Version Date Author0.1 17/11/2014 DC0.2 17/02/2014 DC1.0 06/03/2014 DC

DISTRIBUTION:

Copy Version Issue DateElectronic 1.0 06/03/2014

This spreadsheet forms part of the project "Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0860) accompanies the report of the same title. Ancillary information which supports and informs the project is contained within the worksheets presented, in addition to the results of the literature review. A description of the information presented in each worksheet is described in the section headings below.

This worksheet contains physical and chemical characterising information for each biotope type using information from the EUNIS classification and Marine Habitat Classification for Britain and Ireland (both based on Connor et al. 2004).

This worksheet is split into two sections, 3a and 3b, and indicates how species were selected from the biotopes and refined for the purposes of the project. The species list presented in worksheet 3a is a complete list of taxa taken from the biotope information pages made available through the EUNIS classification and the Marine Habitat Classification for Britain and Ireland (both based on Connor et al. 2004). A total of 155 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed. Worksheet 3b indicates the process used to refine the full list of 155 species to a manageable number for inclusion in the project. Species were selected for each named biotope using an iterative approach. Initially those species listed in the biotope names were included (Columns B to D), followed by any important species listed in the biotope descriptions (Columns E to L). The resulting revised list of 53 species to be included in the project is presented below the table.


The Faunal Traits Matrix presents the results of the literature review for the species and traits selected in Worksheets 3 and 4. Data are entered so that one row represents one reference (identified by the reference code in Column Z). As such, there are multiple rows per species listed, although if one source yielded several pieces of information there may be several entries per row. Confidence in the source is included in Column AA, see Worksheets 8 and 9 for details on how this was assigned.

A summary of the information collected as part of the species traits literature review is presented in Worksheet 6. This Worksheet informs the gap analysis and identifies data gaps in the project, as well as indicating where expert judgement has been used to inform species traits. Entries in the matrix are colour coded according to the legend underneath the table to facilitate interpretation. A list of biotopes relevant to each species is also presented (note that the biotopes listed are those which fauna have been assessed to represent for the purposes of this project, as outlined in Worksheet 3b. It is likely that species are found in other biotopes not identified by this project due to the high degree of species overlaps between

Worksheet 7 contains the Interactions Matrix. This forms the main part of the literature review results and provides information on the relevant environmental drivers, ecosystem functions and ecosystem processes which occur in shallow sublittoral mud habitats around the UK. Information is arranged so that one row represents one piece of information/one link within a model. Several columns provide metadata used for filtering the table when building the output models. The reference code for each row is provided in Column A (see Worksheet 8 for details). Column B provides information on the model level the information informs (see Worksheet 10 for descriptions of model levels and associated report and output models). The model parameter assessed (Column C) refers to the specific model component the information refers to (see Worksheet 10 for definitions). Column D indicates which model component is influenced by the information, and Column E to any specific species named in the information (if applicable). Column F contains metadata on the direction of interaction (input/output or feedback). Column G indicates whether the interaction is positive or negative (if applicable) and Column H shows the magnitude of Interaction. Definitions and criteria for the direction and magnitude of interactions are presented in Worksheet 10. Any limitations of the information are presented in Column I. A summary of the information taken from the source is shown in Column J, along with the source scale or location in Column K. Any additional reference which may be relevant to the source are shown in Column N. The source confidence score is included as the last column, please see Worksheet 9 and the accompanying report for details of how this was assigned.

A full reference list of all sources is presented in Worksheet 8. A unique reference code is assigned to each source which is used to identify the source in all worksheets. The full source reference is presented in Column B, along with the unedited source abstract in Column C. A summary of the information taken from the source is included in Column D (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns E and F in accordance with the criteria laid out in Worksheet 9 and

Worksheet 9 presents the confidence assessment used for assigning confidence to individual sources. Confidence is taken as the lowest common denominator for both quality and applicability. Please see accompanying report for full confidence methodology.

Definitions of key project features are presented in Worksheet 10. For more information regarding how these features relate to the project, please


For further information please contact:

Joint Nature Conservation Committee

Reason/CommentsDraft version submitted for commentsFinal Draft version submitted for commentsFinal version submitted to client

Issued ToJNCC

This spreadsheet forms part of the project "Conceptual Ecological Modelling of Shallow Sublittoral Mud Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0860) accompanies the report of the same title. Ancillary information which supports and informs the project is contained within the worksheets presented, in addition to the results of the literature review. A description of the information presented in each worksheet is

This worksheet contains physical and chemical characterising information for each biotope type using information from the EUNIS classification and

This worksheet is split into two sections, 3a and 3b, and indicates how species were selected from the biotopes and refined for the purposes of the project. The species list presented in worksheet 3a is a complete list of taxa taken from the biotope information pages made available through the EUNIS classification and the Marine Habitat Classification for Britain and Ireland (both based on Connor et al. 2004). A total of 155 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed. Worksheet 3b indicates the process used to refine the full list of 155 species to a manageable number for inclusion in the project. Species were selected for each named biotope using an iterative approach. Initially those species listed in the biotope names were included (Columns B to D), followed by any important species listed in the biotope descriptions (Columns E to L).


The Faunal Traits Matrix presents the results of the literature review for the species and traits selected in Worksheets 3 and 4. Data are entered so that one row represents one reference (identified by the reference code in Column Z). As such, there are multiple rows per species listed, although if one source yielded several pieces of information there may be several entries per row. Confidence in the source is included in Column AA, see

A summary of the information collected as part of the species traits literature review is presented in Worksheet 6. This Worksheet informs the gap analysis and identifies data gaps in the project, as well as indicating where expert judgement has been used to inform species traits. Entries in the matrix are colour coded according to the legend underneath the table to facilitate interpretation. A list of biotopes relevant to each species is also presented (note that the biotopes listed are those which fauna have been assessed to represent for the purposes of this project, as outlined in Worksheet 3b. It is likely that species are found in other biotopes not identified by this project due to the high degree of species overlaps between

Worksheet 7 contains the Interactions Matrix. This forms the main part of the literature review results and provides information on the relevant environmental drivers, ecosystem functions and ecosystem processes which occur in shallow sublittoral mud habitats around the UK. Information is arranged so that one row represents one piece of information/one link within a model. Several columns provide metadata used for filtering the table when building the output models. The reference code for each row is provided in Column A (see Worksheet 8 for details). Column B provides information on the model level the information informs (see Worksheet 10 for descriptions of model levels and associated report and output models). The model parameter assessed (Column C) refers to the specific model component the information refers to (see Worksheet 10 for definitions). Column D indicates which model component is influenced by the information, and Column E to any specific species named in the information (if applicable). Column F contains metadata on the direction of interaction (input/output or feedback). Column G indicates whether the interaction is positive or negative (if applicable) and Column H shows the magnitude of Interaction. Definitions and criteria for the direction and magnitude of interactions are presented in Worksheet 10. Any limitations of the information are presented in Column I. A summary of the information taken from the source is shown in Column J, along with the source scale or location in Column K. Any additional reference which may be relevant to the source are shown in Column N. The source confidence score is included as the last column, please see Worksheet 9 and the accompanying report for

A full reference list of all sources is presented in Worksheet 8. A unique reference code is assigned to each source which is used to identify the source in all worksheets. The full source reference is presented in Column B, along with the unedited source abstract in Column C. A summary of the information taken from the source is included in Column D (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns E and F in accordance with the criteria laid out in Worksheet 9 and

Worksheet 9 presents the confidence assessment used for assigning confidence to individual sources. Confidence is taken as the lowest common

Definitions of key project features are presented in Worksheet 10. For more information regarding how these features relate to the project, please


Biotope MNCR CodeA5.33 SS.SMu.ISaMuA5.331 SS.SMu.ISaMu.NhomMacA5.332 SS.SMu.ISaMu.SundAasp A5.333 SS.SMu.ISaMu.MysAbrA5.334 SS.SMu.ISaMu.MelMagThyA5.335 SS.SMu.ISaMu.AmpPlonA5.336 SS.SMu.ISaMu.CapA5.34 SS.SMu.IFiMuA5.341 SS.SMu.IFiMu.CerAnitA5.342 SS.SMu.IFiMu.AreA5.343 SS.SMu.IFiMu.PhiVirA5.344 SS.SMu.IFiMu.OcnA5.35 SS.SMu.CSaMuA5.351 SS.SMu.CSaMu.AfilMysAnit A5.352 SS.SMu.CSaMu.ThyNtenA5.353 SS.SMu.CSaMu.AfilNten A5.354 SS.SMu.CSaMu.VirOphPmax A5.355 SS.SMu.CSaMu.LkorPpelA5.36 SS.SMu.CFiMuA5.361 SS.SMu.CFiMu.SpnMeg A5.362 SS.SMu.CFiMu.MegMaxA5.363 SS.SMu.CFiMu.BlyrAchi

Biotope NameA5.33 - Infralittoral sandy mud

A5.34 - Infralittoral fine mud

A5.35 - Circalittoral sandy mud

A5.36 - Circalittoral fine mudA5.361 - Seapens and burrowing megafauna in circalittoral fine mud

A5.331 - Nephtys hombergii and Macoma balthica in infralittoral sandy mudA5.332 - Sagartiogeton undatus and Ascidiella aspersa on infralittoral sandy mudA5.333 - Mysella bidentata and Abra spp. in infralittoral sandy mudA5.334 - Melinna palmata with Magelona spp. and Thyasira spp. in infralittoral sandy mudA5.335 - Ampelisca spp., Photis longicaudata and other tube-building amphipods and polychaetes in infralittoral sandy mudA5.336 - Capitella capitata in enriched sublittoral muddy sediments

A5.341 - Cerastoderma edule with Abra nitida in infralittoral mudA5.342 - Arenicola marina in infralittoral mudA5.343 - Philine aperta and Virgularia mirabilis in soft stable infralittoral mudA5.344 - Ocnus planci aggregations on sheltered sublittoral muddy sediment

A5.351 - Amphiura filiformis, Mysella bidentata and Abra nitida in circalittoral sandy mudA5.352 - Thyasira spp. and Nuculoma tenuis in circalittoral sandy mudA5.353 - Amphiura filiformis and Nuculoma tenuis in circalittoral and offshore sandy mudA5.354 - Virgularia mirabilis and Ophiura spp. with Pecten maximus on circalittoral sandy or shelly mudA5.355 - Lagis koreni and Phaxas pellucidus in circalittoral sandy mud

A5.362 - Burrowing megafauna and Maxmuelleria lankesteri in circalittoral mudA5.363 - Brissopsis lyrifera and Amphiura chiajei in circalittoral mud

Substratum

Sandy muds

Mud and gravelly mud

Stones or shells on muddy sediment

Mud with a fine to very fine sand fraction Sandy mud sandy mud Muddy sands and sandy muds Sandy mud

Muddy sediment Mud (occasionally with shells or stones)

Mud Mud occasionally with small stones

Mud with a significant fine to very fine sand fraction Sandy mud Mud occasionally with scattered shells or gravel Sandy mud Sandy mud; shelly and gravelly mud Sandy mud Mud Mud Mud Silty mud

Wave exposure

Moderately exposed, Sheltered, Extremely shelteredSheltered, Very sheltered, Extremely sheltered

Sheltered, Very sheltered, Extremely sheltered Very sheltered Sheltered, Very sheltered

Sheltered, Very sheltered, Extremely sheltered Sheltered, Very sheltered, Extremely sheltered Very sheltered Very sheltered, Extremely sheltered Sheltered, Very sheltered, Extremely sheltered Moderately exposed, Sheltered, Very sheltered, Extremely sheltered Exposed, Moderately exposed, Sheltered, Very sheltered Exposed, Moderately exposed Moderately exposed, Sheltered Moderately exposed Moderately exposed, Sheltered, Very sheltered Exposed, Moderately exposed Moderately exposed, Sheltered, Very sheltered, Extremely sheltered Moderately exposed, Sheltered, Very sheltered, Extremely sheltered Sheltered, Very sheltered, Extremely sheltered Moderately exposed, Sheltered, Very sheltered

Tidal streams

Weak (>1 kn)Moderately strong (1-3 kn), Weak (>1 kn), Very weak (negligible)

Moderately strong (1-3 kn), Weak (>1 kn), Very weak (negligible) Very weak (negligible) Moderately strong (1-3 kn)

Moderately strong (1-3 kn), Weak (>1 kn) Weak (>1 kn), Very weak (negligible) Weak (>1 kn) Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Moderately strong (1-3 kn), Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Very weak (negligible) Very weak (negligible) Weak (>1 kn), Very weak (negligible) Strong (3-6 kn), Moderately strong (1-3 kn), Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible) Weak (>1 kn), Very weak (negligible)

Zone Salinity

Infralittoral Full (30-35 ppt), Variable (18-35 ppt)Infralittoral Full (30-35 ppt), Variable (18-35 ppt)

Circalittoral, Infralittoral Full (30-35 ppt), Variable (18-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt), Variable (18-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt), Variable (18-35 ppt), Low (<18ppt) Infralittoral Full (30-35 ppt), Variable (18-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt), Variable (18-35 ppt) Circalittoral, Infralittoral Full (30-35 ppt), Variable (18-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt), Variable (18-35 ppt) Circalittoral Full (30-35 ppt), Variable (18-35 ppt) Circalittoral Full (30-35 ppt), Variable (18-35 ppt) Circalittoral Full (30-35 ppt)

Depth Band Reference code0-5 m, 5-10 m, 10-20 m R15-10 m, 10-20 m R1

R1R1R1R1R1R1R1R1R1R1R1R1R1R1R1

10-20 m, 20-30 m, 30-50 m, 50-100 m R1R1R1R1R1

0-5 m, 5-10 m, 10-20 m 0-5 m, 5-10 m, 10-20 m 5-10 m, 10-20 m

0-5 m, 5-10 m, 10-20 m 0-5 m, 5-10 m, 10-20 m 0-5 m, 5-10 m 0-5 m, 5-10 m 0-5 m, 5-10 m, 10-20 m 0-5 m, 5-10 m, 10-20 m, 20-30 m 5-10 m, 10-20 m, 20-30 m, 30-50 m, 50-100 m 10-20 m, 20-30 m 20-30 m, 30-50 m, 50-100 m 50-100 m 5-10 m, 10-20 m, 20-30 m

10-20 m, 20-30 m, 30-50 m 10-20 m, 20-30 m, 30-50 m 10-20 m, 20-30 m, 30-50 m, 50-100 m 20-30 m, 30-50 m, 50-100 m

Species UK Status/RarityHydractinia echinata Presumed nativeCerianthus lloydii Presumed nativeMetridium senile Presumed nativeSagartiogeton undatus Presumed nativeNemerteaNematodaNephtys hombergii Presumed nativeScoloplos armiger Presumed nativeMagelona filiformis Presumed nativeChaetozone gibber Presumed nativeCapitella capitata Presumed nativeArenicola marina Presumed nativeEuclymene oerstedii Presumed nativeMelinna palmata Presumed nativeTerebellidaeMyxicola infundibulum Presumed nativeAmpelisca brevicornis Presumed nativeAmpelisca tenuicornis Presumed nativePagurus bernhardus Presumed nativeLiocarcinus depurator Presumed nativeCarcinus maenas Presumed nativeNucula nitidosa Presumed nativeThyasira flexuosa Presumed nativeKurtiella bidentataMacoma balthica Presumed nativeAbra alba Presumed nativeAsterias rubens Presumed nativeAscidiella aspersa Presumed nativePomatoschistusNephtys kersivalensis Presumed nativeCirratulidaeMediomastus fragilis Presumed nativeDexamine thea Presumed nativeMicroprotopus maculatus Presumed nativeAoridaePariambus typicus Presumed nativeAbra nitida Presumed nativeMya spp.Zostera marinaVirgularia mirabilis Presumed nativeChaetopterus variopedatus Presumed nativeLanice conchilega Presumed nativeTurritella communis Presumed nativePecten maximus Presumed nativeAmphiura chiajei Presumed nativeAmphiura filiformis Presumed nativeOphiura albida Presumed nativeOphiura ophiura Presumed nativeEchinus esculentus Presumed native; IUCN: Lower risk(LR / nt)Funiculina quadrangularis Presumed nativePennatula phosphorea Presumed nativeNephtys hystricis Presumed nativeChaetozone setosa Presumed native

Nephrops norvegicus Presumed nativeMunida rugosa Presumed nativeMaxmuelleria lankesteri Presumed nativeOphiodromus flexuosus Presumed nativeCalocaris macandreae Presumed nativeJaxea nocturna Presumed nativeCallianassa subterranea Presumed nativeBuccinum undatum Presumed nativeCorbula gibba Presumed nativeLesueurigobius friesii Presumed nativePomatoschistus minutus Presumed nativePectinaria belgica Presumed nativeBrissopsis lyrifera Presumed nativeSagartiogeton laceratus Presumed nativePhiline aperta Presumed nativeAequipecten opercularis Presumed nativeSaccharina latissimaAlcyonium digitatum Presumed nativeMya truncata Presumed nativeOphiothrix fragilis Presumed nativeOphiocomina nigra Presumed nativeOcnus lacteus Presumed nativeOcnus planci Presumed nativeSpio filicornis Presumed nativeMagelona alleni Presumed nativeAphelochaeta marioni Presumed nativeTharyx spp.Notomastus latericeus Presumed nativeGalathowenia oculata Presumed nativeAmpharete lindstroemi Presumed nativeHarpinia antennaria Presumed nativePhaxas pellucidus Presumed nativePhoronis spp.Kirchenpaueria pinnata Presumed nativeNemertesia ramosa Presumed nativeThysanocardia procera Presumed nativePholoe baltica (sensu petersen)Nephtys incisa Presumed nativeLumbrineris gracilis Presumed nativeDiplocirrus glaucus Presumed nativeScalibregma inflatum Presumed nativeOwenia fusiformis Presumed nativeLagis koreni Presumed nativePomatoceros triqueter Presumed nativePagurus prideaux Presumed nativeAporrhais pespelecani Presumed nativeNuculoma tenuis Presumed nativeOphiuroideaGlycera rouxii Presumed nativeGoniada maculata Presumed nativeNephtys spp.Abyssoninoe hibernicaParaonidae spp.Spiophanes bombyx Presumed native

Spiophanes kroyeri Presumed nativeRhodine gracilior Presumed nativeAnobothrus gracilis Presumed nativeTerebellides stroemi Presumed nativeCylichna cylindracea Presumed nativeLucinoma borealis Presumed nativeThyasira spp.Mya arenaria Alien in UKHydrallmania falcata Presumed nativeSertularia argentea Presumed nativeObelia longissima Presumed nativeOphelina acuminata Presumed nativeAntalis entalis Presumed nativeCellaria fistulosa Presumed nativeEchinocardium flavescens Presumed nativeGobius niger Presumed nativeNephtys cirrosa Presumed nativeMagelona mirabilis Presumed nativeEchinocardium cordatum Presumed nativeMalacoceros fuliginosus Presumed nativePolydora ciliata Presumed nativeOligochaetaTubificoides benedii Presumed nativeHediste diversicolor Presumed nativeChaetozone caputesocisHydrobia ulvae Presumed nativeCerastoderma edule Presumed nativeTubularia indivisa Presumed nativePhascolion strombus strombusPolychaetaAphrodita aculeata Presumed nativeLevinsenia gracilis Presumed nativePectinariidaeNereiphylla lutea Presumed nativePrionospio ehlersi Presumed nativeSpio spp.Chaetozone spp.Amphictene auricoma Presumed nativePseudocuma longicornis Presumed nativeMacropodia rostrata Presumed nativeCorystes cassivelaunus Presumed nativePsammechinus miliaris Presumed nativeCallionymus lyra Presumed nativePleuronectidaeBeggiatoa spp.Diatoms - filmChlamys spp.Gobiidae

Biotopes to be considered:

A5.33 - Infralittoral sandy mud

A5.34 - Infralittoral fine mud

A5.35 - Circalittoral sandy mud

A5.36 - Circalittoral fine mudA5.361 - Seapens and burrowing megafauna in circalittoral fine mud

Fauna selected for inclusion in project:Abra albaAbra nitidaAmpelisca tenuicornisAmpharete lindstroemiAmphiura filiformisAphelochaeta marioniArenicola marinaBrissopsis lyriferaCallianassa subterraneaCalocaris macandreaeCapitella capitataCarcinus maenasCerastoderma eduleCirriformia tentaculataEchinocardium cordatumEchinus escluentusEuclymene oerstediiGalathowenia oculataGoniada maculateHediste diversicolorLabidoplax mediaLagis koreniLeptosynapta bergensisMacoma balthica

A5.331 - Nephtys hombergii and Macoma balthica in infralittoral sandy mudA5.332 - Sagartiogeton undatus and Ascidiella aspersa on infralittoral sandy mudA5.333 - Mysella bidentata and Abra spp. in infralittoral sandy mudA5.334 - Melinna palmata with Magelona spp. and Thyasira spp. in infralittoral sandy mudA5.335 - Ampelisca spp., Photis longicaudata and other tube-building amphipods and polychaetes in infralittoral sandy mudA5.336 - Capitella capitata in enriched sublittoral muddy sediments

A5.341 - Cerastoderma edule with Abra nitida in infralittoral mudA5.342 - Arenicola marina in infralittoral mudA5.343 - Philine aperta and Virgularia mirabilis in soft stable infralittoral mudA5.344 - Ocnus planci aggregations on sheltered sublittoral muddy sediment

A5.351 - Amphiura filiformis, Mysella bidentata and Abra nitida in circalittoral sandy mudA5.352 - Thyasira spp. and Nuculoma tenuis in circalittoral sandy mudA5.353 - Amphiura filiformis and Nuculoma tenuis in circalittoral and offshore sandy mudA5.354 - Virgularia mirabilis and Ophiura spp. with Pecten maximus on circalittoral sandy or shelly mudA5.355 - Lagis koreni and Phaxas pellucidus in circalittoral sandy mud

A5.362 - Burrowing megafauna and Maxmuelleria lankesteri in circalittoral mudA5.363 - Brissopsis lyrifera and Amphiura chiajei in circalittoral mud

Magelona johnstoniMalacoceros fuliginosusMaxmuelleria lankesteriMediomastus fragilisMelinna palmataMicroprotopus maculatusMya truncataMysella bidentataNephrops norvegicusNephtys hombergiiNuculoma tenuisOcnus planciOwenia fusiformisPagurus bernhardusPhaxas pellucidusPhiline aperta

Phoronis muelleriPhotis longicaudataPolydora ciliataPygospio elegansRhodine graciliorSagartiogeton undatusScalibregma inflatumScoloplos armigerSpiophanes bombyxThysanocardia proceraTubificoides (pseudogaster)Virgularia mirabilis

Pholoe inornata (sensu petersen)

Title Species 1 Title Species 2 Title Species 3

Nephtys hombergii Macoma balthicaSagartiogeton undatusMysella bidentataMelinna palmata Magelona Thyasira flexuosaAmpelisca Photis longicaudataCapitella capitata

Cerastoderma edule Abra nitidaArenicola marinaPhiline aperta Virgularia mirabilisOcnus planci

Amphiura filiformis Mysella bidentata Abra nitidaThyasira flexuosa Nuculoma tenuisAmphiura filiformis Nuculoma tenuisVirgularia mirabilis Ophiura albida Ophiura ophiuraLagis koreni Phaxas pellucidus

Maxmuelleria lankesteriBrissopsis lyrifera Amphiura chiajei

Ascidiella aspersaAbra

Description Species 1 Description Species 2 Description Species 3

Abra alba Nucula nitidosa Spiophanes bombyxCarcinus maenas Pagurus bernhardus terebellid polychaetesAmpelisca Scoloplos armiger MyaChaetozone gibber Nephtys hombergii Galathowenia oculataLagis koreni Nucula nitidosa Chamelea gallinaMalacoceros fuliginosus Tubificoides Cirriformia tentaculata

Hydrobia ulvae Hediste diversicolorLabidoplax media Leptosynapta bergensis

Philine aperta

Thysanocardia procera Nephtys incisa PhoronisGoniada maculate Rhodine graciliorOphiura albida Echinocardium flavescens Mysella bidentataPecten maximusMysella bidentata Abra alba Mediomastus fragilis

Nephrops norvegicus Virgularia mirabilis Pennatula phosphoreaNephrops norvegicus Calocaris macandreae Callianassa subterranea

Caulleriella

Description Species 4 Description Species 5 Description Species 6

Lagis koreni Echinocardium cordatum

Thyasira flexuosa Microprotopus maculatusEuclymene oerstedii Ampelisca tenuicornis Ampharete lindstroemiAbra alba Mysella bidentata Echinocardium cordatumPygospio elegans Polydora ciliata

Aphelochaeta marioni

Spiophanes bombyx Owenia fusiformis Scalibregma inflatum

Pholoe

Description Species 7 Description Species 8

Abra alba Phoronis Amphiura brachiata

Species Trait Included in ProjectGrowth form YESMobility/movement YESTypical food types YESBioturbator? YESFragility NOHeight NOAdult dispersal potential NOSociability NOToxic/poisonous NOFeeding method YESEnvironmental position YESHabitat YESFlexibility NOSize YESGrowth rate NODependency NODistribution YESBiogeographic range NOMigratory NODepth range YESSubstratum preference YESBiological zone NOTidal stream preferences YESPhysiographic preferences YESWave exposure preferences NOSalinity preference YESReproduction type NOReproductive season NOReproductive frequency NOLifespan YESGeneration time NOEgg size NODevelopmental mechanism NOReproductive location NORegeneration potential NOAge at reproductive maturity NOFecundity NOFertilisation Type NOLarval/juvenile dispersal NODuration of larval stage NOLarval settlement period NOSensitivity to change YESSensitivity to anoxic conditions YESKey prey taxa YESTemporal variable population YESConnectivity to other habitats/species YESRelationships to other taxa YES

RationaleFundamental trait will inform modelFundamental trait will inform modelFundamental trait will inform modelNeeds consideration as an ecosystem functionNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelFundamental trait will inform modelPosition may influence links within modelHabitat preference will inform modelNot relevant to ecological modelFundamental trait which may inform ecosystem functions and be influenced by driversNot relevant to ecological modelNot relevant to ecological modelDistribution may influence spatial variation within model.Covered by other traitsNot relevant to ecological modelDepth constraints may inform modelFundamental trait will inform modelWill be either circalittoral or infralittoralWill inform model driversWill inform model driversEnergy preference will be covered by tidal stream preferenceWill inform model driversNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelLongevity may inform aspects of modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelNot relevant to ecological modelFundamental trait will inform modelFundamental trait will inform modelWill inform ecosystem functionWill inform temporal aspects of modelFundamental trait will inform modelFundamental trait will inform model

SourceMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMESLMESLMESLMESLMESLMESL

Species (Biotope name) Species (WORMS name) Species Confusion Growth form

Abra alba Bivalved

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida Bivalved

Abra nitida

Abra nitida

Abra nitida

Ampelisca tenuicornis Articulate

Ampelisca tenuicornis







Ampharete lindstroemiAmpharete lindstroemi

Ampharete lindstroemi



Ampharete lindstroemi Vermiform segmentedAmpharete lindstroemiAmpharete lindstroemiAmpharete lindstroemi


Amphiura filiformis Radial / Stellate

Amphiura filiformis


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis




Tharynx marioni superseded combination Recent name changed from Tharynx marioni. Vermiform segmented / Cylindrical

Aphelochaeta marioni is very difficult to identify (Mike Kendall, pers. comm.) and some authors (e.g. Farke, 1979) have commented that specimens that have been the subject of published research may have been misidentified.


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina

Vermiform segmented / Vemiform annulated / Cylindrical

Arenicola defodiens sp. nov. has recently been distinguished from Arenicola marina


Arenicola marina

Brissopsis lyrifera Globose

Brissopsis lyrifera

Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyriferaBrissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera

Callianassa subterranea Articulate

Callianassa subterranea


Calocaris macandreaeCalocaris macandreae

Cancer (Astacus) subterraneus Montagu, 1808 (genus change)


Calocaris macandreae


Calocaris macandreae Articulate



Capitella capitata

Capitella capitata

Capitella capitata

Cylindrical / Vermiform segmented


Capitella capitata

Carcinus maenas Articulate

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule Bivalved

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cirriformia tentaculataCirriformia tentaculataCirriformia tentaculata


Cirriformia tentaculataCirriformia tentaculata

Cirriformia tentaculata




Echinocardium cordatum Globose

Echinocardium cordatum

Vermiform segmented, Cylindrical









Echinus esculentus Globose

Echinus esculentus


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii

Euclymene oerstediiEuclymene oerstediiEuclymene oerstedii Vermiform segmented

Galathowenia oculata

Euclymene oerstedi (Claparède, 1863)

Myriochele oculata Zachs, 1923 (superseded original combination)




Galathowenia oculataGalathowenia oculata

Galathowenia oculata Vermiform segmented

Goniada maculate Goniada maculataGoniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate Vermiform segmented

Hediste diversicolor Vermiform segmented

Hediste diversicolor



Labidoplax media

Labidoplax media

Labidoplax mediaLabidoplax media

Lagis koreni

Lagis koreni

Lagis koreniLagis koreniLagis koreni

Lagis koreni

Lagis koreni

Synonymised name Oestergrenia media (Östergren, 1905)

Pectinaria koreni (supersed recombination)


Lagis koreni

Lagis koreni Vermiform segmentedLeptosynapta bergensis

Leptosynapta bergensis


Leptosynapta bergensisLeptosynapta bergensis

Macoma balthica Bivalved

Macoma balthicaMacoma balthica

Macoma balthica

Macoma balthicaMagelona johnstoni Vermiform segmentedMagelona johnstoni


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Malacoceros fuliginosus Vermiform segmented

Malacoceros fuliginosus






Maxmuelleria lankesteri

Spio fuliginosus Claparède, 1870 (superseded original combination)






Mediomastus fragilis Vermiform segmented

Mediomastus fragilis





Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Melinna palmata Vermiform segmented

Microprotopus maculatusMicroprotopus maculatus

Microprotopus maculatus Articulate

Microprotopus maculatus



Mya truncata BivalvedMya truncataMya truncataMya truncata

Mya truncata

Mysella bidentata Kurtiella bidentata (accepted name)

Mysella bidentata

Mysella bidentata

Mysella bidentata Bivalved


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus Articulate

Nephrops norvegicus


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii Vermiform segmented

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Nephtys hombergii

Nuculoma tenuisNuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis Bivalved

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci

Ennucula tenuis (Montagu, 1808)

Ocnus brunneus may be a smaller form of Ocnus planci. O. planci has also been confused with Aslia lefevrei.


Ocnus planci

Ocnus planci

Owenia fusiformis Vermiform segmented

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus Articulate


Pagurus bernhardus

Phaxas pellucidus Bivalved

Phaxas pellucidusPhaxas pellucidus

Phaxas pellucidus

Philine aperta Globose

Philine apertaPhiline aperta

Vermiform segmented

Pholoe inornata (sensu petersen)Pholoe inornata (sensu petersen)





Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata Articulate

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata

It has been suggested that some other species of Polydora such as P. ligni, P. websteri, P. cirrosa and P. nuchalis may only be varieties of Polydora ciliata

Vermiform segmented / Tubicolous


Polydora ciliata

Polydora ciliataPolydora ciliataPolydora ciliata

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior

Rhodine graciliorRhodine gracilior

Rhodine gracilior

Tubicolous / Vermiform segmented

Rhodine loveni gracilior Tauber, 1879 (superseded original combination)


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior Vermiform segmented

Sagartiogeton undatus Cylindrical / Radial

Sagartiogeton undatus



Scalibregma inflatum




Scoloplos armigerScoloplos armigerScoloplos armiger

Scoloplos armiger Vermiform segmented


Scoloplos armiger

Scoloplos armigerScoloplos armiger

Scoloplos armiger

Spiophanes bombyx Vermiform Segmented

Spiophanes bombyxSpiophanes bombyx

Spiophanes bombyx Tubicolous

Spiophanes bombyx

Thysanocardia procera


Golfingia (Thysanocardia) procera (Möbius, 1875) (subsequent combination)




Thysanocardia proceraTubificoides (pseudogaster)Tubificoides (pseudogaster)Tubificoides (pseudogaster)Tubificoides (pseudogaster)Tubificoides (pseudogaster)

Tubificoides (pseudogaster)

Tubificoides (pseudogaster)Tubificoides (pseudogaster)

Virgularia mirabilis Pinnate

Virgularia mirabilis


Species (Biotope name)

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Mobility/movement Feeding method Typical food types Bioturbator?

Burrower Phytoplankton / detritus

Surficial modifiers

Surficial modifiers

Burrower

Phytoplankton / detritus

Surficial modifiers

Passive suspension feeder / Active suspension feeder / Surface deposit feeder / Sub-surface deposit feeder


Swimmer / Burrower / Temporary attachment


Feeds on detritus, larvae and small organisms from the sediment surface or overlying water.











Ampharete lindstroemiAmpharete lindstroemiAmpharete lindstroemiAmpharete lindstroemi


Amphiura filiformis

Amphiura filiformis


Burrower

Crawler / Burrower Plankton and detritus

Biodiffusors

Upward and downward conveyors

Deposit feeder: surface / Interface feeder / Suspension feeder: facultative

detritus, organic matter in the sediments and plankton from the water column



Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





Surficial modifiers

Burrower Surface deposit feeder Organic debris / diatoms


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Burrower Sub-surface deposit feeder

Micro-organisms (bacteria), benthic diatioms, meiofauna and detritus

Reworking of the surface sediment (bioturbation) by Arenicola marina increase the penetration of oxygen into the upper 2 -10cm of sediment together with rapid mixing of sediment particles.


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Burrower

Biodiffusors

Surface deposit feeder / Sub-surface deposit feeder

Organic detritus / foraminifers and other small organisms within the sediment


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






Crawler / Burrower

Biodiffusors

Biodiffusors

Surface desposit feeder / Sub-surface deposit feeder

Organic content of sediment particles







Capitella capitata

Capitella capitata

Capitella capitata


Principally a deposit feeder

Crawler / Burrower

Burrower

Upward Conveyors

mixture of organic and fine inorganic fragments. Diet components from gut analysis consisted of diatoms, dinoflagellates, algae and terrestrial plant fragments and material of animal origin.


Micro-organisms, phytoplankton and detritus


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



Crawler Omnivore

Regenerators

Crawler / Burrower

Surficial modifiers

Detritus, diatomsSurficial modifiers

Limited mobility / Burrower

plants, algae, molluscs, arthropods (including their own species), annelids and carrion. The diet of large C. maenas mainly consits of molluscs and Mytilus edulis. Smaller C. maenas (<30 mm) have more plant matter and arthropods in their diet.

Passive suspension feeder / Active suspension feeder

Phytoplankton, zooplankton and organic particulate matter










Burrower

Burrower Detritus

Biodiffusors

Deposit feeder / Interface feeder / Suspension feeder: facultative










Echinus esculentus

Echinus esculentus


Crawler Not relevantGrazer (grains/particles/fronds/substratum)

Recorded feeding on worms, barnacles, hydroids, tunicates, bryozoans, macroalagae, bottom material and detritus.


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii

Euclymene oerstediiEuclymene oerstediiEuclymene oerstedii



Sub-surface deposit feeder

Crawler / Burrower

Surficial modifiers

Grazer (grains/particles/fronds/substratum)

Upward and Downward conveyors

Detritus, protists and bacteria






Goniada maculateGoniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




Burrowersubsurface deposit feeder

BiodiffusorsSwimmer / Burrower Predator / Scavenger

Burrower / Swimmer / Crawler

Biodiffusors

Detritus and organic matter in the surrounding sediment

sedentary polychaetes below the sediment surface

Other burrowing polychaetes

Surface deposit feeder / Omnivore / Scavanger / Sub-surface deposit feeder / Passive suspension feeder

Mud, sand & detritus. Phytoplankton & plankton. Other macrofauna



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


Crawler / Burrower

Upward conveyorsBurrower

active suspension feeder / sub-surface deposit feeder

suspended plankton and debris


Deposit feeder, forages with tentacles at the sediment-water interface

Mineral grains and detrital particles. Small copepods and polychaetes


Lagis koreni

Lagis koreniLeptosynapta bergensis




Macoma balthica


Macoma balthica

Macoma balthicaMagelona johnstoniMagelona johnstoni


Surficial modifiers

Crawler / Burrower

detrital material

Crawler / Burrower

Surficial modifiers

Burrower DetritusSurficial modifiers

active suspension feeder / sub-surface deposit feeder



Diatoms, deposited plankton, suspended phytoplankton and detritus

Macoma balthica is classed as a biodestabiliser. Widdows et al. (2000) found a significant relationship between sediment erodability (mass of sediment eroded and erosion rate) and the density of Macoma balthica.


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Selective deposit feeders

Burrower / Swimmer

exploit larger particulate matter including small invertebrates on the surface of the deposits

Upward and Downward Conveyors

Deposit feeder: surface / Interface feeder / Suspension feeder













Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Burrower

Burrower Sub-surface deposit feeder

Upward conveyors

Burrower



Detritus and organic matter in the surrounding sediment

deposit feeder: surface / Interface feeder / Suspension feeder: facultative


Melinna palmata






Mya truncataMya truncataMya truncataMya truncata

Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Surficial modifiers

Omnivore / Predator / Scavenger

Burrowing

Surficial modifiers

Suspension feeder

Burrower Active suspension feeder

Crawler / Burrower

Surficial modifiers


Phytoplankton, small zooplankton, benthic diatoms, suspended particulates and dissolved organic matter


Plankton and organic matter in the surrounding sediment


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


Swimmer / Crawler / Burrower Predator / Scavenger

Biodiffusors

Nephrops is an opportunistic predator feeding on crustaceans, molluscs and to a lesser extent polychaetes and echinoderms.


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Swimmer / Crawler / Burrower Predator / Scavenger

Biodiffusors

Preys upon the burrowing shrimp Calocaris macandreae

Molluscs, crustaceans and other polychaetes


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Surficial modifiers

Burrower

Crawler

Suspension feeder

Scoloplos armiger and Heteromastus filiformis are major prey of N. hombergii

Deposit feeder: subsurface / Deposit feeder: surface

Organic matter in the surrounding mud or sand



Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Burrower

Surficial modifiers

Epifauna

Crawler Organic detritus, carrion Not relevant


Phytoplankton and particulate organic matter

Surface Deposit Feeder / Predator / Active Suspension Feeder


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta

Philine apertaPhiline apertaPholoe inornata (sensu petersen)Pholoe inornata (sensu petersen)





Burrower

Surficial modifiersPlankton

Crawler Predator / Scavenger

Surficial modifiers

Surficial modifiers

Crawler Predator Small invertebrates


Pectinaria koreni, Echinocyamus pusillus, foraminiferans, and small infaunal lamellibranchs and gastropods.


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


Surficial modifiers

Burrower

Swimmer / Burrower

Surficial modifiers

Burrower


Suspended particles in the water


Suspended particles (planktonic, organic) in the water


Detritus, suspended particles and occasionally dead barnacles and other dead invertebrates



Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Burrower

Upward conveyorsDeposit feeder: subsurface





Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Burrower

Temporary attachment / Burrower Passive suspension feeder

Burrower

Biodiffusors

Detritus

Biodiffusors

Burrower Diffusive mixing

Ingest detritus or organisms such as protists and bacteria from the sediments

Carrion, small invertebrates

Surface desposit feeder / Sub-surface deposit feeder

Surface Deposit Feeder / Sub-surface Deposit Feeder


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Burrower

Biodiffusors

Burrower Detritus

Detritus and organic matter in the surrounding sediment.

Passive Suspension Feeder / Active Suspension Feeder / Surface Deposit Feeder / Sub-surface Deposit Feeder

Sediment particles, planktonic organisms, meiobenthic organisms (Dauer et al., 1981).

Upward and downwards conveyors











DetritusBiodiffusors

Deposit feeder

Temporary attachment Passive suspension feeder

Surficial modifiers


Plankton and organic particles


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Environmental positioHabitat Size

infaunal Burrow dwelling Small (1-2 cm)

Small (1-2cm)

infaunal Burrow dwelling

Epifaunal / Epilithic Free living Small-medium (3-10 cm)













Amphiura filiformis

Amphiura filiformis


up to 12 mm

Infaunal Tubiculous

Infaunal Free living Medium (11-20 cm)

Tubiculous (Tube dwelling)

Lives in fragile tubes


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





infaunal Burrow dwelling Small-medium (3-10 cm)


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Infaunal Burrow dwelling Medium (11-20 cm)


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Infaunal Free living Small-medium (3-10 cm)


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






infaunal Burrow dwelling Small-medium (3-10cm)

Small-medium (3-10 cm)







Capitella capitata

Capitella capitata

Capitella capitata


Infaunal Burrow dwelling Largest specimen 50 mm total length

Infaunal Burrow dwelling Small-medium (3-10 cm)


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



Epibenthic Free living Small-medium (3-10 cm)


Free living Medium-large (21-50 cm)










Infaunal










Echinus esculentus

Echinus esculentus


Epifaunal Free Living Medium (11-20cm)


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




Infaunal Tubiculous Medium (11-20 cm)







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




Small - medium (3-10 cm)

Infaunal

Tubiculous


Infaunal

Burrow dwelling




Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


Infaunal up to 50 mm long

Burrow dwelling

Infaunal Tubiculous50 mm


Lagis koreni





Macoma balthica


Macoma balthica



Infaunal up to 300 mm long (easily broken)

20-50 cm

Burrow dwelling

Infaunal Burrow dwelling Small (1-2 cm)



Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Infaunal Small-medium (3-10 cm)

Free living / Burrowing












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Infaunal 120 x 300 mm

Burrow dwelling

Epibenthic Tubicolous Small-medium(3-10cm)

Small-Medium (3-10cm)

Infaunal Tubicolous 10-60 mm


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Very small (<1 cm)

Infaunal


Infaunal Burrow dwelling

Infaunal Burrow dwelling Small (1-2 cm)

Burrow dwelling / Tube builders


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


Demersal / Infauanl Free living Medium-large (21-50 cm)


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Infaunal Free living Medium (11-20 cm)

Constructs and inhabits extensive burrow complexes


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Very small (<1 cm)


Epifaunal

150 mm


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Free living

Infaunal Tubiculous Small-medium (3-10 cm)

Epifauna

Free living Small-medium(3-10cm)EpifaunalEpibenthic


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta








Infaunal / Epifaunal Free living Small (1-2 cm)


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata



Tubiculous

Infaunal

Very small (<1 cm)

Epilithic Tubiculous

Tubiculous Small (1-2 cm)

Tubiculous but free living within their tube

Epibenthic / Epilithic / Epizoic


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Infaunal Small (1-2 cm)

Tubiculous



Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Infaunal Tubiculous

Epibenthic / Epifaunal Attached Medium (11-20 cm)



Infaunal Burrow dwelling Medium(11-20 cm)


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Infaunal Burrow dwelling Small-medium(3-10cm)

Free living Small-medium (3-10 cm)











Infaunal

Free living Small-medium (3-10 cm)

Infaunal

Epifaunal / Infaunal Attached Large (>50 cm)


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Distribution Depth range

Widespread around the British Isles 0 - 60 m

Widespread on the coasts of Britain but less common on the western coast of Ireland.

In the sublittoral zone up to 183 mIntertidal / Continental Shelf (to 200m)

Found on most British and Ireland coasts from the Shetlands to the Scilly Isles.













Amphiura filiformis

Amphiura filiformis


0-500 m

8 - 400m

Along most British and Northern Ireland coasts

5 - 1200 mMost British and Irish coasts although records have not been found for the south of England


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





Mid shore to 5000m

Patchily distributed all around the British coast where suitable substrata exist. Occurs on the south west and south coasts of the Isle of Man and has also been recorded in north east Ireland.


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Found on all coasts around Britain and Ireland. Intertidal


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


5 - 100 mRecorded off the west, north and east coasts of the British Isles, but not off the south coast. Common in deep water.


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






0-20 cm

35-1400 m

Recorded distribution is limited to the south coast of Britain, west coast of Scotland and a single site in the Kenmare River area, southern Ireland. Callianassa subterranea is likely to be more widespread than records suggest.







Capitella capitata

Capitella capitata

Capitella capitata


Reported from all coasts of Britain and Ireland.

1-650 m

West Scotland, Isle of Man, Irish Sea, north east England, southern North Sea


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



Found on all shores of Britain and Ireland. intertidal down to 60 m

Intertidal

Widely distributed in estuaries and sandy bays around the coast of Britain and Ireland.










Lower shoreBritish channel, North Sea

South coast, Devon mid-tidal zone

South coast, Southampton

0 - 230 mfound on sheltered sandy beaches, on all coasts of Britain and Ireland.









Echinus esculentus

Echinus esculentus


0- 1200m

All around the coast of the UK

Most of the coasts of the British Isles, though it is absent form East coast of England, the eastern English Channel and some parts of north Wales.

0 - 40m (sometimes deeper)


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




Common on the west coast of Scotland.

0 - 160 m

British channel, Northwest and Southwest Northsea

From the upper limits of the Laminaria zone to depths .100m.]







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




Northsea

Abyssal regions

British channel, Northsea

Widespread along all British coasts Intertidal

upper sublittoral - 2800 m

Intertidal, bathyal, infralittoral and circalittoral



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


10 m

Shallow conditions

inter- and subtidal

Strangford Lough, Northern Ireland, South Uist in the Hebrides (Scotland)Recorded from the Quoile estuary in Strangford Lough, Northern Ireland and from similar sheltered habitats in the sea lochs of western Scotland.

Recorded in Shetland, Orkney, the east coast from St. Andrews Bay south to Bridlington, the Wash, south east, south and south west coasts of England, the Scilly Isles, Wales, east, south and north west Ireland, and the west coast of Scotland. Widely distributed in the North Sea and English Channel

Highest densities in shallow inshore regions


Lagis koreni





Macoma balthica


Macoma balthica



All round the British Isles

All Great Britain

low on shore to more than 100 m

Common in estuarine environments around the British Isles, with the exception of the south coast.

From mid shore to 190m. In British water M. balthica is mainly an intertidal species


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










All of Great Britain

intertidal (mid to lower shore) to 88m












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Irish Sea, west Scotland

North Sea & English Channel

Around the British Isles and Ireland

Lower shore to shallow sublittoral.


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


0 - 70 m

All British coasts

0 - 70 m

Common around the coast of Britain

Widespread around the Biritsh Isles 0 - 100 m

Wash, Portland, Channel Isles, West Channel, Scilly Isles, Isle of Man, Belfast, Clyde & Argyll, Mich, Mayo

Littoral and shallow sublittoral


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


200 - 800 m

Common around most British coasts but not apparently recorded for the English Channel, the Bristol Channel or the Western Approaches. Populations also exist to the north east of Scotland on the Fladen Ground.


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Found throughout Britain

intertidal and shallow sublittoral


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


20 - 150 m

2 - 175 m

Off the coasts of Scotland and in parts of the North Sea, Dublin, Isle of Man

Recorded from Wider Firth, Orkney; off Runswick Bay, east England; Dogger Bank; Tremadoc Bay, north Wales; southern Ireland; Carlingford and Strangford Lough, Northern Ireland, and sea lochs and inlets of western Scotland.


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Widespread around the British coast

North Sea

Found on all British and Irish coast

Low water and shallow sublittoralIntertidally to 470m. Locally abundant to depths of 27m

From MTL to 140 m, seldom down to 500 m


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






common off all British coasts 5 - 100 m

At a few recorded locations all around the British Isles. 0 - 500 m


Intertidal under stones, shallow sublittoral

Intertidal down to several thousand metres of depths


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


7 - 140 m

Found all around the British coast

0 - 200 m

Widely distributed around Britain

Shetland, Dogger, Portland, Channel Isles, West Channel, Isle of Man, Dublin, Donegal Bay, Mayo, Galway Bay


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Mid-tidal level

All of Great Britain

Northern Northsea

Intertidal and infralittoral

Upper sublitoral to 1000 m


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Firth of Forth and Northumberland coast

Present around the British coasts Lower shore to > 100m

Widespread around the UK

Widely distributed in NW Europe and Britain on

low water mark, shallow sublittoral. Up to 1000m depth

Lower shore and in sublittoral (Fish & Fish, 1996),


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Spiophanes bombyx is found off most British coasts

Low water mark to 60m. Could possibly occur down to 1km.











2 - 200 m

Intertidal to subtidal


Found off all British coasts but less frequent in the south 10-400 m

northern North Sea to the west coasts of Sctoland and Ireland including the Irish and Celtic Seas


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Substratum preference Tidal stream preferences Physiographic preference

Muddy gravel / Sandy mud / Mud / Muddy sand weak (<1 kn)

Mud / Muddy sand / Muddy gravel

Muddy sand

Open coast / Offshore seabed / Strait / Sound / Sealoch / Enclosed coast / Embayment













Amphiura filiformis

Amphiura filiformis


Fine sand

mud mixed with clay, sand or pebbles

fine sand or mud

Offshore seabed

Muddy sand / Sandy mudModerately strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Offshore seabed / Sealoch / Enclosed coast / Embayment


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





Fine clean sand / Mud / Muddy sand / Sandy mudModerately strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Open coast / Offshore seabed / Strait / Sound / Estuary / Enclosed coast / Embayment

Sometimes present in sandy sediments, but predominates in zones of muddy sand or silt


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Salt marsh / Seagrass / Mixed / Muddy gravel / Muddy sand / Sandy mud / Fine clean mud

Very strong (>6 kn) / Strong (3-6 kn) / Moderately strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Strait / sound / Sealoch / Ria / Voe / Estuary / Enclosed coast / Embayment / Isolate saline water (Lagoon)


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Mud / Muddy sand Weak (<1 kn) / Very Weak (negligible)

Offshore seabed / Open coast / Sealoch


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






Muddy sand / Sandy mudModerately strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Open coast / Offshore seabed / Strait / sound / Sealoch / Enclosed coast / Embayment







Capitella capitata

Capitella capitata

Capitella capitata


Mud / Sandy mud

Fine clean sand / Muddy sand / Mud / Sandy mudOpen coast / Strait / sound / Enclosed coast / Embayment / Estuary


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



No preference

organic-rich mud, under stones and amongst sea-grasses

Moderately Strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Open coast / Strait / sound / Sealoch / Ria / Voe / Estuary / Isolated saline water (Lagoon) / Enclosed coast / Embayment

Sandy mud / Muddy sand / Coarse clean sand / Fine clean sand / Seagrass / Muddy gravel

Moderately strong (1-3 kn) / Weak (<1 kn) / Very weak (negligible)

Enclosed coast / Embayment / Open coast / Strait / sound / Sealoch / Ria / Voe / Estuary / Sheltered










muddy sand / Gravel / often between or beneath large pebbles

Coarse clean sand / Fine clean sand / Muddy sand / Sandy mud

Open coast / Offshore seabed / Strait / sound / Enclosed coast / Embayment









Echinus esculentus

Echinus esculentus


Moderately Strong (1-3 kn)

Rocks, stones or seaweed.

Bedrock, Large to very large bouldersSmall boulders, Artificial (e.g. metal/wood/concrete), Rockpools, Under boulders, Caves, Crevices / fissures, Overhangs

Open coast / Strait / sound / Sealoch / Ria / Voe / Enclosed coast / Embayment


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




Algae / Mud

muddy fine sand







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




fine and coarse sand, mixed soil with clay, silt

Silt, mixed soil with coarse sands

muddy and sandy bottoms

Sandy mud / Muddy sand / Mud Weak (<1 kn) / Very weak (negligible)

Estuary / Enclosed coast / Embayment / Ria / Voe



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


Fine mud

Muddy sands or sandy muds

sandy to muddy sediments

Weak, extremely sheltered conditions

Sheltered basins of lagoons and sea lochs


Lagis koreni





Macoma balthica


Macoma balthica



muddy sand

Mud / Muddy sand / Muddy gravel

Mud / Muddy sand / Sandy mud

Coarse clean sand / Fline clean sand

Moderately strong (1-3 kn) / Weak (<1 kn)

Ria / Voe / Estuary / Enclosed coast / Embayment


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Muddy sand / Mud / Under boulders

Moderately strong (1-3 kn) / Strong (3-6 kn)

Open coast / Strait / sound / Enclosed coast / Embayment












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Mud or sand, rock-crevices or under stones

Muddy gravel, Sandy mud, Muddy sand, Mud

Mud / Muddy sand / Mixed bottom deposits / Zostera beds More abundant when mud content exceeds 20%

Enclosed coast / Embayment


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Among algae and on sand

Fine sand with mud

mixed sand / sandy mud / gravel

Muddy sand

Moderately Strong (1--3 kn) / Weak (<1 kn)

Strait / sound / Sealoch / Ria / Voe / Estuary / Enclosed coast / Embayment /


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


Mud / Muddy sand / Sandy mud Weak (<1 kn) / Very weak (nOffshore seabed


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Coarse clean sand / Fine clean sand / Muddy sand / Sandy mudOpen coast / Estuary / Enclosed coast / Embayment


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Sandy mud

Sandy mud / mud / muddy gravel

Open coast / Offshore seab

Soft muddy sediments

Moderately strong (1-3 kn) / Weak (<1 kn)

moderate to weak water flow


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Fine clean sand / Muddy sand / Sandy mud Weak (< 1 kn) Open coast / Offshore seabed / Estuary

Bedrock, Large to very large boulders, Rockpools, Under boulders, Coarse clean sand, Fine clean sand, Gravelly sand

Open coast / Offshore seabed / Strait / sound / Sealoch / Estuary / Enclosed coast / Embayment


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






Fine sand / Muddy sand / Mud and gravel

Muddy sand / Fine clean sand / Sandy mud

Amongst old shells

Soft and hard substrata


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


Mud / Sand

muddy to sandy substrates often with a high organic content and with and abundance of detritus and suspended material

Other species / Algae / Artifical (e.g. metal/wood/concrete) / Mud / Bedrock

Strong (3-6 kn) / Moderately strong (1-3 kn) / Weak (<1 kn)

Open coast / Offshore seabed / Strait / Sound / Estuary / Isolated saline water (Lagoon) / Enclosed coast / Embayment


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Mud / Sandy mud / Crevices / fissures

Mixed sediments of silt, sand and shell/gravel fragments

Sandy shores and mud flats and in mud that has collected in rock crevices


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Sand / Mud

Bedrock / Large to very large boulders / Small boulders / Cobbles / Pebbles / Gravel / Shingle / Coarse clean sand / Fine clean sand / Crevices / fissures

Mud / sand / gravel / attached to stones or shells 10-15 cm below the surface

Coarse clean sandFine clean sandMuddy sand


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Fine clean sand, Sandy mud Moderately Strong (1-3 kn), Weak (,1 Kn)

Open coast / Strait / sound / Estuary / Enclosed coast / Embayment / Sealoch











Muddy sand

Mudflats, Estuarine mud

Coarse clean sand / Fine clean sand / Mud / Muddy sand / Sandy mud

Weak (<1 kn) / Very weak (negligible)

Offshore seabed / Sealoch / Enclosed coast / Embayment


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Salinity preference Lifespan

Full (30-40 psu) 1-2 years

2-3 years

1-2 years













Amphiura filiformis

Amphiura filiformis


3-10 years



Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





3-5 yearsLow (<18 psu) / Reduced (18-30 psu) / Variable (18-40 psu) / Full (30-40 psu)


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


6-10 yearsFull (30-40 psu) / Variable (18-40 psu) / Reduced (18-30 psu)


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Variable (18-40 psu) / Full (30-40 psu) 3-5 years


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera













Capitella capitata

Capitella capitata

Capitella capitata


> 3 years

Up to 10 years

Full (30-40 psu) / Variable (18-40 psu) 1-2 years


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



6-10 years

6-10 years

5-6 years

Low (< 18psu) / Variable (18-40 psu) / Full (30-40 psu) / Reduced (18-30 psu)

Reduced (18-30 psu) / Full (30-40 psu) / Variable (18-40 psu)










Reduced (18-30 psu) / Full (30-40 psu) 11-20 years









Echinus esculentus

Echinus esculentus




Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




3-5 years







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




1-2 years

1 yearLow (<18 psu) / Reduced (18-30 psu) / Variable (18-40 psu)



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


1 year

< 1 year


Lagis koreni





Macoma balthica


Macoma balthica



Little information is available (3-10 years)

6-10 years

3-5 years

Variable (18-40 psu) / Reduced (18-30 psu) / Low (< 18 psu)


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Full (30-40 psu) / Variable (18 - 40 psu)

1-2 years












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


1-2 years

2-3 years

It has been suggested that the species is long-lived with stable populations and low recruitment rates.


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


11-20 yearsFull (30-40 psu) / Low (<18 psu) / Reduced (18-30 psu) / Variable (18-40 psu)


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus




Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Variable (18-40 psu) / Reduced (18-30 psu) 3-5 years


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Full (30-40 psu) Varied information (5-20 years)


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Variable (18-40 psu) / Full (30-40 psu) 3-5 years



Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






3-5 years

3 - 5 years


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


1 year

1-2 yearsLow (<18 psu) / Full (30-40 psu) / Variable (18-40 psu)


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


1 year


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


3-5 years


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Variable (18-40 psu) / Full (30-40 psu)

Short (no length given)











1-2 years

Range into low salinity

Full (30-40 psu)Birkeland, (1974) found the life span of Ptilosarcus guerneyi (same suborder to V. mirabilis) to be up to 15 years, taking 5 or 6 years to reach sexual maturity.


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Sensitivity to change Tolerance to anoxic conditions

Moderate sensitivity to substratum loss. There is a high intolerance to substratum loss with a high recoverability. High intolerance to synthetic compound contamination, alterations in behaviour (no or impaired burrowing) have been recorded after exposure to marine sediments contaminated with pesticides.

Tolerant to changes in oxygenation

Resistant to moderate hypoxia













Amphiura filiformis

Amphiura filiformis


Sensitive to hypoxia

Sensitivity to changes in light, which would alter their migration pattern.

Tolerant of sewage discharge. Appears to be able to tolerate more polluted environments than other Amphipod species.


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





The species has a moderate sensitivty to substratum loss, to an increase in water flow rate and an increase in wave exposure. The species has a high intolerance to synthetic compound contamination (e.g. TBT), hydrocarbon contamination (oil pollution) and to a decrease in salinity.

Low intolerance to changes in oxygenation

Flow regimes and the resulting particle flux, may influence the relative importance of the feeding modes of Amphiura filiformis. Feeding activity is a function of flow velocity with few animals extending their feeding arms in still water.

Rapid changes in population density caused by high mortality and successive reconlonisation in areas of fluctuating hydrodynamic conditions.

Moderate sensitivity to substratum loss and increase in wave exposure. High intolerance to synthetic compound contamination (e.g. ivermectin and TBT).



Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Arenicola marina is capable of anaerobic metabolism in hypoxic conditions.

Moderate sensitivity to substratum loss and a high intolerance to synthetic compound contamination (e.g. xenobiotic ivermectin).



Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Moderate sensitivity to substratum loss. Intolerance to Synthetic compound contamination (e.g. TBT, detergents used to disperse oil oil from the Torrey Canyon oil spill caused mass mortalities of a similar species, Echinocardium cordatum (Smith, 1968))

High intolerance to changes in oxygenation (sensitive to hypoxia). At low oxygen levels, B. lyrifera leave their positions in the sediment and migrate to the sediment surface


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






Highly sensitive to an increase in wave exposure

Opportunistic scavenger, found to increase in abundance after decades of fisheries, relatively tolerant to disturbance

Moderate sensitivity to substratum loss. High intolerance to synthetic compound contamination (TBT, carbaryl pesticide), hydrocarbon contamination (oil pollution) and to an increase in salinity (C. subterranea is stenohaline)








Capitella capitata

Capitella capitata

Capitella capitata


High intolerance to changes in oxygenation. Absent in areas which are de-oxygenated and characterised by a distinctive nutrient enrichment

C. macandreae is capable of withstanding prolonged hypoxia and anoxic conditions by switching to anaerobic respiration

High intolerance of C. capitata to substratum loss. Low intolerance to changes in oxygenation


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



C. capitata is a highly opportunistic species, recolonising quickly and restoring populations to earlier sizes in a short period of time after defaunation.

Intermediate intolerance to synthetic compound contamination and high intolerance to hydrocarbon contamination (oil pollution).

C. maenas is tolerant to changes in oxygenation

Moderate sensitivity to substratum loss. Moderate sensitivity to extraction of other species: Cockles are probably of high intolerance to bait digging although smaller individuals may be more intolerant.

High intolerance to changes in oxygenation










Cirriformia can maintain oxygen uptake down to 10% oxygen saturation. Cirriformia is an oxyconformer, with falling oxygen tensions the rate of oxygen uptake is reduced.

Oil polluted mudflats contained large populations of C. tentaculata. Spawning, growth and mortalitity were unaffected. C. tentaculata was killed by low concentrations of oil-dispersants (Essolvene and BP 1002).









Echinus esculentus

Echinus esculentus


Moderate sensitivity to substratum loss and abrasian & physical disturbance (such as movement of trawling gear over the seabed). High intolerance to synthetic compound contamination (e.g. TBT), hydrocarbon (oil) contamination and changes in nutrient levels. E. cordatum is generally found in sediments with low organic content and the species appears to be intolerant of increases in nutrient concentration.

Hgih intolerance to changes in oxygenation. In the south-eastern North Sea a period of reduced oxygen resulted in the death of many individuals of E. cordatum (Niermann, 1997) and during periods of hypoxia the species migrates to the surface of the sediment (Diaz & Rosenberg, 1995)

Opportunistic scavenger, found to increase in abundance after decades of fisheries, relatively tolerant to disturbance


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




The urchin has moderate sensitivity to substratum loss as it is slow moving and unlikely to escape. Though this does not necessarily apply to algae loss, as the urchin would be able to find other food sources. Echinus appears to be intolerant to synthetic compound, heavy metal and hydrocarbon contamination.

Intermediate intolerance to changes in oxygenation. Under hypoxic conditions echinoderms become less mobile and stop feeding.







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate






Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


High sensitivity to physical impact

Moderate sensitivity to substratum loss and synthetic compound contamination.

Intermediate intolerance to changes in oxygenation. Hediste diversicolor is resistant to moderate hypoxia. After 10 days of hypoxia (10% oxygen saturation) with sulphide (172-187 µmM) only 35% of H. diversicolor had left the sediment compared to 100% of Nereis virens.


Lagis koreni





Macoma balthica


Macoma balthica


Sensitivity to change Tolerance to anoxic conditionsResistant to moderate hypoxia

Very limited dispersal potential and assessed as a low recovery potential

Moderate sensitivity to substratum loss. High intolerance to synthetic compound contamination (TBT), heavy metal contamination and hydrocarbon contamination (oil). Some mortality of Macoma balthica may occur due to harvesting of other species so an intolerance of intermediate is recorded with a high recoverability.

Low intolerance (relatively tolerant) to changes in oxygenation. M. balthica may react to hypoxia by reducing metabolic activity.


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Moderate sensitivity to substratum loss

Vulnerable to sediment mobilisation

Tolerant to changes in oxygenation. Niermann et al. (1990) reported that Magelona sp. remained abundant during the period of hypoxia

Shows resistance to severe hypoxia












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Tolerant to physical disturbance

likely to be vulnerable to dredging, but it may be capable of tolerating sand mobilised by the dredging process

Resistant to moderate hypoxia

Extending their populations in the English Channel possibly as a result of the increase in mud habitats in the subtidal zone due to human disturbances.



Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Moderate sensitivity to substratum loss and hydrocarbon contamination



Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


Moderate intolerance to changes in oxygenation


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Moderate sensitivity to substratum loss.

High intolerance to changes in oxygenation. In moderately hypoxic conditions 38-43% saturation (3.8-4.3 mg O2/l) N. norvegicus compensates by increasing production of haemocyanin (Baden et al., 1990). In severe hypoxia, <20% saturation (<2 mg/l) Nephrops became less active and raised their bodies on their legs.

High intolerance to hydrocarbon contamination. High intolerance to the extraction of other species: Commercially exploitable species such as Cerastoderma edule occur in the same habitat as Nephtys species.


Nephtys hombergii is sensitive to high sedimentation rates of fine clay materials


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Intolerant to physical disturbance

Moderate sensitivity to substratum loss and increase in wave exposure

Intermediate intolerance to Changes in oxygenation. A long term decline of populations of Nucula nitidosa in the German Bight was attributed to an increased frequency of hypoxic events


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Intolerant to substratum loss.

Moderate sensitivity to substratum loss, to an increase in emergence regime (emergence is relevant) and to displacement (Adult Owenia fusiformis probably cannot construct new tubes once removed and therefore are probably highly intolerant to displacement).

Tolerant to changes in oxygenation. O.fusiformis is very tolerant of anoxia and can tolerate anaerobic conditions for up to 21 days by becoming quiescent (Dales, 1958)

Favours areas of physical disturbance as they are scavengers and are known to migrate rapidly into areas of fishing disturbance where they can feed on animals damaged by trawls.


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






Intolerant to disturbance, drastic reduction in numbers after decades of fishing


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


Can survive extended exposure to anoxia for up to 6 weeks


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Moderate sensitivity to substratum loss.

Low intolerance to changes in oxygenation. P. ciliata is repeatedly found at localities with oxygen deficiency

P. ciliata thrive in polluted conditions


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Intolerant to physical disturbance


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Favours Substratum lossOpportunistic polychaete, recovers well from substratum loss.

Tolerance against hypoxia and sulphide is low (Kruse et al., 2004), and worms may ascend into the oxic layer during low tide. Favours sand high in nutrients and turbid, so constantly being replaced.

Moderate sensitivity to substratum change (due to wave exposure etc), to increase in water flow rate and increase in wave exposure. Moderate sensitivity to synthetic compound contamination.

Intermediate intolerance to changes in oxygenation.











Moderately sensitive to physical disturbance

Moderately sensitive to physical disturbance

Virgularia mirabilis live upright with their stalks thrust into a mucus-lined burrow into which the whole colony can withdraw when disturbed.

Moderate sensitivity to substratum loss, increase in water flow rate, increase in temperature, increase in wave exposure and abrasian & physical disturbance. High intolerance to an increae in salinity.

High intolerance to changes in oxygenation. Stratification of the water column and hypoxia in near-bottom water is especially likely to occur during warm temperatures in semi-enclosed water bodies such as sea lochs. Virgularia mirabilis is often found in sea lochs so may be able to tolerate some reduction in oxygenation.


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Key prey taxa? Temporal variable population

Prey for flatfish including flounder Platichthys flesus, common sole Solea solea and plaice Pleuronectes platessa.

Prey for starfish Asterias rubensStrong seasonal and year-to-year variations due to strong recruitment, sedimentological changes and cold winters

Wading birds (e.g. Oystercatcher, Grey Plover and Bar-tailed Godwit)













Amphiura filiformis

Amphiura filiformis


Exhibit diurnal migration.

Highly variable seasonal abundance.

Epibenthic grazers (e.g. peracarid crustaceans, polychaetes, post-larval fish and juvenile decapods). Demersal organisms e.g. predatory polychaetes, crustaceans, demersal fish

Fish (e.g. Whiting Merlangius merlangus)


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





Important in the diet of many fish and invertebrate predators including dab Limanda limanda, haddock Melanogrammus aeglefinus and Norwegian lobster Nephrops norvegicus. These predators do not generally consume the entire brittle star but crop only the arms, which are later regenerated.

Large differences in recruitment success due to varying climatic conditions (strong winters) and food supply.

Flatfish, crabs, shrimps and gobies. Reise (1977) reported that Aphelochaeta marioni (studied as Tharyx marioni) was food for Crangon crangon, Carcinus maenas, Pomatoschistus microps and young Pleuronectes platessa.


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Flatfish, wading birds, ragworm (Nereis virens and Hediste diversicolor). Tail can be niped off by flatfish and wading birds while Arenicola is depositing casts.

Sediment turnover by A. marina has been recorded as highest in spring and summer.


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera



Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






BirdsStarfish, gulls, lobsters, crabs and fish







Capitella capitata

Capitella capitata

Capitella capitata


Nephrops norvegicus is known to prey on C. macandreae

C. capitata provides an important food source for the shrimp Crangon crangon


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



lesser octopus (Eledone cirrhosa)

Carcinus maenas, oystercatchers (Haematopus ostralegus), shrimp (e.g. Crangon crangon), flatfish (e.g. flounder Platichthys flesus)










Rapid growth during spring and summer months









Echinus esculentus

Echinus esculentus


Birds

Starfish, gulls, lobsters, crabs and fish

In coastal areas (20m depth) fast growth is associated with high annual mortality. Recruitment success are most likely related to temperatures of preceding winters.

Juveniles preyed upon by flatfish species (e.g. Solenette and Scaldfish)


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




Starfish, gulls, lobsters, crabs and fish

Is a main prey for Anarhichas lupus (wolf fish)







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




Whilst feeding at the mud surface the worm is particularly prone to predation by wading birds (e.g. curlews and oyster catchers) and fish. The burrow is thus an important refuge in which to retreat.



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


Fish, starfish and crustaceans

Estuarine fish and wading birds. Hediste diversicolor constitutes the main prey of about 15 waders and is the dominant prey for the avocet Recurvirostra avosetta, grey plover Pluvialis squatarola, curlew sandpiper Calidris ferruginea, bar-tailed godwit Limosa lapponica and curlew Numenius arquata (see Goss-Custard et al., 1977 and other references given in Zwarts & Esselink, 1989). Several flatfish prey on the macrobenthos of intertidal mudflats. Sole, Solea solea; dab, Limanda limanda, flounder, Platichthys flesus and plaice, Pleuronectes platessa, all include Hediste diversicolor in their diet.

Important prey for the gastropod Philine aperta

Lagis koreni is a significant food-source for commercially important demersal fish, especially dab and plaice Adult densities may exceed 1000m², e.g. Eagle

(1975), but numbers characteristically fluctuate widely from year to year, owing to variations in recruitment success and mortality.


Lagis koreni





Macoma balthica


Macoma balthica



Fish, starfish and crustaceans

Birds, fish, crustaceans, polychaetes.

Wading birds (e.g. Dunlin and Curlew)

Prey for starfish Asterias rubens


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Flatfish (e.g. plaice)












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Maxmuelleria lankesteri has been recorded in the stomachs of Irish sea cod

Maxmuelleria lankesteri is active all year round but seem to show peaks of activity in December and April when the proportion of easily-degradable organic matter at the sediment surface is at its highest

Fish species e.g. Callionymus lyra


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Fin fish, sandworms, crabs, wading birds


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


Wading birds (e.g. Bar-tailed Godwit)

Nephrops norvegicus is preyed upon by numerous fish e.g. cod Gadus morhua, raj Raja Clavata, small spotted catshark (dogfish) Scyliorhinus canicula


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Birds


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Wading birds (e.g. Oystercatcher)

Plaice (Pleuronectes platessa) and dab (Limanda limanda)


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


Fish e.g. Diplodus sargus (white seabream), starfish and crustaceans

Important food source for the mollusc Acteon tornatilis


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






Fish e.g. haddock and some flatfish


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


Relatively unkown but probably include fish, gastropods and nematodes.


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Fish (e.g. Goby Pomatoschistus microps)


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Nudibranch molluscs, Sea spider Pycnogonum littorale. Possibly some fish species


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




Fish (e.g. Plaice Pleuronectes platessa)Predatory polychaetes e.g. Nephtys hombergii











Fish, birds and invertebrates

Fish (e.g. Goby Pomatoschistus microps) and Brown shrimp (e.g. Crangon crangon)

Possibly important food source for the nudibranch Armina loveni as it is usually in company with V. mirabilis




Haddock, starfish Crossaster papposusV. mirabilis populations are known to completely withdraw into the sediment, little is known of the periodicity of this behaviour.


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Connectivity to other habitats/species Relationships to other taxa

Indicator species of the Abra alba community in the Southern North Sea













Amphiura filiformis

Amphiura filiformis


Host of the parasite Sphaeronella longipes (Copepoda)


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





Important link between the benthic and pelagic environment as it is important in the diets of many fish and invertebrate predators.

The bivalve Mysella bidentata is a cohabitant of A. filiformis


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


Bioturbation by A. marina may inhibit or enhance meiofauna and micro-organisms, depending on species. A. marina can increase aerobic decomposition but decrease anaerobic decomposition, and affect the sediment chemistry and nutrient cycling between the sediment and overlying water (Riisgård & Banta, 1998).

Arenicola marina burrows are a unique microhabitat for a number of meiofaunal species, for example 26 different species of meiobenthic Platyhelminthes (flatworms) were reported from different regions of the burrow in the intertidal mudflats of Sylt, North Sea (Reise, 1987).

Bioturbation by A. marina destabilises the sediment.

Bioturbation by A. marina has strong negative effects on the abundance of the crustacean Corophium volutator, the sediment-reworking stimulates Corompium to emigrate. A. marina was found to have strongly negative effects on juvenile densities of various other worm and bivalve species.


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


Urothoe poseidonis lives in association with A. marina as it benefits from the steady food and oxygen supply and minor temperature fluctuations inside the lugworm burrows

Brissopsis lyrifera plays a role in the enhancement of regional species heterogeneity in an otherwise largely homogenous environment. When burrowing, Brissopsis lyrifera disturbs the sediment in a way that may result in lowered sediment stability (De Ridder & Lawrence, 1982). This disturbance combined with its respiratory activity alters the sediment chemistry, probably increasing oxygenation of the sediment at deeper levels. Consequently, the effects of B. lyrifera on the associated meiofauna will arise through both its non-selective feeding habit and its alteration of the physical and chemical environment of the sediment in which the meiofauna live.

The occurrence of the ascothoracidan parasite Ulophysema öresundense (Brattström) has been observed in the body cavity of Brissopsis lyrifera


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






Amphiura chiajei and Brissopsis lyrifera typically co-occur on some soft bottom areas of the North Sea, the Skagerrak and the Kattegat; they form the so called 'Brissopsis-chiajei association'. Each species affects the other through its feeding and burrowing activities. B. lyrifera can negatively affect the growth of body and gonads of A. chiajei, while A. chiajei seemingly has no effect on the growth of B. lyrifera.

Although population density of Callianassa subterreanea is often high, Rowden & Jones (1994) observed individual shrimps to be aggressive and intolerant of each other.

The bopyrid isopod Ione thoracica resides in the branchial chamber beneath the carapace of Callianassa subterranea.

The opening of the burrows of Callianassa subterranea provide temporary refuge for fish such as the black goby Gobius niger, Pomatoschistus minutus. Occasional errant polychaetes, particularly polynoid worms, inhabit the burrows







Capitella capitata

Capitella capitata

Capitella capitata


Host for nematode parasites


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



the rhizocephalan barnacle Sacculina carcini is the most well known parasite of C.maenas. C. maenas is the first host of the acanthocephalan helminth Profilicollis botulus which infects eider ducks (Somateria mollissima) by ingestion of infected crabs. Small Carcinus maenas (3-11 mm CW) can be attacked by the parasitoid platyhelminth Fecampia erythrocephala.

Host to brucephalid cercariae, Cercaria fulbrighti, the parasitic copepod Paranthessius rostatu and the rhabdocele Paravortex cardiiand Paravortex karlings

Nephtys species occur in the same habitat as C. edule










The bivalve Tellimya (=Montacuta) ferruginosa is a commensal of Echinocardium cordatum, and as many as 14 or more of this bivalve have been recorded with a single echinoderm. Adult specimens live freely in the burrow of Echinocardium cordatum, while the young are attached to the spines of the echinoderm by byssus threads (Fish & Fish, 1996). The amphipod crustacean Urothoe marina (Bate) is another common commensal (Hayward & Ryland, 1995).









Echinus esculentus

Echinus esculentus


several parasitic gregarine protozoans, such as Urospora neapolitana, have been observed in the body cavity of E. cordatum

Copepods Micropontius ovoides and Pseudanthessius sauvagei

Urothoe poseidonis lives in association with E. cordatum

Has species living amongst the spines E.g. polychaete worms (Adyte assimilis), isopods (Astacilla intermedia) and various copepod species.


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




Solitary urchin

As a grazer Echinus clears space enabling other species to colonise. In low numbers the grazing effect is beneficial, in high numbers this effect is destructive.

Common occurrence in kelp forests. Flora within these forests was highest were urchins were in low numbers. It can be assumed that diversity would increase when the predator is absent.

Copepods Asterocheres violaceus, A. echinicola, Cryptopontius brevifurcatus, Pseudoanthessius liber and P. sauvagei







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate






Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


Hediste diversicolor has been implicated as an infaunal species that plays a confounding role in the colonization and establishment of pioneering saltmarsh species. Laboratory experiments have reported that H. diversicolor reduces te succes of Zostera noltii (Hughes et al. 2000). H. diversicolor has also demostrated significant negative effects on the survival of Spartina anglica seeds transplanted to sediment cores (Emmerson 2000).

Copepod Synaptiphilus tridens

Lagis koreni often co-occurs with high densities of Abra alba

Pectinariids and semelid or tellinid clams are often found in association and a trophic interaction seems quite possible.


Lagis koreni





Macoma balthica


Macoma balthica



Possibly associated to the polychaete scaleworm Malmgreniella andreapolis

Macoma balthica is host to at least three gymnophallid trematodes; Lacunovermis macomae (Lebour), Gymnophallus gibberosus (Loos-Franc) and Parvatrema affinis which is known to cause sexual castration (Swennen & Ching, 1974).


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










Host of Ciliocincta julini (endoparasitic)

numerous small bivalves and polychaete worms colonized the walls of Maxmuelleria lankesteri burrows.

the shrimp Jaxea nocturna, which often lives in association with the echiuran worm Maxmuelleria lankesteri (Nickell et al., 1995), may benefit from the organic-rich mud pulled into its burrows by the worm.












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


Nephrops burrows at a site in Loch Sween showed evidence of interactions with other species, including Maxmuelleria lankesteri, Jaxea nocturna and Leseurigobius friesii.


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


M. bidentata is a coinhabitant of the filter-feeding burrowing ophiuroid Amphiura filiformis.

Has been found in the perforations of old oyster shells, in the burrows of the sipunculid Golfingia and in association with the ophiuroid Acrocnida brachiata.


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


M. bidentata lives in the oxidized layers around A. filiformis's burrow.

The rare British Fries' goby Lesueurigobius friesii shares the burrows of Nephrops norvegicus. The minute Cycliophoran Symbion pandora was found in the mouth parts of Nephrops collected in Denmark and the first of its kind to be described. The following species have been observed on specimens of N. norvegicus from the Irish Sea: Triticella koreni, Balanus crenatus, Electra pilosa, Eudendrium capillare, Sabella pavonina, Serpula vermicularis and a forminiferan probably Cyclogyra sp. The polychaete Histriobdella homari has been observed on the pleopods of two N. norvegicus from the Irish Sea and Clyde Sea


Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


Commercially exploitable species such as Cerastoderma edule occur in the same habitat as Nephtys species.


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


Associated with Thasira flexuosa and Abra alba

The Amphipod Tritaeta gibbosa inhabits O. planci by creating a pit and propeling itself into the mantle.


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus



Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






Epifaunal host.

Hermit crab shells are basibiont. Can be thought of as a symbiotic relationship between epibiota and host, though the relationship does vary depending on biological/environmental factors. Large number of epifaunal species can be found on the shell (E.G Hydroids, bryozoans, actinaria, serpulids, spirobids etc.


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


Frequently recorded in Macoma and Amphiura communities

Association with the Anthozoa Cerianthus lloydii.


Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


Associated with sepulid worm heads

In areas of mud tubes built by Polydora ciliata can agglomerate and form layers of mud up to an average of 20cm thick, occasionally to 50cm. These layers can eliminate the original fauna and flora, or at least can be considered as a threat to the ecological balance achieved by some biotopes (Daro & Polk, 1973).

Often associated with oysters and mussels. P. ciliata only invades the shell and not the soft tissue.


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


Megaclausia mirabilis is an associated copepod (Clausiidae family)

Sagartiogeton undatus is often found in the company of Sagartia troglodytes or Cereus pedunculatus

Host for Paranthessius anemoniae Claus, 1889 (parasitic: ectoparasitic)


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




In this article it S.armiger lived in the faecal mounds of A. marina, benefiting from the regular turbation caused by the latter.

Tube building worms, including Spiophanes bombyx, modify the sediment making it suitable for later colonisation and succession .











In the Kattegat populations, individuals appear to act as temporary parasites of the polychaete Aphrodite aculeata L.

Often found with T. benedii


Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra alba

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida

Abra nitida



Additional Notes Reference Code Confidence

R3 Medium

Limited movement R2 High

Brittle shell R4 High

R5 Medium

R6 Medium

R125 Medium

R3 Medium


R7 Medium

R87 High

R142 High

Expert Opinion #N/A

R3 Medium


Solitary animals but with adult densities exceeding 1000m2 in favourable conditions. Smallest recorded specimen 0.34mm













Amphiura filiformis

Amphiura filiformis


R8 High

R9 Medium

R10 Medium

R11 High

R48 High

R12 HighR13 Medium


R35 Medium

R62 High

R104 MediumR127 MediumR142 HighR167 Medium

Expert Opinion #N/A

R3 Medium

slow, free movement through the sediment matrix R2 High

Constructs a flattened flexible mud covered tube, extending 5-10mm above the sediment surface.


Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis

Amphiura filiformis





R14 High

R15 Medium

R16 Medium

R17 Medium

R18 High

R80 Medium


R3 Medium

R19 High

Amphiura filiformis is bioluminescent, arms emit blue light when individuals are mechanically stimulated.

majority of individuals live in the upper 4 cm of the sediment, with the smaller animals nearer the surface.


Arenicola marina

Arenicola marina

Arenicola marina

Arenicola marina


R3 Medium

R20 High

R21 High

R22 Medium

The burrow is irrigated (and therefore aerated) by intermittent cycles of peristaltic contractions of the body from the tail to the head end. Therefore, fresh water is taken in at the tail end and leaves by percolation through the feeding column. Arenicola marina ingests sediment at head end of the burrow forming a feeding column and characteristic funnel or 'blow hole' on the surface

Arenicola marina is presently used routinely as a standard bioassay organism for assessing the toxicity of marine sediments


Arenicola marina

Brissopsis lyrifera

Brissopsis lyrifera

Brissopsis lyrifera


R176 High

R3 Medium

Slow, free movement through the sediment matrix R2 High

R23 HighSea-urchins, especially the eggs and larvae, are used for toxicity testing and environmental monitoring.


Brissopsis lyrifera


Brissopsis lyrifera

Brissopsis lyrifera






R24 Medium

R49 HighR53 High

R121 High

R110 High

R3 Medium

free movement R2 High

R25 Medium

R3 Mediumfree movement R2 High

In the laboratory, Callianassa subterranea showed self-inhibiting burrow construction. Burrows were smaller when individuals were present in high densities

The species is iteroparous, possibly breeding twice a year, producing planktonic larvae and so recovery to substratum loss is expected to be rapid.







Capitella capitata

Capitella capitata

Capitella capitata


R26 High

R51 High

R52 High

R137 Medium

R173 High

R3 Medium


R27 High

C. macandreae is a protandrous hermaphrodite changing from male to female after three years


Capitella capitata

Carcinus maenas

Carcinus maenas

Carcinus maenas

Carcinus maenas

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule

Cerastoderma edule



R28 Medium

R3 Medium

Free movement R2 High

R29 Medium

R55 High

R3 Medium


R30 Medium

R39 Medium

R3 MediumLimited movement R2 High

R35 Medium

In the Wadden Sea and, probably colder, northern parts of Britain, Carcinus maenas migrates to subtidal areas and remains there until spring. During this time the crabs are inactive in shelters and do not feed. Lack of prey in the winter also leads to starvation and inactivity

Commercially fished in areas such as Morecambe Bay, the Wash, Thames Estuary, Dee Estuary, Outer Hebrides and South Wales.










R56 High R57 High

R58 High

R59 High

R61 High

Expert Opinion #N/A

R3 Medium


Does not irrigate its burrows, sediment around the worm is commonly blackened and smells strongly of hydrogen sulfide









Echinus esculentus

Echinus esculentus


R31 High

R49 High

R50 Medium

R53 High

R68 Medium

R121 High

R176 High

R3 Medium

Maximum diameter 16cm R105 High


Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Euclymene oerstedii

Euclymene oerstedii




R106 High

R107 High

R108 High

R109 High

R110 High

R53 High

IUCN Red list, Near threatened R126 High

organisms live in fixed tubes R2 High

R3 Medium

R57 High R60 MediumExpert Opinion #N/A

Organisms live in fixed tubes R2 High

Lower Risk (Red List Category). Largest diameter recorded 17.6cm across.

Echinus are grown in aquaculture due to demand for urchin roe in the Asian market

Described as a key functional species, due to its grazing traits







Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate

Goniada maculate




R3 Medium

R57 High

R63 High R64 High

Expert Opinion #N/A

Slow, free movement through the sediment matrix R2 HighR3 Medium

R65 High

R66 Medium

R57 High

R63 High

Expert Opinion #N/A

R3 Medium


Increase in abundance in moderately organically enriched areas



Labidoplax media

Labidoplax media


Lagis koreni

Lagis koreni


Lagis koreni

Lagis koreni


R32 High

R53 High

R54 Medium

R164 HighExpert Opinion #N/A

R3 Medium

R33 Medium

Live in fixed tubes R2 HighR63 High R56 High

R67 Medium

R123 Medium

Hediste diversicolor may be used as bait by anglers and are often sold commercially. They are harvested using a fork to turn over the substrata and collected. Hediste diversicolor is also used as a food source in aquaculture


Lagis koreni





Macoma balthica


Macoma balthica



R142 High

Expert Opinion #N/ASlow, free movement through the sediment matrix R2 High

R53 High

Hermafrodite R35 Medium

R164 HighExpert Opinion #N/A

R3 Medium

R5 MediumLimited movement R2 High

R34 Medium

R125 MediumR3 Medium


Macoma balthica is not normally considered to be toxic but may transfer toxicants through the food chain to predators. Macoma balthica was implicated in the Mersey bird kill in the late 1970's, owing to bioconcentration of alklyC-lead residues (Bull et al., 1983).


Magelona johnstoni

Magelona johnstoni

Magelona johnstoni

Magelona johnstoni










R35 Medium

R69 Medium

R70 High

R165 Medium

R3 Medium


R35 Medium

R71 High

R56 High

R142 High

Expert Opinion #N/A

R36 Medium












Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata

Melinna palmata


R56 High

R72 High

R73 Medium

Expert Opinion #N/A

Tubes are created from sand R3 Medium

R2 High

Short range mobility R35 Medium

R116 Medium

R142 High

Expert Opinion #N/A

R74 High

R63 High

R75 Medium

R128 Medium

R142 High


Melinna palmata







Mya truncata

Mysella bidentata

Mysella bidentata

Mysella bidentata

Mysella bidentata


Expert Opinion #N/A

R3 MediumSlow, free movement through the sediment matrix R2 High

R11 #N/A

R76 #N/A

R56 High

R77 High

Reproductive maturity at 3 - 5 years R3 MediumLimited movement R2 High

R37 MediumR78 High

R79 Medium

R3 Medium


R80 Medium

R81 High


Mysella bidentata

Mysella bidentata

Mysella bidentata

Nephrops norvegicus

Nephrops norvegicus


R82 Medium

R125 Medium

R142 High

R3 Medium



Nephrops norvegicus

Nephrops norvegicus

Nephrops norvegicus

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii

Nephtys hombergii


R38 High

R137 Medium

R188 High

R3 Medium


R39 Medium

R122 High

In British waters, the Nephrops fishery has grown rapidly since it began in the 1950s


Nephtys hombergii


Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis

Nuculoma tenuis


Ocnus planci

Ocnus planci

Ocnus planci

Ocnus planci


R159 High

R3 MediumSlow, free movement through the sediment matrix R2 High

R83 High

R84 High

R81 High

R103 Medium


R40 Medium

R53 High

R85 #N/A

R86 Medium


Ocnus planci

Ocnus planci

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Owenia fusiformis

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus

Pagurus bernhardus


R164 High

Expert Opinion #N/A

R3 Medium


R41 Medium

R56 High

R111 High

R112 High

R113 Medium

Free Movement R2 High

R3 Medium


Pagurus bernhardus

Phaxas pellucidus


Phaxas pellucidus

Philine aperta






R114 High

R3 Medium

Limited movement R2 HighR81 High

R121 High

R3 Medium

Slow, free movement through the sediment matrix R2 HighR88 High


R3 Medium

R56 High

R63 High

R167 Medium

Philine aperta lives just beneath the surface of fine sediment. The species 'ploughs' through the sediment as it moves and should not really be considered as burrowing species.


Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Phoronis muelleri

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Photis longicaudata

Polydora ciliata

Polydora ciliata


R3 Medium

R42 Medium


R97 High

R178 Medium

R3 Medium


R11 #N/A

R90 High

Expert Opinion #N/A

R3 Medium



Polydora ciliata


Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Pygospio elegans

Rhodine gracilior


Rhodine gracilior


R43 Medium

R91 HighR110 HighR113 Medium

R3 Medium


R44 Medium

R92 High

R124 High

Expert Opinion #N/A

R3 Medium

Organisms live in fixed tubes R2 HighR93 High

R57 High


Rhodine gracilior

Rhodine gracilior

Rhodine gracilior










Scoloplos armiger


R63 High

R94 High

Expert Opinion #N/A

R3 Medium

R45 Medium

R95 #N/A

Rarely present in high abundances R96 High

R3 Medium


R46 Medium

R63 High

R2 HighFavours areas of high nutrients R115 Medium

R116 Medium

R3 Medium

Never form tubes but construct galleries through the sediment down to depths of 60cm


Scoloplos armiger


Scoloplos armiger

Spiophanes bombyx


Spiophanes bombyx

Spiophanes bombyx




R117 Medium

R124 HighR159 High

Expert Opinion #N/A

R3 Medium

R118 HighR119 High

Sand tubes it creates are fixed R2 High

R120 High


R3 Medium

Move freely below the sediment without ever forming burrows. Found 10cm below the sediment surface.

It is often found at the early successional stages of variable, unstable habitats that it is quick to colonize following perturbation (Pearson & Rosenberg, 1978

During suspension feeding captured particles are accumulated in a ciliated groove before being transported to the pharynx, this is termed 'basal' food groove accumulation behaviour (Dauer et al., 1981). Spiophanes bombyx is thought to be the only spionid that displays this unique behaviour.











R98 High

R99 High


Slow, free movement through the sediment matrix R2 HighR100 High R101 HighR116 Medium

R124 High

Expert Opinion #N/AR167 Medium

R3 Medium


R47 Medium

As is the case for all octocorals, sea pens are actually colonies of polyps. What distinguishes sea pens is polyp dimorphism. One polyp grows very large and loses its tentacles, forming the central axis.




R137 Medium

Species Gro

wth

form

Mob

ility

/mov

emen

t

Feed

ing

met

hod

Typi

cal f

ood

type

s

Bio

turb

ator

Env

ironm

enta

l pos

ition

Hab

itat

Siz

e

Abra albaAbra nitidaAmpelisca tenuicornisAmpharete lindstroemiAmphiura filiformisAphelochaeta marioniArenicola marinaBrissopsis lyriferaCallianassa subterraneaCalocaris macandreaeCapitella capitataCarcinus maenasCerastoderma eduleCirriformia tentaculataEchinocardium cordatumEchinus esculentusEuclymene oerstediiGalathowenia oculataGoniada maculateHediste diversicolorLabidoplax mediaLagis koreniLeptosynapta bergensisMacoma balthicaMagelona johnstoniMalacoceros fuliginosusMaxmuelleria lankesteriMediomastus fragilisMelinna palmataMicroprotopus maculatusMya truncataMysella bidentataNephrops norvegicusNephtys hombergiiNuculoma tenuis

Ocnus planciOwenia fusiformisPagurus bernhardusPhaxas pellucidusPhiline aperta

Phoronis muelleriPhotis longicaudataPolydora ciliataPygospio elegansRhodine graciliorSagartiogeton undatusScalibregma inflatumScoloplos armigerSpiophanes bombyxThysanocardia proceraTubificoides (pseudogaster)Virgularia mirabilis

Literature ReviewExpert OpinionData Gap


Dis

tribu

tion

Dep

th ra

nge

Sub

stra

tum

pre

fere

nce

Tida

l stre

am p

refe

renc

es

Phy

siog

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refe

renc

es

Sal

inity

pre

fere

nce

Life

span

Sen

sitiv

ity to

cha

nge

Tole

ranc

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ano

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cond

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s

Key

pre

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Tem

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pop

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Con

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to o

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hab

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Rel

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A5.331; A5.334; A5.335; A5.355A5.341; A5.351A5.333; A5.334; A5.335A5.334A5.353A5.341A5.342A5.363A5.362A5.362A5.336A5.332A5.341A5.336A5.331: A5.335; A5.353 (E.flavescens)N/A - Species of conservation importanceA5.334; A5.334A5.352A5.341; A5.342A5.331; A5335; A5.355A5.342; A5.331A5.334A5.336A5.362A5.355A5.334; A5.333A5.333A5.333; A5.335; A5.351; A5.353; A5355A5.361; A5.362A5.331; A5.334; A5.352; A5.353

Leve

l 5 b

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pes

appl

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ased

on

sele

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A5.344A5.355A5.332A5.355A5.343; A5.344A5.351A5.334; A5.351A5.335A5.336A5.336A5.352A5.332A5.355A5.333A5.331; A5.355A5.351A5.336A5.343; A5.354

Source Code Model Level Model Parameter A

R129 1. Regional to global drivers

Climate

R129 1. Regional to global drivers Water CurrentsR129 1. Regional to global drivers ClimateR130 1. Regional to global drivers ClimateR130 1. Regional to global drivers Climate


R132 1. Regional to global drivers Climate

R133 2. Water column processes

R134 1. Regional to global drivers Depth


R134 1. Regional to global drivers Sediment type


R135 1. Regional to global drivers TemperatureR135 1. Regional to global drivers Water CurrentsR136 1. Regional to global drivers DepthR136 1. Regional to global drivers TemperatureR136 5. Output processes Secondary ProductioR137 1. Regional to global drivers Depth

R137 5. Output processes Bioturbation



R138 2. Water column processes Light attenuation

R1391. Regional to global drivers Depth

R139 2. Water column processes Light attenuation

R1391. Regional to global drivers Climate / Depth

Climate / Temperature

Climate / Water Chemistry

R140 2. Water column processes Primary Production

R140 4. Habitat and Biological AssemblagSediment type

R1412. Water column processes Primary production


R1421. Regional to global drivers Geology

R142 1. Regional to global drivers Temperature


R142 2. Water column processes Temperature

R1422. Water column processes Water Chemistry

R1422. Water column processes Hypoxia

R1423. Local processes at the seabed Hypoxia

R142 3. Local processes at the seabed Microbial activity

R142 1. Regional to global drivers Water currents

R1423. Local processes at the seabed Hypoxia

R142 3. Local processes at the seabed Hypoxia

R142 3. Local processes at the seabed Anoxia

R142 6. Local Ecosystem functions Hypoxia / Anoxia

R142 6. Local Ecosystem functions Microbial activity


R1436. Local ecosystem functions Sediment stability

R1435. Output processes Biodeposition


Water Chemistry and Temperature

Wave exposure / Water currents

R143 6. Local ecosystem functions Sediment stability

R143 5. Output processes Bioengineering




R144 5. Output processes Microbial activity






R1455. Output processes Bioengineering

R1455. Output processes Bioengineering

R145

2. Water column processes Suspended Sedimen



R147 1. Regional to global drivers Wave exposure


R148 5. Output processes Oxygen


R117 6. Local ecosystem functions Bioturbation


R150 1. Regional to global drivers Water Currents




R154 1. Regional to global drivers Water Currents

R1541. Regional to global drivers

Water Currents

R155

1. Regional to global drivers

Geology

R155 5. Output processes Sediment

R156 3. Local processes at the seabed Food Resource



R157 3. Local processes at the seabed Food Resource

R158 6. Local ecosystem functions Food Resource

R158 6. Local ecosystem functions Biogeochemical cycli

R158 6. Local ecosystem functions Ecosystem functions

R158 6. Local ecosystem functions Habitat provision



R160 4. Habitat and Biological AssemblagWater Chemistry

R160 6. Local Ecosystem functions Biodeposition

R160 6. Local Ecosystem functions Food Resource

R161 5. Output processes Biodeposition



R162 6. Local Ecosystem functions Biodeposition

R162 3. Local processes at the seabed Food source


R163 6. Local Ecosystem functions Primary Production

R164 6. Local Ecosystem functions Food Resource

R166 5. Output processes Bioturbation / Bioeng


R168 4. Habitat and Biological AssemblagBioturbation



R169 2. Water column processes Hypoxia



R172 1. Regional to global drivers Climate


R172 2. Water column processes Temperature

R172 5. Output processes Biogeochemical cycli




R174 6. Local ecosystem functions Food ResourceR174 2. Water column processes Suspended Sedimen


R174 6. Local Ecosystem functions Habitat provision

R175 3. Local processes at the seabed Sediment type







R179 6. Local Ecosystem functions Habitat provision


R138 2. Water column processes Suspended Sedimen




R181 4. Habitat and Biological AssemblagBiodeposition






R183 5. Output processes BioengineeringR183 5. Output processes Bioengineering

R183 5. Output processes BioturbationR183 6. Local Ecosystem functions Bioturbation

R183 6. Local Ecosystem functions Microbial activityR183 7. Regional to global ecosystem funBiodepositionR183 5. Output processes Bioengineering


R185 6. Local Ecosystem functions Bioengineering

R185 7. Regional to global ecosystem funSediment stabilityR185 5. Output processes Bioengineering


R185 6. Local ecosystem functions Biodeposition

R185 7. Regional to global ecosystem funR185 6. Local ecosystem functions Bioengineering

R185 7. Regional to global ecosystem funBioengineeringR185 6. Local ecosystem functions BioengineeringR186 5. Output processes Secondary Productio


R187

1. Regional to global drivers

GeologyR147 1. Regional to global drivers Depth

R189 2. Water column processes Primary ProductionR189 5. Output processes Bioengineering

Biogeoch cycling/sediment stability/habitat provision

Linking to Fauna/Species specific focuDirection of interaction

Fauna / Reproduction

InputPropagule supply InputOxygen InputSalinity InputFauna Input

Primary Production Input

Water Chemistry Input

Benthic infauna Input


Epifauna Epifauna Input


Fauna Input

Fauna InputFauna InputBiological Production InputBiological Production InputFauna OutputPrimary production Input

Fauna Output

Oxygen Burrowers Output

Light attenuation Input

Primary ProductionInput

Light attenuationInput

Primary production Input

TemperatureInput

Bivalves Macoma balthica

Food sources Input

Fauna Input

Food sourcesInput

Primary production Input

OxygenInput

Oxygen Input

Oxygen Input

Oxygen Input

OxygenInput

FaunaInput

Microbial activityInput

Organic matter Input

Oxygen Input

Bioengineering / Bioturbation burrowing/tube dwellingInput

Infauna Predators Input

Infauna Polychaetes Input

Nutrient cycling Feedback

Nutrient cycling Input

Oxygen Output

FaunaOutput

FaunaOutput

BiodepositionOutput

Infaunal deposit feeders (Macoma balthica, Cerastoderma edule)Infaunal deposit feeders (Macoma balthica, Cerastoderma edule)Infaunal deposit feeders (Macoma balthica, Cerastoderma edule)

Fauna Tube-builders Output

Water currents Tube-builders Input

Microbial activity Microorganisms and algae Output

Interstitial fauna Benthic meiofauna Output

Benthic infauna Output

Benthic infauna Tube-dwelling infauna Output

Benthic infauna Burrowing anemones Output

Microbial activity Deposit feeders Input

Benthic infaunaOutput

Fauna Grazers Output

Epifauna Epifaunal bivalves Output

Water currents Input

Sediment stabilityInput

Biodeposition

Input

Fauna Output

Fauna Bioturbators Output

Suspended Sediment Input

Suspended Sediment Input

Interstitial Fauna Meiofauna Output

Tube-dwelling infauna (Owenia fusiformis)

Deposit feeding crustaceans and bivalves (Macoma balthica)

Infaunal deposit feeding bivalves (C. edule)



Oxygen Output

Arenicola marina Input

Biogeochemical cycling Output


Biogeochemical cycling Input

Nitrogen cycling Output


Seabed mobility Input

Sediment Feedback

Sediment Input

Bacteria and diatoms Output

Fauna Input

Fauna Input

Sediment Input


Fauna Bivalves Feedback

Benthic infauna Nephtys Output

Fauna Bivalves Output

Epifauna Bivalves Output

Biogeochemical cycling Meiofauna, nematodes Output

Brittle star A. filiformis

Habitat provision/Biod. enhancement

Predatory Infauna (Alitta (Nereis) and Glycera)

Fauna Brittlestars Output

Fauna Brittlestars Output

Higher trophic levels Brittlestars Output

Epifauna Brittlestars Output


Epifauna Output

Microbial activity Input

Epifauna Input

Nutrient cycling Echinocardium Input

Nutrient cycling Echinocardium Feedback

Higher trophic levels sea cucumbers Output

Interstitial fauna Output

Habitat provision Abra alba Output

Fauna Output

Habitat provision Input

Fauna Hermit crabs Output

Fauna Hermit crabs Input

Epifauna Actiniaria Output

Epifauna Actiniaria Output

Water Chemistry Input

Benthic infauna Sea Urchins Input

Propagul supply Input

Fauna Calcifying taxa Output

Biogeochemical cycling Calocaris macandreae Input

Deposit feeding sea cucumbers

Deposit feeding sea cucumbersDeposit feeding sea cucumbersDeposit feeding sea cucumbers

Scavenging predators (Carcinus maenas)

Scavenging predators (Carcinus maenas)

Habitat provision Calocaris macandreae Input

Fauna Bivalves FeedbackFauna Bivalves Input



Fauna Deposit feeders Input

Habitat provision Input

Biodeposition Input

Food resource Input

Recruitment Input

Habitat provision Tube building organisms Input

Habitat provision Tube building Phoronida Input

Sediment stability Input

Water currents Feedback

Light attenuation Input

Fauna Output


Biogeochemical cycling Thalassinid shrimps Input

Fauna Output

Microbial activity Burrowers Input

Thalassinid shrimpsInput

Sediment stability Thalassinid shrimps Input

Microbial activity Thalassinid shrimps Input

Tube building polychaetes (P. elegans, P. ciliata, O. fusiformis)




Tube building polychaetes (L. conchilage, Mellina cristata)Biogenic structures (Polychaete tubes / mollusc shells)

Bioturbating organisms (Upward and downward conveyors)Bioturbating organisms (Upward and downward conveyors)

Biogeochemical cycling / Organic matter

Microbial activity Burrowers Input

Microbial activityThalassinid shrimps

InputFauna Thalassinid shrimps Output

Fauna Callianassidae OutputNutrient cycling Thalassinid shrimps Input

Nutrient cycling Thalassinid shrimps InputExport of organic matter Infauna InputMicrobial activity Infauna Input

Oxygen Primary producers

Ampeliscidae

Biodiversity enhancement AmpeliscidaeWater currents Ampeliscidae

Fauna Ampeliscidae

Biogeochemical cycling Ampeliscidae

Biodiversity enhancement AmpeliscidaeHabitat provision Ampeliscidae

Biodiversity enhancement Tube-buildersSediment stability Tube-buildersFauna Amphipods

Fauna Ampeliscidae

Suspended Sediments InputWave exposure

Dissolved oxygen InputRecruitment Tube builders Feedback

Biogeochemical cycling/Sediment stability

Positive/Negative (if applicabMagnitude of Interactio Limitations in evidence

Medium

MediumNegative Small

MediumSmall

Positive Medium

Large

Negative Medium

Negative Medium

Large

Large

Negative Large

Positive MediumPositive MediumNegative MediumPositive Medium

LargeNegative Large

Large

Large

Large

Large

Positive Large

Positive Large

Positive Medium

Localised location and non-selective habitat typesLocalised location and non-selective habitat types

Localised location and non-selective habitat types

Depth differences may not have been sufficiently large for the relationships between thermal stratification on benthic production processes to become apparent

Medium

Large

Positive Large

Positive Large

Large

Negative Medium

Positive Medium

Negative Medium

Negative Medium

Negative Large

Positive Medium

Positive Small

Medium

Negative Medium

Negative Medium

Small

Medium

Medium

Positive Medium

Negative Medium

Positive Large

Positive Medium

Estuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal waters

Estuarine and shallow coastal waters

Estuarine and shallow coastal waters

Estuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal waters

Estuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal watersEstuarine and shallow coastal waters

Positive Medium

Small

Positive Small

Positive Small

Negative Small

Positive Medium

Positive Medium

Medium

Negative Medium

Negative Medium

Positive Medium

Negative Medium

Negative Medium

Positive Small

Positive Large

Negative Medium

Positive Medium

Positive Medium

Small

Based to some extent on muddy sediments.

Based to some extent on muddy sediments.

Based on an experimental study (Cockles from SW England)Based on an experimental study (Cockles from SW England)

Based on an experimental study (Cockles from SW England)

Positive Medium

Positive Medium Intertidal focused experiment

Positive Large

Medium

Negative Medium

Positive Medium

Negative Medium

Medium

Large

Large

Medium

Large

Large

Large

Medium

Small

Positive Medium

Positive Large

Positive Medium Based on Mytilus edulis

Positive Medium Relatively un-researched.

Study based on Mytilus edulis farms in SW Ireland

Positive Medium

Positive Large

Large

Positive Large

Positive Medium

Positive Medium

Positive Medium

Medium

Positive Large

Medium

Large

Large

Large Subtidal fine-sandy habitats

Large

Large

Small

Negative Medium

Positive Small

Positive Medium

Negative Large

Negative Medium

Positive Large

Feedback Medium

Feedback Medium

Based on Ophiothrix fragilis

Based on Ophiothrix fragilis

Info based on Ophiothrix fragilis

Based on sea cucumber Australostichopus mollis

Based on sea cucumber A. mollisBased on sea cucumber A. mollisBased on sea cucumber A. mollis

Study based on Paguristes eremita in Adriatic SeaStudy based on Paguristes eremita in Adriatic Sea

Sea Urchin Trispneustes gratilla Indo-Pacific

Feedback Medium

Negative SmallPositive Medium

Medium

Positive Medium

Large

Positive Large Intertidal sandflat

Positive Medium Intertidal sandflat

Positive Medium Intertidal sandflat

Positive/Negative Small Intertidal sandflat

Positive Medium

Positive Medium

Positive Small Fine sand

Medium Fine sand

Large

Positive Large

Positive Medium

Medium

Positive Medium

Positive Small

Positive Medium P. tyrrhena

Positive Medium P. tyrrhena

Positive Small P. tyrrhena

Phoronis harmeri, intertidal sand-mudflat

Positive Medium

MediumPositive Large

Positive HighPositive Medium

Positive SmallPositive SmallPositive Medium

Large Shallow estuarine systems

Medium

Medium Haploops niraeFeedback Haploops nirae

Large Haploops nirae

Medium Haploops nirae

Medium Haploops niraeMedium Haploops nirae

MediumMediumHigh

High

Positive Medium

MediumMedium

Information Summary

Currents may alter affecting larval distribution. Deeper water may become isolated through thermal stratification during summer and become deoxygenated.Increased rainfall may influence salinityClimate change will likely change species distribution.

For epibenthos, depth is the major factor and the sediment composition seemed less significant.

Temperature has a positive effect on benthic abundance, diversity and biomassTidal stress has a positive effect on benthic productionBiological production negatively related to water depthBiological production positively related to temperatureHighest biological production recorded for filter feeders, bivalves, omnivores, predators and arthropodsAs depth increases, light attenuation decreases, thus primary production decreases.

Significant relationship between depth and light attenuation

Temperature changes will affect reproduction, growth and mortality of benthic fauna. Currents may alter affecting larval distribution. Water quality may reduce affecting survivalship. Will most likely result in changes to species distribution boundaries which could effect key structural species and associated fauna / changes in biotic interactions. Plankton bloom might change. Has info on some specific species.

In temporal zones, seasonal variation in ambient temperature produces similar variations in water temperature. This increase in temperature causes primary production to begin and temperature will then become a limiting factor in its production.Atmospheric CO2 concentrations are increasing. The uptake of anthropogenic CO2 by the oceans induces changes to the seawater chemistry by decreasing pH levels and carbonate ion availability (Ocean acidification).

Ocean acidification depresses reproduction and growth. M. balthica ia an ecosystem engineer to coastal habitats. Ocean acidification can change population size and distrubtion, affecting coastal diversity and ecosystem functioning.

The major determinant of infaunal community composition is sediment granulometry, with depth being of secondary importance.

The major determinant of infaunal community composition is sediment granulometry, with depth being of secondary importance. For epibenthos, depth is the major factor and the sediment composition seemed less significant.

Depth has a negative influence on benthic production. Primary production decreases with depth, energy input to the benthos is then primarly sourced by pelagic phytoplankton.

Burrowing species create tunnels in the sediment which themselves provide a habitat for other burrowing or iquilinistic species. Oxygenation through the burrowing activity enables the development of a much richer and/or higher biomass community of species living within the sediment and not in contact with the surfaceBurrowing activity in muddy sediments ventilates the burrows ensuring the sediment is oxygenated to a much greater depth than would be the case in un-burrowed sediment.

Primary production by phytoplankton is a light dependent process that provides the energy to drive the plankton and microbial food web, typically takes place in the euphotic zone (depths to which 1% of surface light penetrates).Light (in various wavelengths) penetration of the seawater depends on atmospheric conditions (cloud cover), particles in the atmosphere, the angle of incident which is dependent on season. Within the water column light is absorbed or scattered by particles or organisms. Primary production is limited by the depth of light penetration, usually 150-200m, this is the Photic Zone. If water is turbid this could reduce at as little as 20-30m.The penetration of solar radiation leads to thermal stratification of the water column. In temperate seas during the summer months a thermocline develops at 100-150m below which the temperature markedly reduces. This thermocline disperses in winter months due to strong winds and decreasing atmospheric temperature.

Primary production is limited by the availability of nutrients within the water column.

September storms disperse the hypoxic water, eliminating the summer thermocline

As water temperature increases, the severity of hypoxia increases and tolerance of fauna to hypoxia decreaes.

Bacterial mats are an important source of organic matter in hypoxic conditions.

Poor bottom water circulation creates lower oxygen levels at the seabed

Predation declines significantly in low oxygen saturations.

During anoxia most polychaetes have some ability for facultative anaerobiosis.

Seasonal hypoxia increases the importance of microbes in the energy cyling and carbon remineralisation.

The benthic compartment receives phytodertritus settling out of the water column. Benthic suspension feeders actively filter algae from the lower layer of the water column.Sediment characteristics have a direct influence on the density, biomass, distribution and diversity of benthic communitiesPrimary production is the assimilation of carbon (via photosynthesis) and the uptake of nutrients such as nitrogen and phosphorous, creating biomass. The plants/phytoplankton is consumed directly by animals or their carbon is released upon death or decomposition. This creates detritus (seston) utilised by suspension feeders.

In muddy and silty habitats oxygen only penetrates a few millimeters into the sediment by physical diffusion. At the sediment/water interface there is a thin diffusive boundary layer (1mm thick), through which molecular diffusion is the dominant transport mechanism for dissolved materials.Hypoxia is often seasonally associated with the peak in annual temperature cycle during summer. The seasonal development of a thermocoline (in shallow coastal waters) can result in hypoxic bottom conditions.

The increasing input of anthropogenic nutrients (Eutrophication) increases the flux of organic carbon to the bottom, increasing the oxygen demand at the seabed through bacterial and metazoan respiratory processes (Enhancing hypoxia and anoxia). In a hypoxic zone, stress on benthic animals are exhibited by altered behaviroul patterns, decreased feeding and reproduction activity and changes in physiological functions before mortality. Hypoxia is a major factor in structuring benthic communities and their function.Prolonged hypoxia or anoxia allow sulphate-reducing bacteria to survive in surface sediments where they can produce potentially lethal concentrations of hydrogen sulphide (H2S) and alter other geochemical cycles. Sulphur bacterial mats are formed + increased prominence of anaerobic pathways.

Burrowing fauna leave their burrows and migrate to the sediment surface (sometimes even into the water column) during hypoxia, decreasing their bioengineering potential. Tube dwelling fauna will emerge from their tubes during hypoxia.

Hypoxia and anoxia will enhance the recyling of nutrients from the sediments, increasing primary production and the oxygen demand from the water column and seabed.

Bioturbation by infauna (burrowing and tube building) lead to irrigation of sediments distributing oxygen into deeper sediment layers.

Deposit-feeding infauna decrease the sediment stability due to their feeding activity, through extensive sediment reworkingDeposit-feeding infauna greatly increase biodeposition (material deposited as faeces or pseudo-faeces). This sediment is redistributed in the water column and material is deposited on the bed through the action of biodeposition and natural sedimentation. So deposit feeders mediate tHe stability of the sediments.Deposit-feeding infauna greatly increase biodeposition (material deposited as faeces or pseudo-faeces). This sediment is redistributed in the water column and material is deposited on the bed through the action of biodeposition and natural sedimentation. So deposit feeders mediate te stability of the sediments.

Tube-forming organisms stabilise the seabed, single tubes create turbulence and local erosion.

Irrigation of tubes may promote microbial growth and result in increased mucous binding of the sediment.

Tube-forming organisms alter the flow pattern above the interface. Protruding tubes change the near bed flow (density dependent). Single tubes create turbulence and local erosion.Bacterial films can have significant effects on sediment properties through increasing the adhesion between particles and altering granulometry . Extracellular and autolytic products of microorganisms living on the grains and within the interstices can also foster sediment stability through the accumulation of mucilaginous materials.

Interstitial fauna which excrete mucus (in first instance to trap detritus, bacteria and macromolecules) have a positive effect on sediment stability by binding sediment grains. Copepods and nematodes are considered important, although other taxa which excrete mucus for locomotion are also important, such as ciliates, turbellaria and nemerteans.

Polychaete tubes destabilise the sediment unless at significant densities due to the production of a skimming water flow (near-bed flow).

Burrowing anemones can markedly influence sediment shear strength, likely as a result of mucus spreading over the sediment causing binding. Bioturbation by deposit feeders results in increased depth of the aerobic habitat in the sediment and thus an increase in the surface area available for colonization by microorganisms. Also, as a result of the exchange of sediment pore water with the overlying water, continual bioturbation "increases the rate of nutrient mixing within sediment and accelerates the rate of flushing of metabolites and growth inhibitors out of sediment" thereby stimulating bacterial growth rates

Sediment is disturbed and has an increased erodability due to the burrowing, deposit feeding activity and bioturbation of crustaceans and bivalves.Sediment has an increased erodability due to the disturbance and loosening of the surface sediments by grazers (such as Hydrobia ulvae and Neomysis integer)Sediment stabilisation is influenced by the physical protection and armouring of the surface sediments by epifaunal bivalaves such as mussel beds and oyster reefs.

Increased cockle density led to a reduction in near-bed flow velocity due to the increased bed roughness (bed shear stress) created by the cockles. The exhalent jets of the cockles' siphons can also modify the flow.

Cockles create burrows and furrows which destabilise the sediment, particularly muddy sediments, and increase the bed roughness. Vertical burrowing and horizontal ploughing behaviour.

In response to increased SSC (Suspended Sediment Concentration) C. edule show an increased frequency of valve adduction. In response to elevated SSC levels, bivalves produce copious amounts of mucus which loosely bind the sediment particles together. The mucus bound sediment particles are ejected as pseudofaeces through the inhalent siphon by means of the sudden valve adduction.Stabilizing species at high densities, scaled to diameter of the structure, reduce movement of sediments induced either by water or by biotic forces. Examples include angiosperms, large tube-building polychaetes, and mats of small tube-building crustaceans and polychaetes.

Classical bioturbators lead to increases in re-suspension of surficial sediment, high rates of sedimentary movement, and reduced resistance to lateral forces. Examples include nuculanid bivalves and burrowing urchins. A second distinct group of bioturbators cause high rates of turnover of sediment from depth to surface, may increase rates of re-suspension, and often add emergent structures that cause physical advection; examples are Arenicolidae polychaetes and some thalassinid crustaceans

Waves are generally responsible for resuspending and sorting sediments. Wave action can also lift cohesive muds into suspension. During fair weather, wave action disturbs sediment down to water depths of 10-20m. Storm waves can affect sediment depths up to 200m. Waves are generally responsible for resuspending and sorting sediments, winnowing out the finer particles to be transported and deposited elsewhere by tidal currents that would otherwise be too weak to move them.

The meiofauna of the surficial sediment and the sediment around macrofaunal burrows contribute to the interfacial flux by mixing the pore water within the interstitium of the sediment or by irrigating microcavities and microtubes

Sediment oxygen uptake largely depends on water depth. There is a highly significant decrease with depth.

Hydrodynamics lead to differential paricle-size sedimentation in the English Channel.

Presence of Nephtys burrows increases bacterial film depth and oxygen uptake in burrows.

Buried A. filiformis contribute to the total O2 flux by their respiration and by the O2 uptake of the additional sediment surface that they create (i.e., the inner surfaces of their burrows). Furthermore, the extended arms of A. filiformis can increase the O2 uptake of the superficial sediment by disturbing the boundary layer flow and by moving particles while sweeping the sediment surface in circles

A. marina ventilates its burrows which attracts meiofauna and some macrofauna to distinct regions of the burrows. The deep burrower, Scoloplos armiger benefits from the irrigation and bioturbation of A. marina as the enhanced water inflow lowers the levels of total sulfide and increases oxygen levels in the sediment, possibly also increasing bacteria in the subsurface.Bioturbation and bio-irrigation transport oxygen and organic matter (such as nitrogen) deeper into the sediment and enhance the transfer of excretion products to the water column.Currents play a significant role in the distribution of nutrients and organic carbon recycled by or produced by benthic fauna

Bioturbation is known to influence benthic Nitrogen cycling, typically as enhancing de-nitirification resulting in a subsequent loss of N2 from the system. Nitrogen fixation is less well studied. Bioturbation has the potential to increase sulphate reducing bacteria in burrows which act as nitrogen fixers, thus bioturbation can lead to increased nitrogen fixation.

Ocean acidification (pH decrease of 0.3) significantly reduced the Sediment Community Oxygen Consumption rates (on average by 60%) and benthic nitrification rates (on average by 95%) during the pre-bloom period (Februrary). A slowdown of the nitrogen cycle during winter could have global impacts on couple nitrification-denitrification and on the pelagic nutrient availability.During storms combined with high rainfall in shallow marine environments, large quantities of sediment may be flushed out of estuaries onto the continental shelf by means of mobile fluid muds. Clay has cohesive properties, because electrostatic forces cause the particles to attract each other, thus forming clay flocs or gels. These particle bonds can make a flow strong enough to modulate turbulence or, if turbulent enough, a flow can break relatively weak clay flocs or clay gels into smaller constituents.

Cohesive sediments are composed primarily of clay-sized material with strong interparticle forces. Cohesive sediments consist of inorganic minerals and organic material. Inorganic minerals consist of clay minerals (e.g. silica, alumina, montmorillonite, illite, and kaolinite) and non-clay minerals (e.g. quartz, carbonates, feldspar, and mica, among others).

Discrete particles can become cohesive by the growth of bacteria and diatoms. The particles become covered with the developing growth of the bacteria and diatoms, causing cohesion to increase and roughness to decreases. Further binding is provided by bacterial secretion known as extracellular polymeric substances (EPS) forming cohesive networks between the diatoms.Indicates relative importance of functional groups in food web in fine sediments in terms of carbon flow. Detritus and POM are driving forces. Deposit feeders and bacteria are key functional groups in fine sediments above suspension feeders. Carnivores are more important than omnivores. Meiofauna relatively unimportant. Hydrodynamics seem to be the most important factor in the organisation of the benthic invertebrates. In the English Channel hydrodynamics have a complex pattern due to the large tidal range, the hydrographical influence of large rivers and the coast morphology.

Alitta (Nereis) and Glycera are relatively important predators and determine community structure in benthic soft-bottom environments, preying upon taxa such as Nephtys, Polydora, Streblospio, Scoloplos, phyllodocids and bivalves. Bivalves through filter-feeding (Gosling, 2003) and nutrient regeneration activities can control the phytoplankton diversity in the water column above them which then affects other ecosystem grazers

Ecosystem function of bivalves includes habitat provision to macro- and micro-organisms and species diversity, shell aggregate structure and stability, chemical cycling, flow dynamics, and sediment particle flux. Mussel beds enhance the abundance of other benthic species by providing refuge and a source of organically enriched biodeposits.Meiofauna (especially nematodes) likely to be important as bioengineers at a smaller scale. Bacterial facilitation and biogeochemical cycling functions.

Holothuroidea (sea cucumbers) preyed upon by Fish, Starfish and Crustaceans

Low dissolved oxygen in the water column triggers a response in hermit crabs. High anoxia may cause fatalities.

Ocean acidification is accompanied by a decrease in saturation of the calcium carbonate (CaCO3) minerals.

Acidification reduces the size of sea urchin larvae which will likely affect the benthic adult populations.

Biocalcification plays a crucial role in the carbon cycle

Massive aggregations of suspension-feeding brittlestars can have an important effect on water quality in coastal environments and may even help counteract some of the potentially harmful effects of eutrophication.Brittlestar beds on soft substrata contain a rich associated fauna. Deposit feeding fauna might find conditions more favourable under beds as a result of the increased deposition of organic matter (from brittlestar faeces).Brittlestars preyed upon by crabs, dragonets Callionymus lyra, plaice Pleuronectes platessa, the starfish Asterias rubens and Luidia ciliarisBrittlestars play a major role in pelago-benthic transfer of particles from the water column to benthic habitats due to their suspension feeding activityThe deposit-feeding sea cucumber A. mollis suppressed benthic microalgae and facilitated bacterial activity through grazing and bioturbation. The enhanced bacterial abundance facilitated mineralisation processes, reducing the sediment organic matter content. Nutrient efflux (NH4+) elevated in sediments with A. mollis.Sediment feeding sea cucumbers elevate nutrient concentrations through excretion, creating favourable conditions for bacterial activity.Sediment feeding sea cucumbers elevate nutrient concentrations through excretion, creating favourable conditions for bacterial activity.Grazing by the deposit feeding sea cucumbers significantly reduced surface sediment algal biomass (microphytobenthos).Bioturbation by burrowing Echinocardium enhances nutrient efflux (NH4+) from the sediments to the watercolumn (benthic-pelagic nutrient cycling).The abundance of spatangoid urchins is positively related to primary production as their activities change nutrient fluxes and improve conditons for the production by microphytobenthos. Benthic microphytes at the sediment surface used NH4+ as it was released from the sediment column below.

Bioturbation and bioengineering activities influence the distribution of organisms that are too small to engineer their own habitat e.g. nematodes and bacteria.Abra alba extends the habitat area of Nematodes to deeper sediments. Ecosystem engineering macrobenthos are essential for the survival of lower parts of the food webs e.g. foraminifera and nematodesCrabs continuously disturb and aerate muddy sediment layers, possibly enabling nematodes to penetrate deeper into the sediment. An increased depth of oxygenation was observed in relation to an increasing frequency of biological disturbance and size of the disturber. Crabs continuously disturb and aerate muddy sediment layers, possibly enabling nematodes to penetrate deeper into the sediment. Hermit crabs offer habitat provision to symbionts and epibiota, affecting abundance and distribution of other invertebrates.

Actiniaria provision of habitat, predators and food providers. Not only for amphipods but also fish and crustaceans (copepods, amphipods, porcelain crabs, hermit crabs and spider crabs)).Actiniaria play an important role in benthic–pelagic coupling as part of the benthic suspension feeding community (Sebens and Paine, 1978), transferring energy to the benthos from the water column and releasing metabolites, gametes, and offspring back into the water column

Temperature is considered to be the primary environmental factor controlling the physiology, phenology, planktonic larval duration and biogeography of marine invertebrates. Temperature controls the pace of development in marine larvae.

Through burrowing, burrow irrigation and feeding activities thalassinidean mudshrimps have profound effects on biogeochemical cycling. Burrow irrigation results in an influx of oxygen rich water into the burrow and creates an efflux of water containing dissolved nutrients from microbial decomposition processes.

Filter feeding bivalves can have a top-down control on phytoplankton abundance and bacterial communities.Turbidity can have a positive influencing effect on filter feeding bivalve populations.

Bivalves provide an important resource for the surrounding benthos

Areas with high densities of tube building polychaetes are more diverse in meiofaunal and macrofaunal communities

Provide refuge for other species that would otherwise be susceptible to predation

Through burrowing, burrow irrigation and feeding activities thalassinidean mudshrimps have profound effects on the structure of the benthic communities.

Non-assimilated material is returned to the water column as pseudofaeces and faeces. Bivalves increase deposition rates, enrich sediments and stimulate microbial growth.

Deposit feeders attain higher densities on soft muddy substrata in comparison to suspension feeders. Physical instability of reworked mud surface is stressful for suspension feeders due to clogging of filtering structures.

Tube dwellers reduce the velocity of near-bed flow which results in increased passive biodeposition, which in its turn increases silt/clay fractions and meiofaunal abundances within patches.

Meiofauna use the faecal pellets of Polydora ciliata as a food source

Some taxa recruit more successfully within P. elegans patches

Tube building Phoronis facilitate the abundance and richness of some infauna and provide refuge for small clams from predation by shore crabs. Some uncertainty as this was observed in the laboratory but not in the field.

Tube building organisms promote microphytobenthos and the development of diatom biofilms that have the potential to stabilise the sediment and increase gross primary production and respiration.

Benthic organisms alter water flow, therefore altering the resuspension of sediments which affects the availability of nutrients and oxygen

Strong relationship between suspended particulate material and light attenuation. High levels of suspended particulate material may restricte the availability of light. Statistical evidence confirms this hypothesis in coastal and transitional waters, but was less significant in the offshore.

Upward and downward conveyors ingest organic particles and move them through their guts. These biodeposits modify the overall biogeochemical reactivity of the sediment and the biological functioning

Upward and downward conveyors ingest organic particles and move them through their guts. These biodeposits modify the overall biogeochemical reactivity of the sediment and the biological functioning

Bioturbating thalassinid shrimps are important ecosystem engineers affecting sedimentary and biogeochemical properties and processes. The significance of biodeposition intensity is highest in zones of low-energy hydrological conditions associated with diffusion-dominated microhabitats (sandy/muddy sediment)Bacterial abundances have been shown to be higher along burrow walls compared to either surface or ambient sedimentP. tyrrhena increases the organic matter content in the sediment and benthic metabolism by incorporating organic detritus, such as seagrass debris, in burrow chambersP. tyrrhena is capable of sorting sediment grains; selected fine particles are incorporated into the burrow wall to stabilise the structure, resulting in a lower grain- size homogeneity along burrow wallBacterial abundance within burrow walls of Pestarella tyrrhena was significantly higher than at the sediment surface and in adjacent ambient sediment.

Thalassinid shrimps are among the most important ecosystem engineers in marine soft sediments

Remineralisation rates of organic carbon and nitrogen are positively correlated with the density of shrimp burrows

Biodeposition contributes to the sediment organic content

Local hydrodynamics are altered by the presence of tubes

Offer refuges from predation

Many engineers create very stable conditions for those species that are dependent upond them Amphipods represent a major component of the secondary production in coastal ecosystems

depth has a major influence on the effects of wave exposure to the seabed.

Tube builders provide a settlement surface for larval and postlarval benthic organisms

Infaunal burrows are dynamic systems with intense small-scale gradients that support the growth of complex microbial assemblages. The physicochemical properties of the burrows, their age and the irrigation pattern are amongst the factors that have been shown to affect the composition of the bacterial community along the burrow walls in artificial experimental systemsSome deposit feeding thalassinideans have been described as gardeners, attaching plant and organic material to their burrow walls, enhancing microbial growth and thus their own food availability. There are however interspecific differences.

Callianassidae have the highest sediment turnover rates of all known bioturbators, creating sediment mounds above their burrows. Sediments ejected are unconsolidated and prone to resuspension

Increased nitrogen mineralisation is either directly caused by bacteria or indirectly by increased organic content in thalassinidean burrows.

Benthic macroalgae in shallow estuarine systems have a high importance as oxygen producers, carbon fixers and food sources for grazers.Through tube building and bioturbation activity, ampeliscids could be seen as infaunal hydraulic ecosystem engineers that physically modify their habitat by altering the biogeochemistry fluxes and the composiiton of the surface sedimentsInhabitants of Haploops habitat are not only dependent upon the presence of Haploops but also upon the sediment conditions engineered by this species.

By actively filtering the water column with their antennae and producing large quantity of pseudofaeces and feces, Haploops contribute to enrich the sediment they colonise.By actively filtering the water column with their antennae and producing large quantity of pseudofaeces and feces, Haploops contribute to enrich the sediment they colonise.

Haploops habitat offers potentially rich environment for an abundant benthic fauna which is supported by the high mean densities reported in the Haploops community (1200 ind/m2). Numerous inhabitants of engineered habitats are dependent upon resources indirectly provided by engineer species.

Tubes of any taxonomic group increase spatial complexity of the bottom and promote diveristy and abundances of associated species

Ampeliscid species play significant ecological roles. Feeding primarly on phytodetritus they require a high flux of phytoplankton to the bottom and greatly influence pelagic-benthic coupling.

Sediment consists of organic and inorganic particles. Sediment can be Allochthonous (being imported from the external environment by the chemical or mechanical breakdown of continental rock into their mineral form) or Autochthonous (created by the breakdown of local rock and also containing body-parts from organisms living within the locality, and the precipitation of dissolved minerals).

Photosynthesis together with wind and wave action are the most important sources of dissolved oxygen in the marine environment

Source Scale/Location Author Year

General Hiscock 2004

General Hiscock 2004Mainly in Sea lochs of Scotland Hiscock 2004General Phillipart 2012General Phillipart 2012

General Eppley 1972

General Feely 2009

Lab based Van Colen 2012

Regional - Northern North Sea Basford 1990



National - UK Wide Bolam 2010

National - UK Wide Bolam 2010National - UK Wide Bolam 2010General Cusson 2005General Cusson 2005General Cusson 2005General Jones 2000

General Jones 2000

General Jones 2000

General Devlin et al 2009

General Devlin 2009

General Munn 2004

General Munn 2004

General Munn 2005

Middelburg and Soetaert 2004

Middelburg and Soetaert 2004



General Diaz and Rosenberg 1995














General Paterson and Black1999



General Paterson and Black 1999

General Paterson and Black 1999

General Probert 1984





General Ciutat2007

General Ciutat 2007

General Ciutat 2007

General Ciutat2007

Lab based Ciutat2006

Lab based Ciutat

2006

General Woodin 2010

General Woodin 2010

General Brown 2002

General Brown 2002

General Vopel 2003

Lab based Vopel 2003

Lab based Volkenborn and Reise 2006

Lab based Braeckman 2010

Local - SW Ireland Chamberlain 2001

Global Middelburg and Soetaert 2004

General Bertics 2010


Lab based Baas 2011

Lab based Baas 2011

General Huang 2006

General Huang 2006

Regional - English Channel Garcia 2011



International - Maine, US Ambrose 1984

General Meadows 2012





Regional, UK Hughes 1998



Regional - English Channel Dauvin 2013

Lab based MacTavish 2012




New Zealand Lohrer 2004

New Zealand Lohrer 2004

General, Review Francour 1997



Lab based Schratzberger and Warwick 1999

Lab based Schratzberger and Warwick 1999

Regional - Adriatic Sea Pretterebner 2012

Regional - Adriatic Sea Pretterebner 2012

General Vader 1984

General Daly 2008

General Sheppard Brennand 2010

Lab based Sheppard Brennand 2010



Lab based Pinn and Atkinson 2009

Lab based Pinn and Atkinson 2009

General Saraiva 2011General Saraiva 2011

General Saraiva 2011

General Saraiva 2011

Massachusetts Rhoads and Young 1970

Scotland, Firth of Forth Bolam and Fernandes 2003




Larson et al 2009

Lab based Larson et al 2009

Lab based Passarelli et al 2012

Lab based Passarelli et al 2012

General Devlin 2008

General Kristensen et al 2012



General Mermillod-Blondin 2011

Mediterranean Papaspyrou et al 2005





General Pillay and Branch 2011General Pillay and Branch 2011

General Pillay and Branch 2011General Pillay and Branch 2011

General Pillay and Branch 2011General Pillay and Branch 2011General Pillay and Branch 2011

General Barranguet et al 1998

General Rigolet et al 2014

Off the coast of Brittany, France Rigolet et al 2014Off the coast of Brittany, France Rigolet et al 2014

Off the coast of Brittany, France Rigolet et al 2014

Off the coast of Brittany, France Rigolet et al 2014

Off the coast of Brittany, France Rigolet et al 2014Off the coast of Brittany, France Rigolet et al 2014

Off the coast of Brittany, France Rigolet et al 2014Off the coast of Brittany, France Rigolet et al 2014General Rigolet et al 2012

General Rigolet et al 2012

General Masselink 2003General Brown et al 2002

General Brown et al 2002bGeneral Qian 1999

Comments/Further references

Pearson and Eleftheriou, 1981; Mitchell et al, 1980

Emerson, 1990

Revsbech et al. 1980; Gundersen & Jorgensen 1990

Rosenberg, 1980

Rosenberg, 1980

Rosenberg, 1980

Webb 1969, Frankel and Mead 1973.

Riemann and Schrage 1978

Eckman et al 1981

Rowe 1974, Rhoads and Boyer 1982

Driscoll, 1975, Yingst and Rhoads, 1980.

De Deckere et al., 2000; Widdows et al., 1998b

Orvain et al., 2004; Roast et al., 2004

Widdows et al., 1998b, 2002

Widdows et al., 1998b, 2002

Meyers et al., 1987; Aller and Aller 1992; Pike et al., 2001

Eckman et al. 1981; Eckman 1983; Mills 1967; Featherstone and Risk 1977; Woodin 1978; Brenchley 1982; Rhoads and Boyer 1982

Rhoads and Young 1970; Rhoads et al. 1978; Brenchley 1982; Lohrer et al. 2004 Huettel et al. 1996; de Vlas 1979; Suchanek 1983; Thayer 1983.

Ducheˆne and Rosenberg 2001

Reise and Ax, 1979; Reise, 1981, 1987; Lackschewitz and Reise, 1998

Yingst & Rhoads 1980, Mermillod-Blondin et al., 2004; Blackburn, 1988.

Hayter, 1983

Paterson (1997)

Blanchet et al., 2004

Prins et al., 1995

Mermillod-Blondin and Rosenberg (2006)

(Gounin et al., 1995; Davoult and Gounin, 1995; Allen, 1998

Uthicke 2001; Plante et al., 1990

Uthicke 2001; Plante et al., 1990

Reise , 1985; Mermillod-Blondin et al. 2004

Bouchet et al., 2009; Van Colen et al., 2009

Warwick et al., 1990

Brooks and Mariscal, 1986, Williams and McDermott, 2004).

Tyrrell 2008

Zemlys 2003

Noji, 1994

Bolam and Fernandes 2002

Woodin 1978, Haines and Mauer 1980, Woodin 1981, Stachowicz 2001

Larson 2007

Nowell & Jumars 1984, Friedrichs et al 2009

Heip et al 1995

Rigolet et al 2011

Jones et al 1994

Young and Rhoads 1971Jones et al 1997

Grebmeier and McRoy, 1989

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Code Full Reference

R1

R2

R3 MarLin Biotic. http://www.marlin.ac.uk/biotic

R4

R5

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R7

Connor,D.W., Allen,J.H., Golding, Howell,K.L., Lieberknecht,L.M., Northen,K.O., Reker,J.B., (2004) The Marine Habitat Classification for Britain and Ireland Version 04.05. JNCC, Peterborough ISBN 1 861 07561 8 (internet version) www.jncc.gov.uk/MarineHabitatClassification

Queiros, M., Birchenough, S., Bremner, J., Godbold, J., Parker, R., Romero-Ramirez, A., Reiss, H., Solan, M., Somerfield, P., Van Colen, C., Van Hoey, G., Widdicombe, S. 2013. A bioturbation classification of European marine infaunal invertebrates, Ecology and Evolution, 3, 11.

Budd, G. 2007. Abra alba. A bivalve mollusc. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 21/10/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2307>

Allen P.L. 1983. Feeding behaviour of Asterias rubens (L.) on soft bottom bivalves: A study in selective predation. Journal of Experimental Marine Biology and Ecology, 70, 79-90.

Van Hoey, G., Vincx, M. and Degraer, S. 2007. Temporal variability in the Abra alba community determined by global and local events. Journal of Sea Research, 58, 144-155

Saskiya Richards 2007. Abra nitida. A bivalve mollusc. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 21/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=2308>

R8

R9

R10

R11

R12

R13

R14

R15

Sheader, M., 1979. Production and population dynamics of Ampelisca tenuicornis (Amphipoda) with notes on the biology of its parasite Sphaeronella longipes (Copepoda). Journal of the Marine Biological Association of the United Kingdom, 57, 955-968

Dauvin,J.C. and Zouhiri,S. 1996. Suprabenthic crustacean fauna of a dense Ampelisca community from the English Channel. Journal of the Marine Biological Association of the United Kingdom. 76. 909-929.

de-la-Ossa-Carretero, J.A, Del-Pilar-Ruso, Y., Giménez-Casalduero, F., Sánchez-Lizaso, J.L,, Dauvin, J-C. Sensitivity of amphipods to sewage pollution. 2012. Estuarine, Coastal and Shelf Science. Volume 96. 129-138

Lincoln, R.J. British Marine Amphipoda: Gammaridea.1979. London British Museum (Natural History), 658 pp.Marine Species Identification Portal http://species-identification.org/index.phpHolthe, T. 1986. Polychaeta Terebellomorpha. Norwegian University Press. 192

Jacqueline Hill and Emily Wilson 2008. Amphiura filiformis. A brittlestar. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 22/10/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2503>

Loo, L.-O., Jonsson, P.R., Skold, M. & Karlsson, O. 1996. Passive suspension feeding in Amphiura filiformis (Echinodermata: Ophiuroidea): feeding behaviour in flume flow and potential feeding rate of field populations. Marine Ecology Progress Series, 139, 143-155.

R16

R17

R18

R19

R20

R 21

Bruggeman, O.; Dupont, S.; Mallefet, J.; Bannister, R.; Thorndyke, M.C. (2003). Bioluminescence in the ophiuroid Amphiura filiformis (O.F. Müller, 1776) is not temperature dependant, in: Féral, J.-P. et al. (2003). Echinoderm Research 2001: proceedings of the 6th European Conference on Echinoderm Research, Banyuls-sur-mer, 3-7 September 2001. pp. 177-180

Skold, M., Loo, L.-O., Rosenberg, R. 1994. Production, dynamics and demography of an Amphiura filiformis population. Marine Ecology Progress Series, 103, 81-90.

Duineveld, G.C.A., Kunitzer, A., Heyman, R.P. 1987. Amphiura filiformis (Ophiuroidea: Echinodermata) in the North Sea. Distribution, present and former abundance and size composition. Netherlands Jounral of Sea Research, 21 (4), 317-329.

Rayment, W. 2007. Aphelochaeta marioni. A bristleworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 23/10/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2560>

Dr Harvey Tyler-Walters 2008. Arenicola marina. Blow lug. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 23/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2592>

Retraubun, A.S.W., Dawson, M., Evans, S.M. 1996. Spatial and temporal factors affecting sediment turnover by the lugworm Arenicola marina (L.). Journal of Experimental Marine Biology and Ecology, 20 (1-2), 23-35.

R22

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R27

Flach, E.C. 1992. Disturbance of benthic infauna by sediment-reworking activities of the lugworm Arenicola marina. Netherlands Journal of Sea Research, 30, 81-89.

Budd, G. 2004. Brissopsis lyrifera. Spiny mudlark. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 23/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2801>

Hollertz., K., Skold, M. and Rosenbergm R. 1998. Interactions between two desposit feeding echinoderms: the spatangoid Brissopsis lyrifera (Forbes) and the ophiuroid Amphiura chiajei Forbes. Hydrobiologia, 375-76, 287-295.

Hill, J. 2005. Callianassa subterranea. A burrowing mud shrimp. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 23/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2836>

Pinn, E.H., Atkinson, R.J.A., Rogerson, A. 1998. The diet of two mud-shrimps, Calocaris macandreae and Upogebia stellata (Crustacea: Decapoda: Thalassinidea). Ophelia, 48 (3), 211-223.

Riley, K. and Bilewitch, K. 2009. Capitella capitata. Gallery worm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2875>

R28

R29

R30

R31

R32

R33

R34

Tsutsumi, H. 1987. Population dynamics of Capitella capitata (Polychaeta, Capitellidae) in an organically polluted cove. Marine Ecology Progress Series, 36 (2), 139-149.

Neal, K. and Pizzolla, P. 2008. Carcinus maenas. Common shore crab. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2885>

Dr Harvey Tyler-Walters 2007. Cerastoderma edule. Common cockle. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=2924>

Hill, J. 2008. Echinocardium cordatum. Sea potato. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=3228>

Budd, G. 2008. Hediste diversicolor. Ragworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=3470>

Mayhew, E. and Bilewitch, J. 2009. Lagis koreni. A bristleworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 28/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=3610>

Budd, G. and Rayment, W. 2001. Macoma balthica. Baltic tellin. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesimportance.php?speciesID=3749>

R35

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R37

R38

R39

R40

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R42

Marine Ecological Surveys Limited, 2008. Marine Macrofauna Genus Trait Handbook. Marine Ecologcial Surveys Limited, 24a Monmouth Place, Bath, BA1 2AY. 184pp. ISBN 978-0-9506920-2-9.

Hill, J.M. 2004. Sea pens and burrowing megafauna in circalittoral soft mud. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/habitatecology.php?habitatid=131&code=2004>

Susie Ballerstedt 2002. Mya truncata. Blunt gaper. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=3840>

Sabatini, M. and Hill, J. 2008. Nephrops norvegicus. Norway lobster. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=3892>

Budd, G. and Hughes, J. 2005. Nephtys hombergii. A catworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=3897>

Dr Harvey Tyler-Walters 2002. Ocnus planci. A sea cucumber. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=3935>

Neal, K. and Avant, P. 2008. Owenia fusiformis. A tubeworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesimportance.php?speciesID=4001>

Barnes, M. 2008. Phoronis muelleri. A horseshoe worm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=4109>

R43

R44

R45

R46

R47

R48

Hill, J. 2007. Polydora ciliata. A bristleworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=4165>

Penny Avant 2005. Pygospio elegans. A bristleworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=4227>

Emily Wilson 2007. Sagartiogeton undatus. A sea anemone. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=4289>

Emma Snowden 2008. Scalibregma inflatum. A polychaete. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=4299>

Hill, J. and Wilson, E. 2000. Virgularia mirabilis. Slender sea pen. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29/10/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=4579>

Sheader, M. 1998. Grazing predation on a population of Ampelisca tenuicornis (Gammaridae: Amphipoda) off the south coast of England. Marine Ecology Progress Series. 164, 253-262.

R49

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R51

R52

R53

R54

R55

R56

Sievers, D., Friedrich, J.P. and Nebelsick, J.H. 2014. A feast for crows: bird predation on irregular Echinoids from Brittany, France. Palaios, 29 (3) 87-94.

Beukema J.J. 1985. Growth and dynamics in populations of Echinocardium cordatum living in the North Sea off the Dutch north coast. Netherlands Journal of Sea Research. 19 (2), 129-134.

Perez-Calderon, J.A. Occurrence of nematode parasites in Calocaris macandreae (Crustacea: Decapoda) from an Irish Sea population. Journal of the Marine Biological Association of the United Kingdom. 1986. 2, 293-301.

Ingle, R.W. and Christiansen M.E 2004. Lobsters, Mud Shrimps and Anomuran Crabs. Synopses of the British Fauna. No. 55

Southward, E.C and Campbell, A.C 2005. Echinoderms. Synopses of the British Fauna. No. 56.

Dr Harvey Tyler-Walters 2005. Labidoplax media. A sea cucumber. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 30/10/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=3597>

Grisley, MS.., Boyle, P.R., Pierce, G.J., Key, L.N. Factors affecting prey handling in lesser octopus (Eledone cirrhosa) feeding on crabs (Carcinus maenas)

Hayward, P.J. and Ryland, J. S. 1995. Handbook of the marine Fauna of North-West Europe.

R57

R58

R59

R60

R61

R62

R63

R64

R65

Hartmann-Schroder G. 1996.Die tierwelt Deutschlands und der angrenzenden Meersteile nach ihren Merkmalen und ihren Lebensweise: 58. Teil: Annelida - Borstenwurmer - Polychaeta. 2, neubearbeitate Auflage. VEB Gustav Fischer Verlag Jenam pp.648

Bestwick, B.W., Robbins, I.J., Warren, L.M. Metabolic adaptations of the intertidal polychaete Cirriformia tentaculata in an oxygen-sink environment. Journal of Exprimental Biology and Ecology, 125, 193-202.

George, J.D. 1971. The effects of pollution by oil and oil-dispersants on the common intertidal polychaetes, Cirriformia tentaculata and Cirratulus cirratus. Journal of applied Ecology, 8 (2), 411-420

Clavier J. 1984. Production due to regeneration by Euclymene Oerstedi (Claparede) (Polychaeta: Maldanidae) in the maritime Basin of the Rance (Northern Britanny). Journal of Experimental Marine Biology and Ecology, 75, 97-106.

Fauchald, K. (2008). Cirriformia tentaculata. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=129964

Michael, R. (2014) Ampharete lindstroemi Malmgren, 1867 sensu Hessle, 1917. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=129781

Rouse, G.W. and Pleijel, F. 2001. Polychaetes. Oxford University Press

Fauchald, K., Jumars, P.A. 1979. The diet of worms: A study of Polychaete feeding guilds. Oceanographic Marine Biology Anniversary Review. 17. 193-284.

Fauchald, K. (2008). Goniada maculata. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=130140

R66

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R68

R69

R70

R71

R72

R73

Mattson, S. 1981. Burrowing and feeding of Goniada maculata Orsted (Polychaeta). Sarsia, 66 (1) 49-51

Dobbs, F.C. and Scholly, T.A. 1986. Sediment processing and selective feeding by Pectinaria koreni (Polychaeta: Pectinariidae). Marine Ecology Progress Series, 29, 165-176.

Schuckel, S., Sell, A., Kroncke, I and Reiss, H. 2011. Diet composition and resource partitioning in two small flatfish species in the German Bight

Rijnsdorp, A.D. and Vingerhoed, B. 2001. Feeding of plaice Pleuronectes platessa L. and sole Solea solea (L.) in relation to the effects of bottom trawling. Journal of Sea Research, 45, 219-229

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Abstract

Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical processes in marine systems. In situ quantification of bioturbation can be achieved in a myriad of ways, requiring expert knowledge, technology, and resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption of a trait-based approach to estimate community bioturbation potential (BPc). This index can be calculated from inventories of species, abundance and biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking mode and mobility of benthic species. Based on information from the literature and expert opinion, we provide a functional classification for 1033 benthic invertebrate species from the northwest European continental shelf, as a tool to enable the standardized calculation of BPc in the region. Future uses of this classification table will increase the comparability and utility of large-scale assessments of ecosystem processes and functioning influenced by bioturbation (e.g., to support legislation). The key strengths, assumptions, and limitations of BPc as a metric are critically reviewed, offering guidelines for its calculation and application.

The behaviour of Asterias rubens (L.) feeding on a selection of common infaunal bivalves was investigated on a sand bottom in aquaria. A seasonal preference for Abra alba (Wood) and Spisulasubtruncata (da Costá) was noted. Nucula turgida (Leckenby & Marshall), although relatively numerous, was an insignificant component of the seastar diet. “Fitness” of prey as food species (mg AFDW of prey obtained per feeding bout) appeared to be the major factor determining preference. This was affected seasonally by accessibility of the highest ranked prey, Abra, through its increased burrowing activity. The investigation revealed that when presented with a selection of bivalves of varying accessibility, abundance, ease of opening, and food value, Asterias rubens was able to implement a flexible feeding policy consistent with the general rules of diet optimization.Shelf over a period of nine years (1995 – 2003). During this period, the community did not show a cyclic pattern, but a shift between the years 1995 – 1997 and 1999 – 2003 that was possibly triggered by changes in the hydroclimatic state of the North Sea and was reflected by a small shift of the dominant species in the A. alba community. In the years 1995 – 1997, the temporal pattern was dominated by strong year-to-year differences, coinciding with different successive events (strong recruitment, sedimentological changes, cold winters). Therefore, these years were characterised as the unstable period. Such events may affect the macrobenthic density, diversity and species composition. The mass recruitment of S. subtruncata may have been responsible for an initial decrease in the density and diversity of the macrobenthos, whereas the increase of mud content was responsible for a crash of the species richness and macrobenthic density in the winter of 1996. After those events, the A. alba community needed time to recover (1996 – 1998). The recovery was possibly retarded by a slow amelioration of the habitat conditions, unsuccessful recruitment and the occurrence of a cold winter. This period was followed by some years in which the seasonal dynamics in the community exceeded the lower year-to-year variability and therefore these years were characterised as the stable period. The seasonal dynamics in the study were characterised by high macrobenthic densities and diversity in spring and summer, followed by declines

Ampelisca tenuicornis Lilljeborg, 1855, is a member of a widely distributed genus of benthic tube-dwelling amphipods. Population studies of A. macrocephala Lilljeborg in the Øresund show it to have a generation length of 2 years (Kanneworff, 1965), and A. vadorum Mills off Massachusetts, U.S.A., produces two generations per year (Mills, 1967). A. brevicornis (Costa) in the Mediterranean breeds throughout the year with a generation time of 5–7 months (Kaim-Malka, 1969), but has only one generation per year in Helgoland Bight (Klein, Rachor & Gerlach, 1975).

Ninety-six species (97,677 individuals) were collected over the course of 6 h in five suprabenthic sledge hauls from a very dense Ampelisca fine sand community from the Bay of Morlaix (western English Channel). All the species migrated into the water column at night (98% of the specimens collected in the suprabenthos were found in the night hauls). The 23 most abundant species collected were classified into five groups based on their height within the water column, but two groups predominated: the upper suprabenthic species, abundant at 0-80-145 m above the sea-bed; and the lower suprabenthic species which were abundant only near the sea bottom (-0-1-0-75 m high). Three different patterns of nocturnal vertical migration were distinguished based on the timing of maximum swimming activity: at dusk; at the beginning of the night; or later in the night. Sexually dimorphic patterns of free-swimming behaviour was observed in Ampelisca and some other species of Amphipoda (Bathyporeia teniupes, Metaphoxusfultoni), and Cumacea (Bodotria pulchella, Pseudocuma longicornis), with many more males than females migratinginto the water column at night. Finally, the density of suprabenthic crustaceans in nocturnal hauls was amongst the highest reported from infralittoral or circalittoral suprabenthic studies on other parts of the Atlantic Ocean sampled during spring.

Amphipods are considered a sensitive group to pollution but here different levels of sensitivity were detected among species, by analysing the impact of five sewage outfalls, with different flow and treatment levels, on amphipod assemblages from the Castellon coast (NE Spain). Sewage pollution produced a decrease in the abundance and richness of amphipods close to the outfalls. Most of the species showed high sensitivity, particularly species such as Bathyporeia borgi, Perioculodes longimanus and Autonoe spiniventris, whereas other species appeared to be more tolerant to the sewage input, such as Ampelisca brevicornis. These different responses could be related to burrowing behaviour, with fossorial species being more sensitive and domicolous species being less affected. Benthic amphipods, which live in direct contact with sediment, are widely used for bioassay and numerous species are usually employed in ecotoxicology tests for diverse contaminants. In order to consider amphipods for monitoring and biodiversity programmes, it is important to establish the degree of sensitivity of each species to different sources of pollution

Experimental studies in a laboratory flume show that the sediment-living brittle-star Amphiura filiformis captures suspended particles. Feeding activity is a function of now velocity with few animals extending feeding arms in still water. Flow velocity also affects the orientation of feeding arms, and we suggest that this orientation is partly controlled by A. filiformis. By combining field measurements of current velocity and seston concentration with morphometrics and filtration models, a theoretical encounter rate of suspended particles was calculated for A. filiformis. In terms of organic content, A. filiformis can potentially balance growth and respiration with ingested seston although balance will strongly depend on retention efficiency and particle quality. Detailed measurements of flow around feeding arms revealed complex now patterns that will limit the applicability of available models of food encounter for passive suspension feeders, but our sensitivity analysis indicates that suspended aggregates may be especially important in the nutrition of this species.

Influence of temperature on the bioluminescence in the ophiuroid Amphiura filiformis was experimentally tested. Amputated arms from individuals acclimated at 6° or 14° were incubated at five different temperatures ranging from 6° to 14° for 2 hours before KCl stimulation. Light response comparisons do not reveal any difference neither for the two temperatures of acclimation nor for the five temperatures of incubation. Therefore, the temperature does not affect the studies parameters of the light emission.

Secondary production and population dynamics of the brittle star Amphiura filiformis (O. F. Muller) were studied between June 1990 and November 1991 at 40 m depth in the Skagerrak, west Sweden. Mean abundance of individuals > 1. mm in disc diameter was stable around 280 ind. m(-2) After a spawning maximum in June and July, settling post-larvae (ca 7000 ind. m(-2)) occurred in autumn 1991. Disc growth and gonad production accounted for ca 68.9 % (1.8 g AFDW m(-2) yr(-1)) of the total annual production in the population. About 13.3 % (0.34 g AFDW m(-2) yr(-1)) of the total production was allocated to regeneration of arms, probably a result of cropping by predators. Mean regenerated biomass in percent of total biomass for adult A. filiformis was between 12 and 30 % (mean 22 %). Annual production/biomass ratio was 0.46 yr(-1). The input of energy to arm regeneration indicates the importance of A. filiformis as an important food source.During the North Sea Benthos Survey undertaken by the ICES Benthos Ecology Working Group in spring 1986, a synoptic inventory was made of the benthos in the southern, central and part of the northern North Sea. The present paper deals with the status of the population of the infaunal ophiuroid Amphiura filiformis on the basis of measurements from 150 stations. A. filiformis was found at all the offshore muddy stations, but densities were generally highest in the shallow area south of the Dogger Bank. Stations with more than 1000 ind·m−2 were mainly situated near the frontiers between turbid and summer-stratified water masses, viz. along the southern border of the Oyster Ground, the southern slope of the Dogger Bank and in the northern part of the Pleistocene Elbe river valley. The size-distributions of A. filiformis show that juveniles were generally scarce at stations with high numbers of adults, whereas highest numbers of juveniles occur at stations with few adults. An unequivocal relation between adults and juveniles was, however, absent. This stresses the importance of other factors involved in recruitment. Comparison between the present and former (1938 and 1950) density of A. filiformis suggests that density has increased in the shallower parts but has remained stable in the deeper northern North Sea. An increase of A. filiformis shallow part of the North Sea could point to an enhanced food supply for the benthos, which may have some relation to the eutrophication in nearshore areas.

There were substantial differences in the amounts of sediment reworked by Arenicola mal ina (lugworms) on different parts of the shore and at different times of the year. The amounts were affected by the density, distribution and movement of worms. Rates of cast production were also important, and they were affected by the nature of the substrate and temperature. The production of casts occurred at regular intervals during periods of high activity.

The influence of the lugworm Are nicola marina on the abundance of other benthic species was studied in the westernmost part of the Wadden Sea. Small squares (1 m2) within depopulated 144-m2 plots were recolonized with various (naturally-occurring) lugworm densities (0-10-20-40-80 and 0-25-50-75-100 per m2). These plots were sampled during the summer. Lugworms were found to have a strongly negative effect on the densities of C. volutator. At 0-density lugworms, the numbers of C. volutator were high. These were halved at 17 lugworms per m2 (i.e. the mean density on the tidal flats of the Dutch Wadden Sea), and were further reduced at higher lugworm densities (e.g. 20% remained at 40 lugworms per m2). Laboratory observations of Corophium behaviour in the presence of Arenicola suggest that sediment-reworking lugworms stimulate Corophium to emigrate. Effects of lugworms on other benthic species were also studied in the same way. Lugworms were found to have strongly negative effects on the juvenile densities of various worm and bivalve species (Nereis diversicolor, Nephtys hombergii, Heteromastus filiformis, Scoloplos armiger, Pygospio elegans, Capitella capitata and Mya arenaria, Cerastoderma edule, Macoma balthica, Angulus tenuis, respectively).

Amphiura chiajei and Brissopsis lyrifera typically co-occur on some soft bottom areas of the North Sea, the Skagerrak and the Kattegat; they form the so called 'Brissopsis-chiajei association'. Both species are deposit feeders that live partly (A, chiajei) or completely (B. lyrifera) burried in the sediment. In this association, each species is expected to affect the other one, notably through its feeding and burrowing activities. This study investigates the effects on body and gonads growth in A. chiajei and B. lyrifera as a result of their co-occurrence. The experiments were performed in aquaria with natural sediments (3 months observations) and have taken in account the population densities of both species and the availability of food. The results show that B. lyrifera can negatively affect the growth of body and gonads of A. chiajei, while A. chiajei seemingly has no effect on the growth of B. lyrifera. This situation probably results from the extensive bioturbation of the sediment by B. lyrifera, who also increased its surface feeding activity when food was added to the sediment surface.

The monthly variations in the gut contents of Calocaris macandreae, principally a deposit feeder, and Upogebia stellata, principally a suspension feeder, were examined over the period of a year. The diet of both species consisted of a mixture of organic and fine inorganic fragments. The identifiable components of their respective diets included diatoms, dinoflagellates, algal and terrestrial plant fragments, and material of animal origin. The contribution of these different dietary components varied between the two species, and also over the year within each species. Using Strauss' Linear Food Selection Index (L) it was found that U. stellata positively selected diatoms, dinoflagellates and plant and algal fragments whilst avoiding animal material. In contrast, C. marandreae showed little evidence of food selectivity, except in the case of algal material which was positively selected. In general, variations observed in the diet were related to changes in food abundance and availability within their benthic habitat.

The population dynamics of a sibling species of Capitella capitata (Fabricius) in a cove organically polluted by fish farming were characterized by early recolonization in azoic areas, very rapid population growth, and extinction following development of extremely anaerobic conditions during summer. The remarkably high potential for population growth results from the short life cycle and continual reproduction as a population. Previously, the production of planktonic larvae with the ability to disperse over a wide area was considered one of the most important life history characteristics of opportunistic polychaetes in unpredictable habitats. In the present study, however, Capitella sp.exhibited opportunistic population dynamics during benthic recovery after defaunation in heavily polluted areas, despite production of a small number of large eggs with restricted dispersal ability via lecithotrophic larvae. The size composition of the recolonizing population suggested that some of the worms were recruited as mature adults from a very small population maintained in the immediate vicinity of the sampling station in an environment disturbed by organic pollution. Although the population dynamics of Capitella sp. appear to be characteristic of highly opportunisitc organisms, results suggest that populations may be maintained within each habitat by reliance only on a remarkably large potential for population growth. In general, organically polluted areas may not be temporary habitats that Capitella sp. can utilize only during benthic recovery immediately after defaunation, but rather such areas may be their native habitats.

Grazing impact was assessed for a population of the tubicolous amphipod Ampeljsca tenuicornis from a shallow sublittoral muddy-sand community off the east coast of the Isle of Wight, England. Damage resulting from grazing predation or failed predator attack was indicated by the presence of tanned wounds. Almost all damage recorded was to appendages, with the pnncipal feeding structures, the antennae, accountlng for 84 O/o of total injuries and the urosomal appendages accountlng for 12%. Pereion and pleon limbs showed little damage. The pattern of injury among appendage groups and the intensity of grazing were both found to be dependent on body size and sex. At maturity, a proportion of time is spent in the water column and this is associated with changes in the pattern of injury and intensity of grazing. Seasonal grazing intensity correlated with temperature and was related to life-cycle charactenstics. The intensity, though not the pattern, of grazing was negatively correlated to the abundance of A. tenu~cornis.T he. impact of grazing on reproductive output was dctcrnined. Although grazing on the population was high, rapid regeneration and compensatory feeding appeared to minimlse the impact The length of antennae and their relative proportions are important taxonomic characters used to separate species; if antennal grazing is a common feature of ampel~scldp opulatlons, then due care should be taken in selecting undamaged holotype matenal and in the use of antennal characters in identification keys.

Selective feeding behavior of birds on burrowing irregular spatangoid sea urchins, normally out of reach for such predators, is described. Behavioral observations were made on Carrion Crows (Corvus corone) feeding on spatangoid sea urchins (Echinocardium cordatum) during low tide on two different beaches in Brittany, France. The on-site searching behavior of individuals and flocks of crows on the surf line was observed together with characteristic feeding traces on the sediment. Wound morphologies on collected tests were documented with respect to possible preservation potential and recognition in the fossil record. Documented traces on feeding sites allow for clear identification of predator species and can be linked to observed bird behavior. The two sites differ in sea urchin size and the resulting predation traces on the echinoid tests. Predation on smaller specimens fragments the test, whereas predation on larger specimens leaves a characteristic wound morphology that is mainly restricted to the aboral side of the test. The wound morphology resulting from test puncturing involves both extensive inter- and intraplate fragmentation but does not completely destroy the tests. These predation traces are compared to other observed records of bird predation on echinoids. The potential for recognition and preservation in the rock record is then discussed. For 20 years (1969-1988), larger bottom animals have been sampled quantitatively once or twice annually at 15 stations scattered over Balgzand (a 50 km 2 tidal flat area in the westernmost part of the Dutch Wadden Sea). In 29 species, numbers were sufficiently high to allow a statistical evaluation of the fluctuation patterns of their annual densities. The results revealed two main trends: (1) a sensitivity to low winter temperatures in 12 species, causing low densities in these species immediately after a severe winter (1979, 1985, 1986 and 1987) and relatively high densities during a period with some mild winters in succession (1973-1975); (2) an upward long-term trend in 11 (other) species, causing upward trends (viz. roughly a doubling) of total macrozoobenthic biomass and production over the 20-year period of observation, probably as a consequence of increasing eutrophication. By far the major part of the species thus exhibited either of these two patterns, causing total biomass and species number to be governed largely by the above two trends. Results of less frequent sampling (once per 5 or 10 years) of 26 transects scattered over the --500 km 2 of tidal fiats of the whole western half of the Dutch Wadden Sea showed that the two trends also represent the changes occurring in a much larger area. Some local departures from the general patterns are discussed and related to specific causes.A number of nematodes are known to develop in decapod crustaceans. These parasite nematodes are present in the coelom of the host either free or surrounded by different types of host cells. All belong to the order Ascaridida or Spirurida and most of them develop only to the third larval stage in the decapod host; further development takes place in a predator of the crustacean which is generally a teleost or elasmobranch (Berland, 1961; Ouspenskaia, 1960; Petter, 1970; Poinar & Kuris, 1975;Uspenskaja, 1953; Yamaguti, 1961). The life-cycle in most cases is not fully understood. Ouspenskaia (1960) and Uspenskaja (1953, 1963) deduced the life-cycle for Ascarophis morrhuae van Beneden and A. filiformis Poljanski in the Barents Sea by relating the larvae found in decapod crustaceans through affinity of characters to the adults present in cod (Gadus morhua L.) and haddock (Melanogrammus aeglefinus L.). Similarly, the life-cycle of the spirurid Proleptus obtusus was described by Lloyd (1928); the larvae occur in a decapod crustacean, usually the hermit crab Pagurus bernhardus L. and in some cases the shore crab Carcinus maenas L. and the adults are found in the lesser spotted dogfish (Scyliorhinus canicula L.). A more complex life-cycle has been proposed for some anisakids such as Anisakis, Contracaecum and Hysterothylacium (Berland, 1961; Norris & Overstreet, 1976; Wootten, 1978) in which more than one intermediate host is required.

Octopuses (Eledone cirrhosa) feeding on crabs (Carcinus maenus) may penetrate the crab by a carapace borehole or puncture of the eye. In ad libitum feeding trials (632 crabs eaten), 31% of the predated crabs had a punctured eye, 57% had a borehole in the dorsal carapace. Eye puncture and boring occurred together in 6% of cases but 18% were neither punctured nor bored. Feeding trials in which size of prey and size of octopus were controlled showed that the incidence of boreholes was greatest (> 70%) in small crabs (<50 mm carapace width). Incidence of eye puncture (10% in small crabs) rose to 25% in crabs of over 50 mm carapace width and to over 40% in the largest crabs used (65-80 mm carapace width). Large octopuses used eye puncture less frequently than small octopuses. Increasing the proportion of small crabs in the diet increased the subsequent incidence of carapace boring at all crab sizes. The results are discussed in relation to differences in prey handling efficiency at different prey sizes.

The sediment around the burrow of the intertidal polychaete Cirriformia tentaculata (Montagu) is often hydrogen-sulphide rich and, thus, constitutes an oxygen-sink environment. Oxygen supply is maintained by filamentous branchiae that are spread over the sediment surface. Cirriformia is faced with tidally induced periods of environmental hypoxia and, in order to escape predation, burst-activity-induced anaerobiosis. Anaerobic energy metabolism, initiated at pO2 values of <30% saturation, is typically exploitative, with the accumulation of succinate and alanine. Pyruvate oxidoreductase activities are negligible and no cytosolic compensatory anaerobic pathways appear to be involved during intense muscular activity. It is suggested that this is an adaptation to life in such an oxygen-sink environment, as it avoids both the extreme acidosis and the incurring of an oxygen-debt synonymous with compensatory metabolic pathways.

When the Torrey Canyon ran aground on Seven Stones reef in 1967 there was little published information about the effects on marine life of oil and oil dispersants. This paper describes some observaions made in the field and the laboratory on two species of intertidal cirratulid polychaete which were exposed to an oil spill in September 1960.

The polychaete species Euciymene oerstedi (Claparede) forms one of the dominant populations of the muddy tine sand community in the maritime basin of the Rance (Northern Britanny). Analyses of regular samples of this species indicated that regeneration was an important process within the population. Of the individuals examined 22% were found to be regenerating the anterior and 41 y0 the posterior regions of the body. The distribution of the incision sites demonstrated a distinct preference for the body sections extending from both extremities to segment 3 or 18 inclusive, with the mean recovery times being 1.5 and 1 month respectively. Biomass production from regeneration within the population is estimated as 2 g. m - z. yr - i. From a comparison between the total amounts regenerated by individuals protected by cages and those in the surroundmg environment, it is suggested that posterior regeneration is essentially linked to epibenthic predation, while regrowth of the anterior region may also be a result of the activities of infaunal predators. The occurrence of these phenomena may be related to the behaviour of E. oerstedi within its tube.

Orth, 1973), in man-made harbours (Reish, 1959), and on coral reefs (Kohn & Lloyd, 1973), polychaetes arealso among the most 'species-rich' groups . They often comprise over one third the number of macrobenthic species and may be even more dominant in numbers of specimens (Knox, 1977) . Polychaetes may be numerically less important on hard substrata, and bivalves and various peracarid crustaceans may co-dominate in soft sediments, but of all metazoans only the nematodes are more ubiquitous . Polychaetes must thus be included in calculations of community trophic structure and in community energy budgets (e .g. Banse, Nichols & May, 1971 ; Pamatmat, 1977) . Despite their obvious importance the literature on ecological roles of polychaetes remains largely anecdotal. This review attempts to summarize current information about the feeding biology of these animals . We have organized the information into a limited number of patterns, using the guild concept to define our patterns . The concept of a guild (in the sense of a functional grouping) has proved to be a valuable tool both in generalizations and for continuing investigations in various animal and plant taxa (e.g. Grime, 1974 ; Karr & James, 1975) . Provisional attempts at delineating feeding guilds among benthic polychaetes (Jumars & Fauchald, 1977) allowed generalizations to be drawn and revealed several unexpected trends. There are other useful ways to form functional groups of polychaetes (e.g. by reproductive behaviours or

The burrowing and feeding habits are described in broad outline and compared summarily with those of Glycera alba. Whereas the latter is stationary, Goniada maculata is not. The two species may be found in the same locality and there is a clear division of food resources between them. Faecal food remains show that G. maculata feeds primarily on ‘sedentary’ polychaetes below the sediment surface in contrast to G. alba which feeds mainly on ‘errant’ polychaetes and amphipods on the sediment surface.

Pectinaria (Lagis) koreni (Malmgren) is an abundant, deposit-feeding, infaunal inhabitant of shallow-water marine environments in northern Europe. Laboratory experiments were performed to quantify the polychaete's sediment processing in 2 distinct sediments, 1 fine-grained and high in combustibles, the other coarse-grained and low in combustibles. Gut passage time and time to pseudodefecation were predictable only in coarse-grained sediment. In both sediments, reworking rate increased with worm size and over time, although temporal patterns differed in the 2 sediments. The ratio of pseudodefecated sediment to defecated sediment did not differ significantly over time in either sediment, but the ratio was greater in the fine-grained sediment during the first measurement period.2010 stomach contents of solenette and scaldfish and benthic infauna were sampled seasonally in a study area in the German Bight. The objectives were to investigate the seasonal variability of feeding activity and diet composition of both flatfish species related to benthic prey availability. For both flatfish, the highest feeding activity was found in summer, at the same time that the highest prey densities occurred in the study area. A reduced feeding activity was observed during the winter of 2010, but not in the winter of 2009, probably related to higher 2009 water temperatures. In all seasons, diet composition of solenette was dominated by meiofauna, mainly harpacticoid copepods. Macrofauna prey species, namely juveniles of bivalves and echinoderms became important in spring. An increase in amphipods and cumaceans was found in the stomach contents during summer and autumn, simultaneously with their increased abundance in the benthic infauna. In contrast, polychaetes were rarely found in the diet, but dominated the infauna during all seasons. Diet composition of scaldfish was dominated by larger and mobile prey, and, during all seasons, was mainly comprised of crustaceans. Amphipods characterised the diet in both winters, while decapods such as Crangon spp. and Liocarcinus spp. were the dominant prey from spring to autumn. Additionally, juveniles of flatfish (Pleuronectids) and bivalves were found in the scaldfish diet in spring, replaced by cumaceans in summer. No dietary overlap between both flatfish Stomachs of plaice and sole were collected in 1996 within and just outside the ‘plaice box’ (PB), an area in which fishing by vessels larger than 300 hp has been prohibited since 1989. In the mid-1990s the beam trawl fishing effort was reduced by 85% of the pre-closure level. In addition, a comparison was made of the diet composition of plaice and sole between the present and the beginning the 20th century. The diet of both species comprises mainly short-lived, highly productive benthic organisms. No difference could be found between the diets of fish sampled at grounds with different trawling intensities. The comparison of the present-day diet and the diet at the beginning of the 20th century suggests that the preponderance of polychaetes has increased and that of bivalves decreased. These results are consistent with the hypothesis that beam trawling has improved the feeding conditions for the two flatfish species by enhancing the abundance of small opportunistic benthic species such as polychaetes in the heavily trawled areas. However, the changes in diet may also be related to eutrophication and pollution.

The identification of magelonids with mucronate chaetae on chaetiger 9 has long been confused. Until 1977 all corresponding European specimens were erroneously referred to Magelona papillicornis; a Brazilian species. Since then, but without any detailed study, the name M. mirabilis (originally given to a species from Scotland) has been widely employed. However, in recent years, it has become clear that two morphologically similar species coexist in European waters. Magelona mirabilis is redescribed and a neotype designated, and M. johnstoni sp. nov. is formally distinguished. Following re-examination of the other five species present in the region, a dichotomous key and a synoptic table of characters is provided for all seven European species.

Since late 1990s the annelid polychaete Melinna palmata and the mollusc bivalve Ensis directus have been collected in the eastern part of the Bay of Seine (English Channel), indicating changes in the benthic communities. Melinna palmata was never collected prior to 2002, whereas it was reported in the muddy fine sands of the western part of the Channel, along the French (e.g. Bay of Cherbourg) and southern UK (e.g. Southampton Waters) coasts. Ensis directus was first reported in 1998 and now appears to be well implanted, given the abundant population collected in 2006. The colonization of Melina palmata seems to be a consequence of recent increase of the fine sediment in the eastern part of the Bay, while that of the invasive Ensis directus seems more likely to be related to its southwest expansion, from the Scheldt estuary (Belgium and Netherlands) towards the Bay. Since both species have complex life cycles including planktonic larval phases, their colonisation may also be favoured either by an accidental introduction via ballast waters or by larval dissemination from neighbouring populations.

The earliest detailed account of the nature of the sea bottom near Plymouth is that of Allen (1899), wherein analyses of the soils on the 30 fm. line are coupled with lists of the animals collected by trawl and dredge. Ford (1923) described a number of soils in shallower water, and gave a quantitative list of the bottom fauna, collected with a grab which covered an area of 0·1 sq. m. Smith (1932) described in great detail the soils of the area of shell-gravel which surrounds the Eddystone Lighthouse. By none of these workers, however, was special attention paid to the smaller burrowing Crustacea, which are often overlooked unless they are made the special object of collecting. Some species, e.g. of Bathyporeia and Ampelisca, may be very common, and certainly play an important part in the ecology of the sea-bottom.

of the facultative filter-feeding burrowing ophiuroid Amphiura filiformis and its coinhabitant, the bivalve Mysella bidentata in two gradients of pelagic productivity. In a temporal gradient, 1971-1994, with high productivity in the 80 s, abundance of Amphiura at 100 m depth showed logistic population growth (R(2) = 0.90, n = 33) with carrying capacity reached in 1986 (ca. 3000 ind m(-2)). Mysella abundance versus time was best modelled by an exponential increase, occurring mainly during the period when Amphiura had reached maximum density (1986-1994) (R(2) = 0.75, n = 18). Amphiura showed somatic growth during the 'plateau period' of similar magnitude as predicted from independent data from the Kattegat. These observations indicate that Amphiura not was food limited in the plateau period. In the spatial gradient, inferred from chlorophyll concentrations in the surface sediment at 60 sites bisecting the area of the Skagerrak-Kattegat pelagic front, where primary productivity is elevated, Amphiura reached maximum abundance at the same level as in the temporal gradient, while Mysella and total benthic biomass peaked in the area of highest phytoplankton input to the sediment (Josefson & Conley, 1997). High content of the plant pigments chlorophyll-alpha (Chl-alpha) and phaeopigments in Amphiura suggested ingestion of labile algal matter, and a tentative chlorophyll budget indicated that Amphiura metabolic demand could be accounted for by ingestion of relatively fresh algal derived matter if half-lives of total chlorophyll in Amphiura were on the order of 1/2-1 h. The results suggest that space may be a limited resource in some soft-

The survey of the sublittoral fauna of the Clyde Sea Area from 1949 onwards has shown that five species of the Protobranchiata are abundant throughout this region on a variety of substrata. Pelseneer (1891, 1899, 1911), Heath (1937), and Yonge (1939) have contributed much to the knowledge of the group as a whole, but little comparative work has been done at species level. Verrill & Bush (1897, 1898) studied the shell characters of the American Atlantic species. Moore (1931 a, b) worked on the faecal pellets of the British Nuculidae and attempted to distinguish the species by this means, while Winckworth (1930,1931), mainly in the light of the latter work, attempted to clarify the nomenclature of these species. Winckworth (1932) lists six British species of the family Nuculidae: Nucula sulcata Bronn, N. nucleus (Linné), N. hanleyi Winckworth, N. turgida Leckenby & Marshall, N. moorei Winckworth and N. tenuis (Montagu); and four species of the family Nuculanidae: Nuculana minuta (Müller), Yoldiella lucida (Loven), Y. tomlini Winckworth and Phaseolus pusillus (Jeffreys). All species of Nucula, except N. hanleyi, were taken from the Clyde Sea Area, although the latter species is included in the Clyde fauna list (Scott Elliot, Laurie & Murdoch, 1901). Only Nuculana minuta of the Nuculanidae has been taken on the present survey. Yoldiella tomlini is included in the 1901 list but is noted as being ‘insufficiently attested’. Nucula hanleyi was obtained from the Marine Station, Port Erin, but Yoldiella and Phaseolus were unobtainable.

Although dexaminid amphipods have been observed interacting with numerous other taxa, the particular relationships they share with echinoderms remain unexplored. This study examines interactions between Tritaeta gibbosa and the holothurian Ocnus planci, which the amphipod inhabits. Each amphipod creates its pit and propels itself through the mantle by actively pulling the mantle tissue using its pereopods. Many individuals were observed living in high densities in all areas of the O. planci mantle, with higher preference of the oral and "dorsal" sides, a trend that was corroborated by behavioral experiments. Experiments on behavior and anatomical studies of the amphipod and the holothurian host were performed in order to clarify the mechanisms behind settlement and pit formation, placement and location, as well as the amphipod's morphological adaptations to this peculiar life style. To investigate parasitism and allow for future identification, T. gibbosa, O. planci and Cucumaria montagui were also barcoded (CO1 and 16S), unfortunately with lower success.

1.5 mm s(-1). Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve. Simultaneous with stoppage of a particle, the oral hood is rapidly lifted so that water from the surrounding area flows in to occupy the volume created under the hood. Due to the proximity of the tentacle with the arrested particle, this suction will draw the particle away from the laterofrontal cilia into the increased space under the oral hood with a velocity of about 0.6 min s(-1) before it is subsequently carried to the mouth. If a particle is stopped near the tip of a tentacle this may trigger a tentacle flick, with a tip velocity of about 5-7 mm s(-1) which brings the particle down toward the lifting edge of the hood. A stopped particle may cause a local disruption of the metachronism of the lateral cilia for about 0.14 s. Likewise in the adult, when an incoming particle with a velocity of about 1.6 min s(-1) is stopped near the tip of a tentacle this triggers a flick, which brings the particle down towards the mouth. The duration of the active flick phase is about one-tenth of the flick cycle. Only when a particle is stopped on the outer part of a tentacle is a flick triggered. Otherwise the particle is either transported down along the frontal surface of the tentacle by means of the frontal cilia, or transferred into the downward current created by the lophophore. The metachronal wave velocity is about 0.25 mm s(-1), the wavelength about 12 mum, and hence the ciliary beat frequency about 21 Hz (at similar to16degreesC). Essential features of the A hitherto unreported association between the isaeid amphipod Photis longicaudata and the hexacorallian Cerianthus llopdii is described and illustrated from material collected near Millport, Clyde Sea area. By building their tubes around the outside rim of the anemones' tubes, this amphipod presumably gains proximity protection from predators.

Polydora ciliata is a spionid polychaete found below mid-tidal level burrowing in a variety of rocks all of which contain calcium carbonate. It can also penetrate some non-calcareous materials such as rotten wood. It excavates a U-shaped burrow which it lines with a tube composed of mucoprotein and sand grains. This tube has a smooth inner lining of mucoprotein. Both food and tube-building material is collected by the palps and it is suggested that discrimination is made chiefly on size of the particle. Selection is made between the limits of 003-0-05 mm, the smaller particles passing down the gut whilst the larger are used in construction of the tube. A particle selection mechanism is suggested. The function of the segmental mucus glands is to provide the inner lining of the tube. The problem of chemical penetration of rocks by animals is discussed. P. ciliata uses both mechanical and chemical methods. No acid has been identified, and the use of a sequestering or chelating agent linked with the biochemistry of mucus is suggested.

Megaclausia mirabilis n.gen., n.sp. a new copepod commensal with the maldanid polychaete Rhodine gracilior is described from material collected in the Firth of Forth and off the Northumberland coast. The species is a member of the family Clausiidae and is readily distinguished by the unusually large size of the female and by its leg structure. Other genera of the family are briefly reviewed and the position of Megaclausia is discussed.

Echinus esculentus plays a key role in the structure of subtidal communities. Large numbers were removed from Skomer MNR during the 1970s when divers targeted the population for the curio trade and a population survey was completed in 1979. No repeat surveys were completed until 2003 when data was collected to establish the status of both the E. esculentus population and conspicuous starfish species. This survey repeated the methods used in 2003 and established fixed surveys sites that can be used in future surveys. The survey was completed over 4 days by a team of 20 volunteer divers. E. esculentus, Marthasterias glacialis, Crossaster papposus and Luidia ciliaris were counted and the diameter of E. esculentus were measured along 30m transects. The study sites were selected from the north and south coasts of the island and the north coast of the mainland. The mean densities of E. esculentus and M. glacialis were 6.1 and 3.47 per 100m2 respectively for the whole MNR, but density varied between sites and depth. A normal size frequency distribution for E. esculentus was found and little variation in size range was observed no matter what the depth. Comparison of results with previous surveys and other areas in the UK suggest a naturally low density of E. esculentus in Skomer MNR possibly due to low recruitment. The same may be true for C. papposus and L. ciliaris. Plankton samples collected from June to September identified Echinopluteus larvae in samples in the last two weeks of July, confirming that spawning occurs during July.Several species of sea urchins are now being cultivated for commercial purposes and with the continued increased demand for sea urchin gonads as a food product, new species are being assessed for their aquaculture and market potential. This study focussed on establishing protocols for the production of common sea urchin Echinus esculentus larvae and juveniles to assess its potential as an echinoculture species. Two trials were carried out, the first trial evaluated the influence of three microalgal diets (D=Dunaliella tertiolecta only, mixed D/P= D. tertiolecta plus Phaeodactylum tricornutum and P= P. tricornutum only) on larval morphology. Larval length, width, post-oral arm length and rudiment length were significantly effected by diet. Diets D and D/P prompted more rapid metamorphosis. In the second trial, the effects of different rations of D. tertiolecta were tested. The food ration, standard ration (SR; 1000, 3000, and 5000 cells ml 1) and high ration (HR; 3000, 9000, and 15,000 cells ml 1) were increased as the larvae acquired the 3rd and 4th pair of larval arms. Larvae fed the SR were significantly larger (longer and wider) and had significantly longer rudiments than those in the HR treatment. The number of larvae metamorphosing and settling onto substrates was significantly higher in treatment SR compared to HR. Optimising the larval diet shortened the larval stage from 21–23 days in the first trial to 16 days in the second trial. The maximum percentage of metamorphosing individuals which survived to post-larvae or juveniles (10 days after they were first judged competent to settle) was 46.6%, suggesting E. esculentus is a viable aquaculture candidate.hypothesized. On the Western coast of Norway, a rocky reef with a highly complex topography has been chosen to be the first full-scale offshorewind farm in the country. Underwater video analyses and multibeam bathymetry data with a generalized linear model were used opportunistically to investigate the influence of geomorphic explanatory variables on the occurrence of selected taxa (algae, sea urchins and sea stars) identified in the study area. Combining video observations and multibeam bathymetry in a generalized linear model revealed that the geomorphic descriptors: aspect, slope, rugosity, and benthic position indexes (BPI), were of significance for algae, sea urchins and sea stars at Havsul and served in showing their habitat preferences. Kelp occurred in areas of high rugosity, on gentle slopes, at elevated areas with a southerly orientationnd on the sheltered side of rock or bedrock. Thus, construction disturbance that modify those variables may lead to a change in the area preferred by kelp. Turbines that shade southerly aspects may affect small kelp plants in reducing their available habitat. Sea urchins were more abundant on steep slopes and both sea stars and sea urchins showed a preference for a complex local relief (high rugosity) and heterogeneity infine and broad elevation (shown by BPI). Thus, foundations and cable route preparation may significantly change the slope, rugosity of BPI broad, which will change the basis for sea urchin populations. It may likewise significantly change the rugosity or BPI (fine or broad), which may change the distribution of sea stars. The combination of video data and models using multibeam bathymetry yields useful

As part of a long-term study to examine the ecological effects of beam-trawling, we investigated the immediate impact of fishing on the megafaunal component of a benthic community and the extent to which it had recovered 6 months later. A quantitative dredge was used to collect megafaunal samples following a replicated, paired control and treatment design to maximize the chances of detecting any effects due to trawling. There were two different habitats with distinct communities in the experimental area, one with stable sediments and a rich fauna, the other with mobile sediment and a relatively impoverished fauna. Immediately after fishing the composition of the community in the stable sediments was significantly altered. While the abundance of some species decreased (e.g. sea mice Aphrodita aculeata), others apparently increased (e.g. hermit crabs Pagurus bernhardus). Variation between samples from the fished areas was higher than those from the control areas. This suggests that the effects of trawling were not uniform, even though the treatment area was entirely swept at least once. The effects of fishing were not detectable in the mobile sediments. Six months later, seasonal changes had occurred in both communities and the effects of the trawling disturbance were no longer evident.Isle of Cumbrae (Scotland) were analyzed. The epibionts found were: (1) the protozoans Acineta, Conchacineta, Corynophrya, Zoothamnium, Cothurnia mobiusi, Cothurnia longipes, Chilodochona, Cryptacineta, Ephelota gemmipara and Ephelota plana; (2) the hydrozoans Leuckartiara, Clytia and Phialella; (3) the entoproct Barentsia; (4) the cirripeds Balanus balanus and Balanus crenatus; (5) the polychaetes Pomatoceros triqueter, Circeis amoricana paguri and Hydroides norvegica. Among these epibionts, a protozoan (E. plana), two hydrozoans (Leuckartiara, and Phialella), and the entoproct (Barentsia) have not been described previously as epibionts on crustaceans. The protozoan, entoproct and hydrozoan epibiont species found on P. bernhardus represent the first mention of their presence on this hermit crab. The protozoan epibionts were found only on the crab, while the hydrozoan epibionts were observed on the shell and on the crab. The cirriped species were observed only on the external surface of the shell, and the polychaete species were observed on the external and internal surfaces of the shell, although Circeis was found only internally. An analysis of the distribution and density of each epibiont species on the anatomical units of the crab, as well as on the different shell areas were made. The comparison between the data of the two years showed differences with respect to the size of crabs, diversity of epibiont groups and density of epibionts. There was a significant difference between the two years with respect to the distribution of epibionts on the anatomical units of the crab, both in terms of density and biomass. The influence of some During the last 20 yr the western half of the Dutch Wadden Sea has undergone significant eutrophication: concentrations of P and N compounds and planktonic algae have roughly doubled, as has primary production. Though oxygen levels are often low in summer, anoxic areas are small and rare due to strong tidal mixing. During the 1970 to 1990 period, macrozoobenthos was sampled annually at 15 stations at Balgzand, a 50-km 2 tidal-flat area in the westernmost part of the Wadden Sea. Not only did the estimates of total numbers, biomass, and production double during these two decades, but significant changes in the composition of the benthic community were observed, too: (1) the numerical proportion of polychaetes increased at the expense of molluscs and crustaceans, (2) the overall mean weight per individual of the macrozoobenthos decreased (numbers of individuals of small-sized species increased more rapidly than those of large-sized species), and (3) though absolute numbers and biomass of all feeding types increased, the share of carnivores declined and that of deposit feeders increased; the proportion of suspension feeders showed little change. This study refers to true macrobenthos only (l-mm sieve) and further excludes two taxa (Corophium spp. and Hydrobia ulvae) which occasionally exercised an undue influence on numbers. Mass mortalities caused by low oxygen concentrations were of a small-scale nature only. Total number of species fluctuated without a clear trend. As a consequence of the increasing numerical densities, trends in species numberswere slightly increasing when expressed per unit of area and slightly decreasing when estimated per 100 individuals(by rarefaction).The effects of trawling disturbance on a benthic community were investigated with a manipulative field experiment in a fine muddy habitat that has been closed to fishing for over 25 yr. We examined the effects of extensive and repeated experimental trawl disturbance over an 18 mo period on benthic community structure and also followed the subsequent patterns of recovery over a further 18 mo During the penod of trawl disturbance the number of species and individuals increased and measures of diversity (Shannon's exponential H' and S~rnpson'sre ciprocal D) and evenness decreased in the trawled area relative to the reference site. The cirratulid polychaetes Chaetozone setosa and Caulleriella zetlandica were found to be most resistant to disturbance. whilst the bivalve Nucula nitidosa and polychaetes Scolopolos armiger and Nephtys cirrosa were identified as sensitive species. Multivariate analysis and abundance biomass comparison plots confirmed that community changes occurred following disturbance. with some differences between treatment and reference sites still apparent after 18 mo of recovery. Physical effects, examined with Side-scan and RoxAnn, were identifiable immediately after disturbance, but were almost indistinguishable after 18 mo of recovery. Such long recovery times suggest that even fishing during a restricted period of the year may be sufficient to maintain communities occupying fine muddy sediment habitats in an altered state.

generates a pit-and-mound topography at the sediment surface from intertidal sands near the island of Sylt, Germany. This experiment was used to test whether other abundant deposit feeding polychaetes (the discretely motile and surface feeding ragworm Nereis diversicolor and the subsurface-feeding, motile orbiniid polychaete Scoloplos cf. armiger) benefit from competitive release. Ragworms took advantage from the absence of lugworms. Presumably they responded to a more stable and nutritious surface layer at lugworm exclusion plots (relief from inhibitive bioturbation). Contrary to this, S. cf. armiger was negatively affected by theexclusion of A. marina. It may have suffered from higher sulfide concentrations in the less irrigated and less permeable sediment where lugworms were absent. For adult worms of both species these results were consistent in 2 out of 3 years examined. Recruitment by N. diversicolor was highly variable between years and occurred either irrespective of experimental treatments or the response was inconsistent. Juveniles of S. cf. armiger benefited from the presence of A. marina and aggregated near lugworm tail shafts where inflow of oxygen rich water was high and sulfide concentrations were low. Biogenic habitat mediated effects of lugworms on both deposit feeders were in the same order of magnitude as abundance variation in space and time. Thus, A. marina was one of the key factors structuring the deposit feeding community. It is suggested that arenicolids modify the composition of the colonisation were investigated. Values of abundance and total numbers of species were significantly lower ð p < 0:05Þ in an area most recently exposed to the highest level of dredging intensity compared with samples taken from an area of low intensity, and those from a reference site. Differences between previously dredged sediments and the reference location were due to the reduced abundance of a range of macrofaunal species characterising nearby sediments. Multivariate measures of community structure also indicated that there were significant differences ð p < 0:01Þ between the macrofaunal assemblages in the areas exposed to different dredging intensities. Sediment from the area exposed to the highest dredging intensity contained proportionally more sand than other sampled sediments. The extent to which dredging intensity contributed to these differences was difficult to determine owing to the absence of any baseline data. Despite this, univariate and multivariate analyses indicated a strong relationship between macrofaunal community structure and dredging intensity at this site. Correlation analyses also demonstrated that the predominant influence on the macrofaunal community was that of the level of dredging that took place in 1995, the last year that the licensed site was dredged heavily. Preliminary observations indicated that the fauna remained in a perturbed state some 4 years after cessation of dredging. Therefore, relatively rapid recovery rates, commonly cited as 2–3 years for European coastal gravelly areas, should not be assumed to be universally applicable. Implications for the future management and scientific study of marine aggregate Sublittoral benthic coastal communities of the North Sea and of the Western Mediterranean were studied before and after sand extraction between 1993 and 1995 at borrow sites in Denmark, The Netherlands, and Spain. Recolonization of disturbed areas was fast owing to the rapid increase of opportunistic species. At the North Sea sites, the benthic community largely recovered within 2–4 years, whereas in Spain recovery is expected to take longer. The response of zoobenthos to sand extraction is discussed, taking into account differences in site characteristics, extraction methods, and recovery time of the habitats. The effects on the benthic community appear to be related to the physical impact on the sea floor. Small-scale disturbances in seabed morphology and sediment composition result in short-term effects on the benthic community. However, larger disturbances mainly caused by sediment composition may have a prolonged effect, particularly in low dynamic systems such as those present in the Mediterranean.

Qualitative historical benthos data (1902-1912) were compared with recent data (1986) to find long-term trends in epifauna species composition in the southern North Sea that may be attributed to fishery-induced changes. In general, the frequency of occurrence of bivalve species declined, whereas scavenger and predator species (crustaceans, gastropods, and sea stars) were observed more frequently in 1986. We suggest that these shifts can be attributed not only to the physical fishery impact, but also to the additional potential food for scavenging and predator species provided by the large amounts of discards and moribund benthos. Our findings are put into the perspective of the general development of the demersal fishery in the southern North Sea. Despite the problems with the historical data set, the comparison presented may be the best illustration achievable of the changes in the benthos from a near-pristine situation to the present conditions after long-term disturbance. (C) 2000 International Council for the Exploration of the Sea.

This investigation of the bottom fauna of St Austell and Mevagissey Bays was prompted by the feeling among local fishermen that the increasing amount of china-clay waste being deposited in the bays was adversely affecting the local fishery. Physical pollution on a large scale can harm a fishery in a number of ways. The smothering of shellfish grounds is often to be expected, and where the pollutant contains a high proportion of very fine particles there is the danger that unstable clay banks will build up and constitute a purely mechanical hazard to trawling and potting. In this paper, however, we are concerned only with the effects of clay-waste pollution upon the bottom fauna, since it had been suggested that clay deposition might be impairing St Austell and Mevagissey Bays as feeding grounds for demersal fish. Previous work on the biological effects of china-clay waste includes that of Miss N. Sproston (1945, unpublished), who examined the effects of suspended clay upon some species of pelagic fish. This worker was unable to demonstrate any recognizable effect of mica particles (present in clay waste) upon the gill tissues of fish taken from polluted water at Pentewan. However, Herbert et al. (1961) recorded cases of gill damage in Salmo trutta from china-clay-polluted reaches of the River Fal. These workers also showed that ‘the bottom fauna in the control streams was on average 3.3 times greater than in the polluted part of the Fal and 19 times greater than in the polluted part of the Par’.experiments. The benthos dredge Triple-D was used to sample megafauna (>1 cm), while macrofauna (>1 mm) were sampled by means of a Reineck boxcorer and, in some cases, a van Veen grab. Direct mortalities ranging from about 5 up to 40% of the initial densities were observed for a number of gastropods, starfishes, small and medium-sized crustaceans, and annelid worms. For bivalve species, direct mortalities were found from about 20 up to 65%. Mortality per m(2) trawled area due to fishing with a 12-m beam trawl was not higher than that due to a 4-m beam trawl. For all species considered, the direct mortality was largely attributed to animals that died in the trawl track, either as a direct result of physical damage inflicted by the passage of the trawl or indirectly owing to disturbance, exposure, and subsequent predation. In 1994, the 12 m beam trawl with tickler chains was the dominant gear type in the Dutch sector, resulting in a mean annual trawling Frequency of 1.23. The mean annual trawling frequencies with the 4 m beam trawl using tickler chains, the 4 m beam trawl with a chain mat, and the otter trawl were 0.13, 0.01, and 0.06, respectively. The annual fishing mortality in invertebrate megafaunal populations in the Dutch sector ranged From 5 up to 39%, with half of the species showing values of more than 20%. For all species studied, the 12 m beam-trawl fisheries caused higher annual fishing mortalities than the concerted action of the other fisheries. Only with respect to species restricted to sandy coastal areas did the 4 m beam-trawl fleet contribute substantially to the annual mortality. Implications of the impact of trawling on the composition of benthic The Culbin Sands lagoon ecosystem in NE Scotland was studied during a three-year period (1994-1996) to identify the major trophic links from benthic invertebrates to epibenthic predators, and to assess impacts of overwintering fish on their prey communities. Every 2-4 weeks, samples of mobile fauna were collected to study their diets. The major trophic links identified between benthic invertebrates and epibenthic predators were from benthic invertebrates to the shrimp Crangon crangon, and to the common goby Pomatoschistus microps and the plaice Pleuronectes platessa. The energy flow from benthic invertebrates to overwintering fish was estimated at 133 kJ m(-2)yr(-1). A flow of 10 kJ m(-2)yr(-1) was also observed from eggs and larval stages of the overwintering shrimp Crangon crangon to the overwintering fish. Nevertheless, manipulative field experiments showed no significant impacts of the most abundant overwintering fish Pomatoschistus microps on prey community densities, despite an overall individual ingestion rate of 89 J day(-1).

The tube-building polychaete Lanice conchilega can form dense populations, often called reefs, which promote benthic community change and constitute feeding grounds for secondary consumers. The aim of this study was to quantify the role of the L. conchilega reef of the Bay of the Mont Saint-Michel (BMSM) for feeding waders, by combining macrobenthos data, bird counts and bird diet information. Wader densities in the reef were on average 46.6 times higher than in non-reef areas. According to faecal analyses, waders in the reef mainly selected the accompanying fauna and especially crustaceans. The attractiveness of the reef to feeding birds may be largely explained by the high abundance, richness and biomass of macrobenthic species in the reef compared with the rest of the BMSM.

The diets of gurnards Aspitrigla cuculus and Eutrigla gurnardus, lesser-spotted dogfish Scyliorhinus canicula and whiting Merlangius merlangus were examined to determine whether they migrated into recently trawled areas to feed on animals that may be damaged or dislodged by the action of a 4 m beam trawl. Gurnards and whiting increased their intake of prey after an area had been fished. In particular, they increased the proportion of the amphipod Ampelisca spinipes in their diets. Beam trawling damaged the purple burrowing heart urchin Spatangus purpureus, scallop Aequipecten opercularis, Ensis spp. and Laevocardium sp., exposing internal tissues which were then eaten by whiting. Some mobile invertebrate scavengers, such as Pandalus spp., only occurred in diets after the area had been fished, suggesting that these animals were also scavenging over the trawl tracks. Observations of the seabed using a side-scan sonar revealed a greater concentration of fish marks around the trawl tracks than in adjacent unfished areas. Our results indicate that fish rapidly migrate into beam trawled areas to feed on benthic animals which have been either damaged or disturbed by fishing or on scavenging invertebrates. In areas where certain benthic communities occur, beam trawling intensity may be such that it creates a significant food resource for opportunistic fish species. This is a possible mechanism whereby long-term community structure could be altered by fishing activity.

The contribution of infauna and mussel-raft epifauna to the diet of 3 dominant species in the demersal fish community of the Ria de Arosa, N. W. Spain - Lesueurigobius friesji (Gobiidae), Callionymus lyra (Callionymidae) and Tn.sopterus luscus (Gadidae) - was determined. Intense raft mussel culture in the Ria de Arosa supports a rich epifauna which constitutes the main food resource for the fishes studied. In contrast, infauna density is low and contributes only a small proportion to fish diets. Prey consumed was similar in the 3 fish species. Gut contents consisted mainly of the small crab Pisjdja longicomis. This decapod is a dominant component of the raft epifauna, and electivity indices indicate that it is selected by the fishes. In the Ria de Pontevedra, which contains fewer mussel rafts, these flsh fed on infauna. Thus, one effect of intense mussel aquaculture has been to change the food habits of these 3 fishes from predominantly infauna to raft epifauna dietdistribution and abundance of species. Initially, there will not be a wholesale movement northwards of southern species or retreat northwards of northern species, because many additional factors will influence the responses of the different organisms. Such factors include the hydrodynamic characteristics of water masses, the presence of hydrographical and geographical barriers to spread and the life history characteristics (reproductive mode, dispersal capability and longevity) of species. Survey data over the past century show how organisms react to changes of the order of 0.58C, and in the last two decades, when sea temperatures have risen by as much as 18C, there have been significant local changes in the distribution of intertidal organisms. These past changes provide a clue to more extensive changes expected in the future if global warming develops as predicted. Where species affected by climate change are dominant or key structural or functional species in biotopes, there may be a change in the extent and distribution of those biotopes. Some, dominated by predominantly northern species such as the horse mussel Modiolus modiolus, may decline and reduce their value as rich habitats for marine life. Others, characterized by southern species, for example the sea fan Eunicella verrucosa and the alcyonacean Alcyonium glomeratum, may increase in extent. Using information on the life history characteristics of species, their present distribution and other factors, a key supported by a decision tree has been constructed to identify ‘types’ of organism according to their likely response to temperature rise. Conspicuous and easily identified rocky substratum species are temperature has been generally higher in northern than in southern European seas, and higher in enclosed than in open seas. Although European marine ecosystems are influenced by many other factors, such as nutrient enrichment and overfishing, every region has shown at least some changes that were most likely attributable to recent climate change. It is expected that within open systems there will generally be (further) northward movement of species, leading to a switch from polar to more temperate species in the northern seas such as the Arctic, Barents Sea and the Nordic Seas, and subtropical species moving northward to temperate regions such as the Iberian upwelling margin. For seas that are highly influenced by river runoff, such as the Baltic Sea, an increase in freshwater due to enhanced rainfall will lead to a shift from marine to more brackish and even freshwater species. If semi-enclosed systems such as the Mediterranean and the Black Sea lose their endemic species, the associated niches will probably be filled by species originating from adjacent waters and, possibly, with species transported from one region to another via ballast water and the Suez Canal. A better understanding of potential climate change impacts (scenarios) at both regional and local levels, the development of improved methods to quantify the uncertainty of climate change projections, the construction of usable climate change indicators, and an improvement of the interface between science and policy formulation in terms of risk assessment will be The variation in growth rate with temperature of unicellular algae suggests that an equation can be written to describe the maximum expected growth rate for temperatures less than 40 degrees centigrade. Measured rates of phytoplankton growth in the sea and in lakes are reviewed and compared with maximum expected rates. The assimilation number (i.e. rate of photosynthetic carbon assimilation per weight of chlorophyll a ration in the phytoplankton). Since maximum expected growth rate can be estimated from temperature, the maximum expected assimilation number can also be estimated if the carbon/chlorophyll a ratio in the phytoplankton crop is known. Many investigations of phytoplankton photosynthesis in the ocean have included measures of the assimilation number, while fewer data are available on growth rate. Assimilation numbers for Antarctic seas are low as would be expected from the low abmient temperatures. Tropical seas and temperate waters in summer often show low assimilation numbers as a result of low ambient nutrient concentrations. However, coastal estuaries with rapid nutrient regeneration processes show seasonal variations in the assimilation number with temperature which agree well with expectation. The variation in maximum expected growth rate with temperature seems a logical starting point for modelling phytoplankton growth and photosynthesis in the sea. dramatic impacts on biological ecosystems in the upper ocean. Estimates based on the Intergovernmental Panel on Climate Change (IPCC) business-as-usual emission scenarios suggest that atmospheric CO2 levels could approach 800 ppm near the end of the century. Corresponding biogeochemical models for the ocean indicate that surface water pH will drop from a pre-industrial value of about 8.2 to about 7.8 in the IPCC A2 scenario by the end of this century, increasing the ocean's acidity by about 150% relative to the beginning of the industrial era. In contemporary ocean water, elevated CO2 will also cause substantial reductions in surface water carbonate ion concentrations, in terms of either absolute changes or fractional changes relative to pre-industrial levels. For most open-ocean surface waters, aragonite undersaturation occurs when carbonate ion concentrations drop below approximately 66 µmol kg-1. The model projections indicate that aragonite undersaturation will start to occur by about 2020 in the Arctic Ocean and 2050 in the Southern Ocean. By 2050, all of the Arctic will be undersaturated with respect to aragonite, and by 2095, all of the Southern Ocean and parts of the North Pacific will be undersaturated. For calcite, undersaturation occurs when carbonate ion concentration drops below 42 µmol kg-1. By 2095, most of the Arctic and some parts of the Bering and Chukchi seas will be undersaturated with respect to calcite. However, in most of the other ocean basins, the surface waters will still be saturated

processes of the Baltic tellin (Macoma balthica), a widely distributed bivalve that plays a critical role in the functioning of many coastal habitats. We demonstrate that ocean acidification significantly depresses fertilization, embryogenesis, larval development and survival during the pelagic phase. Fertilization and the formation of a D-shaped shell during embryogenesis were severely diminished: successful fertilization was reduced by 11% at a 0.6 pH unit decrease from present (pH 8.1) conditions, while hatching success was depressed by 34 and 87%, respectively at a 0.3 and 0.6 pH unit decrease. Under acidified conditions, larvae were still able to develop a shell during the post-embryonic phase, but higher larval mortality rates indicate that fewer larvae may metamorphose and settle in an acidified ocean. The cumulative impact of decreasing seawater pH on fertilization, embryogenesis and survival to the benthic stage is estimated to reduce the number of competent settlers by 38% for a 0.3 pH unit decrease, and by 89% for a 0.6 pH unit decrease from present conditions. Additionally, slower growth rates and a delayed metamorphosis at a smaller size were indicative for larvae developed under acidified conditions. This may further decline the recruit population size due to a longer subjection to perturbations, such as predation, during the pelagic phase. In general, early life history processes were most severely compromised at similar to pH 7.5, which corresponds to seawater undersaturated with respect to aragonite. Since recent models predict a comparable decrease in pH in coastal waters in the near future, this study indicates that future populations of This paper compares the infaunal and epifaunal assemblages from surveys encompassing 121 grab stations and 152 Agassiz trawl samples respectively, collected between 1980 and 1985. The area surveyed is delimited by the Scottish, Norwegian and Danish coasts lying between 56°15'N and 60°45'N. Samples for infauna and environmental parameters were collected by Smith-Mclntyre grab and Craib corer. The epifaunal and infaunal assemblages were analysed separately by ordination techniques (DECORANA and TWlNSPAN) to detect the major environmental gradients underlying the distribution and abundance of the fauna and to indicate which taxa were characteristic of different zones within the survey area. The major determinant of infaunal community composition was sediment granulometry, with depth being of secondary importance. For the epibenthos, depth was the major factor and the sediment composition seemed less significant. Assemblages identified by TWlNSPAN were characterised by particular species, but these 'community types' were seen to grade into one another along continuous environmental gradients. These findings are discussed in relation to previous North Sea benthic classification schemes.presented. Values for individual sampling stations varied from 0.21 to 4.1 y−1 for community P/B and 3.1 to 897.2 kJm−2 y−1 for total production. Such data fills an important gap pertaining to our understanding of the spatial variation in production estimates for this region. Benthic production estimates varied primarily at small (inter-station) scales (24 nm), although larger-scale differences were observed. In general, the highest production estimates were exhibited by benthic communities in Cardigan Bay (Irish Sea) and East English Channel, while the lowest estimates were observed for the mid- and northern North Sea areas. The former were typified by shallow, gravelly areas of seabed which exhibit high bed tidal stress and do not thermally stratify during the summer months. On average, annelids contribute an overwhelming majority of the total production with different regions varying in the relative contributions from other phyla such as molluscs, crustaceans and echinoderms. Spatial heterogeneity of sediment granulometric variables occurred primarily between stations while those of other variables (e.g., depth, stratification, and tidal bed stress) were more regional. Although a large proportion of the spatial variation in secondary production estimates was not explained by environmental characteristics, the data indicate that such relationships are scale-dependent. Average bed temperature was a significant factor in creating some of the observed differences at large spatial scales. The possible reasons why a larger proportion Using data published in 15 major marine ecology journals (from 1970 to 1999), we examined global patterns of marine benthic macroinvertebrate production and its distribution among feeding guilds and taxonomic groups and physical variables such as substratum type, water depth and temperature. Our database contains 547 production datasets, from 147 studies including 207 taxa, assessed by classical methods (cohort and size-based methods), from 170 sites (77° 50’ S to 69° 35’ N; 0 to 930 m depth). In general, higher values of production to biomass (P/B) ratios were observed in the Northern Hemisphere than in the Southern Hemisphere. High values of P/B ratios were observed in mid-latitudinal zones while low values of P/B ratios were observed in high (80 to 60° S) and low latitudinal zones (40° S to 20° N). Highest production was observed on hard substrata, for filter feeders and for mollusc (e.g. bivalves) species. Highest P/B ratios were observed on algae (or high organic substrata), omnivores and predators, and arthropods (e.g. amphipods). Regression models explained a significant percentage of the amount of variance of benthic production (92%) and P/B ratios (50 to 86%). Production and P/B ratios were negatively related to water depth and positively related to water temperature, but these abiotic variables did not greatly improve the predictability of production by biotic variables (e.g. life span, mean body mass). Biotic variables were more important than environmental variables in explaining observed variations in production and P/B ratios. For the latter, life span explained most (45 to 83%) of the variation of the models.

mixing and salinity. There was a strong statistically significant linear relationship between SPM and Kd for the full data set. We show that slightly better results are obtained by fitting separate models to data from transitional waters and coastal and offshore waters combined. These linear models were used to predict Kd from SPM. Using a statistic (D) to quantify the error of prediction of Kd from SPM, we found an overall prediction error rate of 23.1%. Statistically significant linear relationships were also evident between the log of Secchi depth and the log of Kd in waters around the UK. Again, statistically significant improvements were obtained by fitting separate models to estuarine and combined coastal/offshore data – however, the prediction error was improved only marginally, from 31.6% to 29.7%. Prediction was poor in transitional waters (D ¼ 39.5%) but relatively good in coastal/offshore waters (D ¼ 26.9%). SPM data were extracted from long term monitoring data sites held by the UK Environment Agency. The appropriate linear models (estuarine or combined coastal/offshore) were applied to the SPM data to obtain representative Kd values from estuarine, coastal and offshore sites. Estuarine waters typically had higher concentrations of SPM (8.2–73.8 mg l 1) compared to coastal waters (3.0–24.1 mg l 1) and offshore waters (9.3 mg l 1). The higher SPM values in estuarine waters corresponded to higher values of Kd (0.8–5.6 m 1). Water types that were identified by large tidal ranges and exposure typically had the highest Kd ranges

ecological importanCe to coastal marine ecosystems that has changed so drastically in such a shon period as dissolved. oxygen. While hypoxic and anoxic environments have existed through geological time, !heir occum:oce in shallow coastal and estuarine areas appears to be ineuuiog. moSt likely accelerat-::d by human activilies. Ecological problems associated with the occurrence of low oxygen au increasing on a global scale. The oxygen budgets ofrnost major eSfllarine and coastal ecosystems have been adversely affected mainly through !he procw of eutrophication. whichaclS as an acceleranl or enhancing factor to hypoxia and anoxia, and when coupled with adverse meteorologicaland. hydrodynamic events. hypoxia ioeuas« in frequency and severity. The area of hypo:dc and anoxic bonom water is even increasing within systems thaI historically are considered oltygen stressed. Many ecosystems that are II()W seveuly slfesscd by hypoltla appear to be near or al a threshold. Should oxygen concentTlitions become sJighfly lower, catastrophic events may oven:ome the systems tl1d alter !he productivity base that leads to fiSheries spo::ci<:::s. fuamples of such events are ~com;nZ increasinll:ly com· mono At wltat poim permanent damage will result is difficult to say. To date there is II() large system that Itas recovered after development ofpcrsistent hypoll;la or anoxia. The only exception may be small systems wltere pollution inputs have ceased and recovery initiated from surrounding non-affected aTeas. The eltpanding occurunce of Itypoxia and tl10ltia continues to

Previous studies of marine soft-bottom communities have shown (1) that natural disturbances (especially biologically-mediated disturbances, which are usually localized and recur reasonably frequently) help maintain spatio-temporal heterogeneity of communities, and (2) that biogenic modification of sediment can affect sediment stability with respect to fluid forces and geotechnical properties and that this is an important factor in community organization, particularly in the trophic structure of the macrofauna. It is argued in this paper that natural disturbances, and the ensuing biogenic alterations to sediment stability, may be important in maintaining trophically-mixed communities where deposit feeders do not have an overriding influence on sedimentary properties. The hypothesis is presented that an initial post-disturbance response by micro- and meiobenthos leads to an increase in sediment stability as a result of mucous-binding of sediment, and that this stage may be of critical significance to potential suspension-feeding colonists if they are competing with deposit feeders for space. It is suggested, partly as a corollary to this hypothesis, that there may be marked differences in the structure and function of meiofaunal communities co-occurring with deposit-feeding and suspension-feeding macrofaunas. Implications for macrofaunal trophic structure of seasonal changes in sediment stability are also examined. Several areas for future research are recommended.

steps, and current velocities, turbulent kinetic energy (TKE), and suspended sediment concentration (SSC) were measured. Current speeds decreased and turbulence increased within 3 cm of the sediment surface as a function of cockle density. Sediment resuspension increased with increasing cockle density: SSC at 0.5 m s− 1 = 156, 574, 1045 and 2253 mg L− 1 for 0, 47, 141 and 312 animals m− 2 respectively. Enhanced sediment erosion was due to increased bioturbation and bed roughness. Critical erosion velocity decreased from 26 to 8 cm s− 1 with increasing cockle density. Shear stress, measured in terms of TKE, increased at 0.5 and 1 cm above the bed for 141 and 312 animals m− 2 and up to 2 cm above the bed for 312 animals m− 2, reflecting the increased bed roughness due to bioturbation. However, the critical erosion shear stress was relatively independent of cockle density (0.225 and 0.151 N m− 2 for 0 and 312 animals m− 2 respectively). The valve adduction frequency of the cockles was also measured in response to increasing SSC. The frequency increased from 1.2 to 16.3 adductions h− 1 with increasing SSC from 13 to 308 mg L− 1. It represents a significant behavioural response, creating positive feedback with increased SSC further enhancing sediment disturbance and resuspension.

Biogenic forces alter sediment characteristics along several axes with important consequences for structure of benthic communities. The usual axes discussed are those of sediment stabilization versus resuspension and mobile versus temporally persistent organisms. A third axis of bioadvection is typically subsumed within the others. Here we argue that given the complex fluid dynamics resulting from the bidirectional forces that organisms exert on porewater, bioadvection needs to be examined separately. The probable major players in generation of bioadvection are described with impacts on transport both of materials and heat. Illustrations are given of the bidirectionality of bioadvection and the resultant changes in oxygenation either surficially or at depth, as well as of heat transport both laterally within the sediment and vertically.

O2 plays a key role in early sedimentary diagenetic processes, but the effect of most macrofaunal species on the pathways and rates of supply of O2 into the seabed are not well known. We investigated the effect of the ophiuroid Amphiura filiformis,one of the dominant macrobenthic species on soft bottoms in the northeast Atlantic, at depths of ;15-100 m, in a laboratory environment. We determined how the presence of the ophiuroid changed the total O2 uptake of macrofauna-free sediment by combining measurements from a microcosm approach and an approach that uses microelectrodes and a flushed aquarium. We suggest that natural populations of A. filiformiscan account for 80% of the total flux of O 2 into the soft bottom. At least 67% of this portion is due to the diffusion of O 2 across additional sediment-water interfaces excavated by the brittle star.investigated to estimate the effects of density declines and species loss on benthic ecosystem functioning. Two laboratory experiments were performed: before (winter, temperature = 10°C) and after (summer, temperature = 18°C) sedimentation of the spring phytoplankton bloom. Single species treatments of key species (Abra alba, Lanice conchilega and Nephtys sp.) with different functional traits were added to microcosms at 3 density levels (natural, lower, lowest) to account for possible density declines. Sediment–water exchanges of oxygen and nutrients, denitrification and bioturbation were measured. In absence of fauna, benthic mineralisation in the summer experiment was 2.0 times higher than in winter. Fauna stimulated microbial respiration more in summer (up to 100% in L. conchilega treatments) than in winter (negligible fauna effect). As chlorophyll a concentrations were similar in both seasons, the stronger fluxes in summer must be explained by a higher macrobenthic activity owing to the elevated temperature and better condition of the animals. Stimulation of mineralisation by the 3 species in the microcosms was different, and behaviour-related. Owing to its irrigation activity, the tube dweller L. conchilega had more pronounced influences on benthic respiration, nutrient release and denitrification than did the biodiffusers, A. alba and Nephtys sp. A. alba appeared to be a more effective bioturbator than Nephtys sp. Processes such as benthic respiration, nutrient fluxes, denitrification and bioturbation seem to

The effects of increased sedimentation on the macrobenthic community, physical structure, and biogeochemistry of the surficial sediment around two farms in southwest Ireland were examined in conjunction with current characteristics. Both farms had been in production for over eight years, were of reasonably large size (>100 MT) and located in low-energy environments. At one site, the benthic community was subjected to bulk sedimentation and organic enrichment and reduced macrobenthic infaunal diversity and elevated levels of organic carbon were recorded close to the farm. In general, effects were restricted to a radius of 40 m around the farm. Conversely, at the second site, there were no observed effects of mussel biodeposits on the benthos and a diverse macrobenthic community persisted. We propose that variations in the dispersion of biodeposits caused by local current patterns had a significant influence on the impact observed, and that this could also account for differences reported in other studies.

contributing to the biological functions of all organisms. Because biologically available N often limits marine productivity, microbial processes leading to its loss and gain (e.g. denitrification and N2 fixation, respectively) play an important role in global biogeochemical cycles. Bioturbation is known to influence benthic N cycling, most often reported as enhancement of denitrification and a subsequent loss of N2 from the system. N2 fixation has rarely been addressed in bioturbation studies. Instead, sedimentary N2 fixation typically has been considered important in relatively rare, localized habitats such as rhizosphere and phototrophic microbial mat environments. However, the potential for N2 fixation in marine sediments may be more widespread. We show here that nitrogenase activity can be very high (up to 5 nmol C2H4 cm–3 h–1) in coastal sediments bioturbated by the ghost shrimp Neotrypaea californiensis and at depths below 5 cm. Integrated subsurface N2-fixation rates were greater than those previously found for un-vegetated estuarine sediments and were comparable to rates from photosynthetic microbial mats and rhizospheres. Inhibition experiments and genetic analysis showed that this activity was mainly linked to sulfate reduction. Sulfatereducing bacteria (SRB) are widespread and abundant in marine sediments, with many possessing the genetic capacity to fix N2. Our results show that N2 fixation by SRB in bioturbated sediments may be an important process leading to new N input into marine sediments. Given the ubiquity of bioturbation and of SRB in marine sediments, this overlooked benthic N2 fixation may play an important role in Research so far has provided little evidence that benthic biogeochemical cycling is affected by ocean acidification under realistic climate change scenarios. We measured nutrient exchange and sediment community oxygen consumption (SCOC) rates to estimate nitrification in natural coastal permeable and fine sandy sediments under pre-phytoplankton bloom and bloom conditions. Ocean acidification, as mimicked in the laboratory by a realistic pH decrease of 0.3, significantly reduced SCOC on average by 60% and benthic nitrification rates on average by 94% in both sediment types in February (pre-bloom period), but not in April (bloom period). No changes in macrofauna functional community (density, structural and functional diversity) were observed between ambient and acidified conditions, suggesting that changes in benthic biogeochemical cycling were predominantly mediated by changes in the activity of the microbial community during the short-term incubations (14 days), rather than by changes in engineering effects of bioturbating and bio-irrigating macrofauna. As benthic nitrification makes up the gross of ocean nitrification, a slowdown of this nitrogen cycling pathway in both permeable and fine sediments in winter, could therefore have global impacts on coupled nitrification-denitrification and hence eventually on pelagic nutrient availability.concentrations. Ultrasonic Doppler velocity profiling was used to measure the flow velocity within these concentrated suspension flows. The development of current ripples under decelerated flows of differing kaolin concentration was documented and evolution of their height, wavelength and migration rate quantified. This work confirms past work over smooth, fixed beds which showed that, as clay concentration rises, a distinct sequence of flow types is generated: turbulent flow, turbulence-enhanced transitional flow, lower transitional plug flow, upper transitional plug flow and a quasi-laminar plug flow. Each of these flow types produces an initial flat bed upon rapid flow deceleration, followed by reworking of these deposits through the development of current ripples during the subsequent steady flow in turbulent flow, turbulence-enhanced transitional flow and lower transitional plug flow. The initial flat beds are structureless, but have diagnostic textural properties, caused by differential settling of sand, silt and cohesive mud, which forms characteristic bipartite beds that initially consist of sand overlain by silt or clay. As clay concentration in the formative flow increases, ripples first increase in mean height and wavelength under turbulence-enhanced transitional flow and lower transitional plug-flow regimes, which is attributed to the additional turbulence generated under these flows that subsequently causes greater lee side erosion. As clay concentration increases further from a lower transitional plug flow, ripples cease to exist under the upper transitional plug flow and quasi-laminar plug flow conditions investigated herein. This disappearance of ripples appears due to both

the basic parameters for each benthic compartment (i.e. detritus, bacteria, meiofauna and macrofauna). Our study focuses on the eastern basin of the English Channel and the Southern Bight of the North Sea. Trophic web modelling was used to assess the functioning of the three main benthic community assemblages. To test and assess the relative importance of factors assumed to influence trophic structure (geographical environment and sedimentary particle size distribution), the study area was subdivided into divisions defined a priori according to the two main structural factors of community distribution; geographic distribution and sedimentary patterns. Then, a steady state trophic model utilising the inverse method was applied to a diagram composed of eight compartments, including detritus, bacteria, meiofauna, macrobenthos and fish. For each compartment, six physiological parameters were assessed, based on our own data, empirical relationships and literature data. This method allowed estimation of the flux of matter and energy within and between the units of the benthic system and assessment of the amount of trophic energy stored in these units (available mostly to fish). Our results showed that suspension-feeders control most of the matter transfer through the macrobenthic food-web, except in the fine sand community, where deposit-feeders play a dominant role. The results also showed that, whatever the geographic area, trophic structure is strongly linked to the sedimentary conditions. As benthic communities are dibrunchiata Ehlers, and epibenthic predators in structuring an intertidal soft-bottom community in Maine. Epibenthic predators were excluded from portions of the soft bottom using cages which also enclosed elevated densities of the predatory polychaetes. The experiments ran 10 wk and 20 wk beginning in June 1979. Exclusion of epibenthic predators had no effect on infaunal densities after 10 wk but produced ;2: IJ-fold increase in total density after 20 wk. Since gulls (Lams spp.) avoided all cages, including those not designed to exclude epibenthic predators, the effect of gull predation on infaunal abundances was not tested using exclusion cages. Crabs, Carcinus maenas (Linnaeus) and Cancer irrorutus Say were observed in cages not designed to exclude predators. Densities of Nephtys in&a Malmgren, Poiydora Zigni Webster, Streblospio benedicti Webster, Scoloplos robustus Verrill, phyllodocids, and bivalves were highest in cages containing elevated Glycera dibranchiata density and lowest in cages containing elevated NereL virens density. N. virens was the only taxon whose abundance was reduced in the presenci of Glycera dibranchiata which may account for high infaunal densities in the G. dibranchiata treatment. Laboratory experiments demonstrated that G. dibranchiata are capable of preying on Nereis v&ens. Additional field experiments revealed that the presence of N. virens reduced the abundance of some taxa within the tirst 10 days of colon~ation. N. &ens may have reduced infaunal densities by predation andjor disturbance at the sediment surface. These results suggest that complex interactions within the infauna are important in

Benthic organisms in marine ecosystems modify the environment on different spatial and temporal scales. These modifications, many of which are initially at a microscale, are likely to have large scale effects on benthic seascapes. This is especially so if the species are ecosystem engineers. Most species of infaunal and epifaunal invertebrates and macrophytes contribute at a geophysical or geochemical level. Microorganisms also play a key but currently neglected role. In the intertidal and immediately sublittoral zone, algae and seagrasses, and mussels in mussel beds have received considerable attention. A substantial fossil record also exists. Mathematical modelling of these systems is still in its infancy, although several sophisticated mathematical tools have been applied. The effects of bioturbation and of microorganisms have been less studied, and little is known about the activities of benthic organisms in the deep sea. This paper addresses all these effects, and places them in the context of large scale benthic seascapes and of the extensive literature on species defined as ecosystem engineers in the sea.

Negative relations between abundance of the predatory polychaete Nephtys hombergii and values for biornass and rate of increase in 2 of its prey species, the polychaetes Scoloplos armiger and Heteromastusfiliformis, are evident in long-term (18 yr) data from tidal flats in the westernmost part of the Wadden Sea. Values for prey biornass tended to decline at high N. hombergii biomass (above ca 0.3 g m-' AFDW), whereas they tended to increase at lower levels of N. hombergii biomass. These results corroborate Schubert & Reise's (Mar. Ecol. Prog. Ser. 34: 117-124, 1986) conclusion, based on short-term enclosure experiments, that N. hornbergl~ is an important infaunal predator in the Wadden Sea

There is a paucity of studies showing long-term changes in the population dynamics of dominant benthic epifaunal species, especially echinoderms, in relation to biological and environmental factors. In the English Channel, the brittle-star Ophiothrix fragilis is a common epifaunal species, mainly found in strong tidal currents characterised by benthic habitats with pebbles. However, in the Bay of Seine, O. fragilis lives on gravel and coarse sandy sediments; more locally, it occurs where there are unexpected amounts of fine particles for such high hydrodynamic areas. This species forms dense aggregations, supporting large populations up to 7450 ind m_2. This paper analyses the long-term spatio-temporal changes of O. Fragilis aggregations over the last 25 years in the eastern part of the Bay of Seine through observations obtained from several scientific programmes from 1986 to 2010. This area is characterised as a tidal environment affected by the Seine estuary and is subject to potential sediment supply from the dumping site of the Le Havre harbour dredging operations. During all surveys, there was a similar pattern: persistent patches with high abundances of O. fragilis and sites without O. fragilis, showing that there was a high heterogeneity of the spatial population pattern. Interactions between environmental conditions and ophiurid aggregations (e.g., storm waves, Seine floods and patches) are suggested to explain these patterns.Background: Bioturbators affect multiple biogeochemical interactions and have been suggested as suitable candidates to mitigate organic matter loading in marine sediments. However, predicting the effects of bioturbators at an ecosystem level can be difficult due to their complex positive and negative interactions with the microbial community. Methodology/Principal Findings: We quantified the effects of deposit-feeding sea cucumbers on benthic algal biomass (microphytobenthos, MPB), bacterial abundance, and the sediment-seawater exchange of dissolved oxygen and nutrients. The sea cucumbers increased the efflux of inorganic nitrogen (ammonium, NH4+) from organically enriched sediments, which stimulated algal productivity. Grazing by the sea cucumbers on MPB (evidenced by pheopigments), however, caused a net negative effect on primary producer biomass and total oxygen production. Further, there was an increased abundance of bacteria in sediment with sea cucumbers, suggesting facilitation. The sea cucumbers increased the ratio of oxygen consumption to production in surface sediment by shifting the microbial balance from producers to decomposers. This shift explains the increased efflux of inorganic nitrogen and concordant reduction in organic matter content in sediment with bioturbators. Conclusions/Significance: Our study demonstrates the functional role and potential of sea cucumbers to ameliorate some of the adverse effects of organic matter enrichment in coastal ecosystems.Predicting the consequences of species loss is critically important, given present threats to biological diversity such as habitat destruction, overharvesting and climate change(1). Several empirical studies have reported decreased ecosystem performance ( for example, primary productivity) coincident with decreased biodiversity(2-4), although the relative influence of biotic effects and confounding abiotic factors has been vigorously debated(5-7). Whereas several investigations focused on single trophic levels ( for example, grassland plants)(8,9), studies of whole systems have revealed multiple layers of feedbacks, hidden drivers and emergent properties(10,11), making the consequences of species loss more difficult to predict(12). Here we report functionally important organisms and considerable biocomplexity in a sedimentary seafloor habitat, one of Earth's most widespread ecosystems. Experimental field measurements demonstrate how the abundance of spatangoid urchins - infaunal ( in seafloor sediment) grazers / deposit feeders - is positively related to primary production, as their activities change nutrient fluxes and improve conditions for production by microphytobenthos (sedimentatry microbes and unicellular algae). Declines of spatangoid urchins after trawling are well documented(13,14), and our research linking these bioturbators to important benthic - pelagic fluxes highlights potential ramifications for productivity in coastal oceans.

In the literature, 69 references altogether have reported 76 predators on holothurians. In terms of the number of predatory species, the most important predators are fishes (26 species), seastars (19 species), and crustaceans (17 species). Seastars are the predators most often cited as regularly ingesting large quantities of holothurians. Toxicity seems to be an effective defense against a generalized predator but, against a specialist on holothurians, escape by swimming movements or shedding of a piece of body wall are the only behaviors that occasionally end with a successful escape. Escape behaviors may be a factor in providing the apparent size refuge from predators. Impacts of predators on holothurian populations have rarely been reported or evaluated, and predation on the earliest life stage is unknown.

also affect the spatial distribution of these smaller organisms. In a controlled laboratory experiment, also affect the spatial distribution of these smaller organisms. In a controlled laboratory experiment, we studied the effects of 3 functionally different macrobenthic species on the vertical distribution of nematodes. Abra alba, a suspension-deposit feeding bivalve reworking the sediment randomly, Lanice conchilega, a suspension-deposit feeding, tube-irrigating polychaete and Nephtys hombergii, a burrowing predatory polychaete, were added in single-species treatments to sediment from a coastal subtidal station in the Belgian part of the North Sea, sieved (1 mm) to remove macrofauna. After 14 d, the control treatment without macrobenthos was found to be detrimental to nematode density and diversity, which points to the importance of macrobenthic engineering to sustain the smaller components of the food web. Nematode densities were highest at the sediment surface in all treatments, but subsurface density peaks were observed in A. alba (to 3 cm depth) and L. conchilega (to 7 cm depth) microcosms. In the A. alba treatment, the dominant non-selective deposit feeders and the epistrate feeders shifted downwards probably to avoid disturbance and exploitative competition by the bivalve siphons at the surface, while they might have benefited from the faecal pellets deposited in the subsurface. In the L. conchilega treatment, the several dominant species were redistributed over depth layers, indicating polychaete-mediated habitat extension from surface into depth. Nematode communities seemed hardly affected by the presence of

A microcosm experiment was conducted to investigate the effects of continuous and episodic biological disturbance by Carcinus maenas on estuarine nematode assemblages from sand and mud for a period of 57 days. Univariate methods of data evaluation failed to reveal major changes in community structure. Distributional techniques (dominance curves) were more sensitive in detecting changes in diversity patterns in the sand assemblage. Results of multivariate analyses indicated that nematode assemblages had changed characteristically due to biological disturbance. The observed changes in nematode community stucture were the result of confounded effects of predation and modification of the sediment due to crab activity. Nematode assemblages from the organic-poor sand were mainly affected by crab predation, those from the organic-rich mud were mainly affected by changes to the sediment due to crab feeding activity. Effects of biological disturbance on both nematode assemblages were dependent on the frequency of disturbance events.bottom in 24 m depth in the Gulf of Trieste. This approach successfully simulates the seasonal low dissolved oxygen (DO) events here and enabled studying the behaviour and mortality of the hermit crab Paguristes eremita. The crabs exhibited a sequence of predictable stress responses and ultimately mortality, which was correlated with five oxygen thresholds. Among the crustaceans, which are a sensitive group to oxygen depletion, P. eremita is relatively tolerant. Initially, at mild hypoxia (2.0 to 1.0 ml l−1 DO), hermit crabs showed avoidance by moving onto better oxygenated, elevated substrata. This was accompanied by a series of responses including decreased locomotory activity, increased body movements and extension from the shell. During a moribund phase at severe hypoxia (0.5 to 0.01 ml l−1 DO), crabs were mostly immobile in overturned shells and body movements decreased. Anoxia triggered emergence from the shell, with a brief locomotion spurt of shell-less crabs. The activity pattern of normally day-active crabs was altered during hypoxia and anoxia. Atypical interspecific interactions occurred: the crab Pisidia longimana increasingly aggregated on hermit crab shells, and a hermit crab used the emerged infaunal sea urchin Schizaster canaliferus as an elevated substrate. Response patterns varied somewhat according to shell size or symbiont type (the sponge Suberites domuncula). Mortality occurred after extended anoxia (~1.5 d) and increased hydrogen sulphide levels (H2S ~128 μmol). The Published and unpublished records of amphipod-sea anemone associations are reviewed. They involve at least 22 amphipod species in 7 families, and 8 families of sea anemones. The associations are of 4 main types: protection only, ectocommensals, endocommensals and micropredators. Morphological adaptations are not conspicuous, except for the specialised mouthparts of Acidostoma spp., but most obligate symbionts show inborn immunity against the toxic substances released by the host. Sex ratios are normal, sexual dimorphism small, and fecundity low compared to related free-living species. The obligate commensal associates are usually host-specific, although able to survive in alternative hosts in the laboratory, while the micropredators and the facultative associates show low host specificity. The amphipod symbionts usually do not occupy the entire geographical and ecological range of their hosts' distribution. Amphipod-sea anemone associations have evolved independently many times and do not seem to be of great evolutionary antiquity.

Sea anemones (order Actiniaria) are among the most diverse and successful members of the anthozoan subclass Hexacorallia, being found at all depths and latitudes and in all marine habitats. Members of this group exhibit the greatest variation in anatomy, biology, and life history in Hexacorallia, and lack any morphological synapomorphy. Nonetheless, previous molecular phylogenetic studies have found that Actiniaria is monophyletic with respect to other extant hexacorallians. However, relationships within Actiniaria have remained unresolved, as none of these earlier works have included sufficient taxon sampling to estimate relationships within Actiniaria. We have analyzed sequences from two mitochondrial and two nuclear markers for representatives of approximately half of the family-level diversity within the order, and present the first phylogenetic tree for Actiniaria. We concur with previous studies that have suggested that molecular evolution is unusually slow in this group. We determine that taxonomic groups based on the absence of features tend not to be recovered as monophyletic, but that at least some classical anatomical features define monophyletic groups.

hypercapnia suppresses metabolism. However, this may be buffered by enhanced growth and metabolism due to warming. Methodology/Principal Findings: We examined the interactive effects of near-future ocean warming and increased acidification/P-CO2 on larval development in the tropical sea urchin Tripneustes gratilla. Larvae were reared in multifactorial experiments in flow-through conditions in all combinations of three temperature and three pH/P-CO2 treatments. Experiments were placed in the setting of projected near future conditions for SE Australia, a global change hot spot. Increased acidity/P-CO2 and decreased carbonate mineral saturation significantly reduced larval growth resulting in decreased skeletal length. Increased temperature (+3 degrees C) stimulated growth, producing significantly bigger larvae across all pH/P-CO2 treatments up to a thermal threshold (+6 degrees C). Increased acidity (-0.3-0.5 pH units) and hypercapnia significantly reduced larval calcification. A +3 degrees C warming diminished the negative effects of acidification and hypercapnia on larval growth. Conclusions and Significance: This study of the effects of ocean warming and CO2 driven acidification on development and calcification of marine invertebrate larvae reared in experimental conditions from the outset of development (fertilization) shows the positive and negative effects of these stressors. In simultaneous exposure to stressors the dwarfing effects of acidification were dominant. Reduction in size of sea urchin larvae in a high P-CO2 ocean would likely impair their performance with negative consequent effects for benthic adult populations.

Using aquaria containing natural sediment, medium-term burrow development was investigated. After initial construction, and over the first month, relatively little burrow development was observed in terms of the number of openings and tunnels created. After an additional six months, however, there was a significant increase in the number of openings and tunnels constructed. In contrast to what might be expected from the number of openings and tunnels created during this period, a significant proportion of the sediment movement was sub-surface and relatively little sediment expulsion occurred. The concentration of nitrite, ammonia and phosphate in the burrow water was generally greater than that of the surface water, whilst the nitrate and sulphate measurements showed no particular pattern. Only the differences in phosphate concentrations were found to be significantly different. These results were indicative of Calocaris macandreae influencing rates of denitrification. This study also provided further evidence of carnivory and caching behaviour in this species.Many bivalves species inhabit coastal waters where fluctuations in both quantity and quality of suspended particulate matter occur. The study of interactions between the organism and its environment requires thus a certain level of detail concerning the feeding process, not only from the bivalve point of view - which material can they actually use as food - but also from the ecosystem point of view - to what extent are bivalves able to clear the water column and change ecosystem dynamics? However such detail is commonly neglected in ecosystem modelling and a mechanistic description of the feeding process is still lacking. In this study, the Synthesizing Units concept, part of the Dynamic Energy Budget (DEB) theory, is used to describe the main feeding processes in bivalves. Filtration, ingestion and assimilation are assumed as three different steps and pseudofaeces production computed as the difference between filtered and ingested fluxes. Generic formulations are proposed and discussed, considering several types of food, with type-specific ingestion and assimilation efficiencies. The model performance is evaluated by comparison with the literature data for the blue mussel for a wide range of experimental conditions. The lack of data and of detailed information on the experimental setups adds some uncertainty to the parameters estimation. Nevertheless, the model results are in good agreement with observations. The model has the desired flexibility to be implemented as an extension to the standard DEB model, to simulate bivalve growth in estuaries and coastal areas where the organisms experience different food quantity and quality.

over large areas, and it is not known whether single tubes also affect the surrounding fauna. In this study a low density population of L. conchilega (15.7 +/- 15.6 m(-2)) was investigated on an exposed beach in South Wales, UK, from May 1998 to April 1999. Effects of single tubes and small groups of 2 to 5 tubes on the benthic community were examined over 1 yr. The relationship between L, conchilega and an associated amphipod (Urothoe poseidonis) was studied more closely in the field and the laboratory. Of a total of 56 species, 27 were found exclusively in samples with L. conchilega tubes. In comparison with tube-free samples, species richness and abundance of individuals was significantly higher in samples containing L. conchilega tubes. The community structure differed significantly between samples containing groups of tubes and tube-free samples in 10 out of 11 cases and in 9 of 11 cases for samples with 1 tube compared to samples with no tubes. Throughout the year, the polychaete Eumida sanguinea and the haustoriid amphipod U. poseidonis benefited from the presence of L. conchilega. E. sanguinea lives among the fringe filaments of the tube top, and U. poseidonis inhabits areas deep in the sediment in close vicinity to the tube. Laboratory experiments indicated that, unlike other haustoriids, the amphipod is not prone to benthopelagic migration but remains in the sediment for long periods of time and may benefit from an improved oxygen supply arising from L. conchilega's activity inside the tube. It is concluded that not only groups of tubes, but also single polychaete tubes bioengineer their environment.

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 m2) 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 Habitat modifying organisms can alter the distribution of associated species. We surveyed soft-sediment patches in Bodega Harbor, California and found that patches with high densities of the phoronid Phoronopsis harmeri (Pixell, 1912), a chemically-defended tube-building lophophorate, have higher infaunal abundance and richness than similar patches with low densities of P. harmeri. To determine whether this difference was driven by P. harmeri and whether this difference is attributable to the activities of the organism, or simply its physical structure, we conducted a field experiment with four treatments: live phoronids, mimics of phoronid structure, phoronid-free sediments (bare) and unmanipulated sediments. Although the field experiment did not detect differences in the overall abundance or richness of infauna among the manipulated treatments, some of the individual species did show a positive response to the presence of phoronids and phoronid structure (i.e., mimics). In particular, the polychaete Boccardia proboscidea, the amphipod Monocorophium uenoi, and harpacticoid copepods were facilitated by the presence of phoronids and phoronid structure when there was sediment disturbance. The inconsistency between the results of the survey and of the manipulative experiment may be largely driven by the disturbance caused by the manipulation. However, where P. harmeri has an effect, it is generally positively associated with infaunal abundance that may be attributable to the stabilization of sediments.intertidal habitat complexity, modifying water flow, promoting sediment accretion and affecting nutrient fluxes at the water–sediment interface. Understanding how such structures affect the benthic ecosystem’s functioning requires the assessment of their influence on all benthic components and how the related ecosystem services may be modified. We performed an in situ experimental study, involving the use of artificial mimics of polychaete tubes, to investigate the purely physical impacts of the structures without the complexity of worm activity. Benthic chambers of different mimic densities were used, and their effect on the recolonisation of defaunated natural sandy sediments by microorganisms, meiofauna and macrofauna was monitored. We also measured air–sediment CO2 fluxes and sediment stability as they constitute crucial ecosystem services provided by benthic habitats. We showed that the biogenic structures stimulated the development of diatom biofilms (microphytobenthos) and their associated extracellular polymeric substances (EPS). Impacts of tubes on meiofaunal and macrofaunal assemblages were significant; in most cases, species and groups were more abundant in treatments with few or no tubes. In response to the tube density increase, the whole system tended towards heterotrophy and higher sediment stability, probably as a consequence of the development of the diatom biofilm. Biogenic structures are, therefore, of critical importance for soft-bottom intertidal communities in terms of both structure and function.been referred to as particle reworking, while water movement (if considered) is referred to as bioirrigation in many cases. For consistency, we therefore propose that, for contemporary aquatic scientific disciplines, faunal bioturbation in aquatic environments includes all transport processes carried out by animals that directly or indirectly affect sediment matrices. These processes include both particle reworking and burrow ventilation. With this definition, bioturbation acts as an ‘umbrella’ term that covers all transport processes and their physical effects on the substratum. Particle reworking occurs through burrow construction and maintenance, as well as ingestion and defecation, and causes biomixing of the substratum. Organic matter and microorganisms are thus displaced vertically and laterally within the sediment matrix. Particle reworking animals can be categorized as biodiffusors, upward conveyors, downward conveyors and regenerators depending on their behaviour, life style and feeding type. Burrow ventilation occurs when animals flush their open- or blind-ended burrows with overlying water for respiratory and feeding purposes, and it causes advective or diffusive bioirrigation exchange of solutes between the sediment pore water and the overlying water body. Many bioturbating species perform reworking and ventilation simultaneously. We also propose that the effects of bioturbation on other organisms and associated processes (e.g. microbial driven biogeochemical transformations) are considered within the conceptual framework of ecosystem engineering.Benthic invertebrates have important ecosystem engineering functions (bioturbation and biodeposition) in freshwater and marine benthic systems. Bioturbation and biodeposition affect the metabolism of the water-sediment interface through modification of water-sediment fluxes or organic-matter enrichment of sediments by biodeposits. The functional significance of these processes depends strongly on the type of invertebrate activities (the functional traits of the invertebrates) and on the modulation of this activity by environmental conditions. The aim of my article is to propose a common framework for the role of bioturbation/biodeposition in benthic habitats of both marine and freshwater environments. In these ecosystems, hydrological exchanges between the water and sediments (interstitial flow rates) control the microbial activity inside sediments. The ability of ecosystem engineers to influence benthic microbial processes differs strongly between diffusion-dominated (low interstitial flow rates) and advection-dominated (high interstitial flow rates) habitats. Bioturbation/biodeposition may play a role in diffusion-dominated habitats where invertebrates can significantly modify water and particle fluxes at the water-sediment interface, whereas a slight influence of ecosystem engineers is expected in advection-dominated habitats where fluxes are predominantly controlled by hydrological processes. A future challenge will be to test this general framework in marine and freshwater habitats by quantifying the interactions between the functional traits of species and the water-sediment exchanges.

Chemical properties of burrow wall sediment from burrows of the thalassinidean shrimp Pestarella (=Callianassa) tyrrhena located at Vravrona Bay (Aegean Sea, Greece) were studied and found to be very different from the sediment surface and ambient anoxic sediment. P. tyrrhena burrow walls had significantly higher amounts of silt and clay, while total organic carbon (TOC) was up to 6 times higher than in surrounding sediment. Chlorophyll a (chl a) accounted for a small fraction of TOC and showed similar values in burrow walls and surface sediment, whereas the low chl a: chl a + phaeopigment ratio indicated the presence of more fresh material in the latter. Biopolymers (carbohydrates, proteins and lipids) were 4 to 11 times higher in burrow walls than in the surrounding sediment, accounting for 47% of TOC. The low protein:carbohydrate ratio indicated that the high TOC in the burrow walls was caused by the presence of aged detritus of low nutritional quality, such as seagrass detritus. The distinct conditions along the burrow wall also affected the bacterial community and resulted in a 10-fold increase of bacterial abundance. Molecular fingerprints of the bacterial communities showed that the bacterial composition of the burrow wall was more similar to the ambient anoxic sediment and showed less seasonal change than the sediment surface. These results suggest that burrow walls have distinct properties and should not be considered merely as a simple extension of the sediment surface.The specific microclimates they create within their burrows are particularly important for microbes and meiofauna, which in turn play important roles in organic and inorganic nutrient cycling. The physical turnover of sediments from burrows to the sediment surface significantly influences macro-invertebrate community structure, generally by negatively affecting surface fauna or organisms such as filter-feeders and epibenthic grazers that are dependent on the interface of sediment and water for feeding. Their burrowing activities increase sediment penetrability and porosity, which can favour burrowing macrofaunal species. Sediment turnover can also reduce recruitment of macro-invertebrates, either indirectly by diminishing microbial biofilms that act as food, sediment stabilizers and biochemical cues for larval settlers or directly by burying recruits. Thalassinidean bioturbation also influences marine vegetation, in some instances excluding seagrasses; together with the ecosystem services these plants provide for co-occurring species. Thalassinideans also affect commercial aquaculture operations, for oysters and penaeid shrimps. Sediment turnover by thalassinideans buries adult and juvenile oysters, and their propensity to increase fluxes of toxic nutrients and sulphides, allied with their high oxygen consumption, reduces yields of cultured shrimps, leading to financial losses. Harvesting of thalassinideans for bait has important consequences for soft-sediment ecosystems as the physical disturbance induced by bait-collectors, associated with types of sediment: sandy and muddy sands. The changes observed were compared with abiotic factors and the biomass abundance, which was dominated by benthic diatoms. C-14 uptake values obtained from incubations in a photosynthetron were used for the construction of P-I (photosynthesis-irradianee) curves. Annual averages indicate that both sediments were equally productive (34.5 +/- 23.6 mg C m(-2) h(-1) and 41.1 +/- 11.6 mg C m(-2) h(-1) for the sands and muddy sands respectively), but production rates were highly variable on monthly time scales and were regulated by different mechanisms. Light and temperature played an important role in determining the production rates, especially in the muddy sediments, where changes in I-k (light saturation) were correlated with temperature. I-k showed seasonal changes, suggesting that algae adapted to the seasonal light conditions but there was not a significant correlation between the I-k and PAR (photosynthetic available radiation) at any of the stations. Vertical migration of the algae, as followed by spectroradiometric measurements, probably accounted for a general absence of photoinhibition. In the sandy sediments, production appeared to be limited by the low biomass of algae, due to resuspension and export. The fact that gross oxygen production rates measured on intact cores using microelectrodes were not lower than potential production obtained from 14C fixation suggests that short-term limitation of production due to nutrients and/or carbon is not frequent in the microphytobenthos of the Westerschelde.engineer species on ecosystem functions. We showed that the occurrence of Haploops tubes and individuals significantly modifies sediment features (e.g. change in sediment grain size, increase in C and N organic content) but also largely affect species diversity and benthic composition. The species richness was significantly higher in Haploops community but the species assemblage associated with Haploops habitat was very homogeneous compared to the neighboring habitats and unique with 33% of all species exclusively found in this community. Multivariate analysis (dbRDA) revealed that Haploops density was by far the factor explaining the variation in species composition of benthic communities. No differences in species diversity and assemblage were detected in relationship to Haploops density. A biological trait analysis performed on the whole ecosystem (Haploops included) revealed that Haploops largely dominates the functional structure of the Haploops community by its own functional traits. When performed on selected traits of the associated fauna only (Haploops excluded) the functional structure of the Haploops community was characterized by a greatly reduced proportion of small to medium long lived, sensitive to disturbance, free living or burrowing/tube-building filter-feeding species. H. nirae appears to be a bioengineer and a foundation species that largely modifies its hydro-sedimentary features, controlling diversity and abundances of associated species, and creating a complex set of positive and negative interactions so that a unique benthic assemblage is found in sediments they colonized. and the Bay of Concarneau). To assess the ecological consequences of its recent expansion, the pattern of population dynamics of H. nirae was determined from January 2010 to March 2011 at a 29 m deep station in the Bay of Concarneau with a sampling interval of ca. 3 weeks (23 sampling dates). Modal analysis of size-frequency distribution at each date was used to describe the life cycle and the population dynamics of H. nirae , and to estimate the annual secondary production using the increment summation method. We showed that H. nirae is a semelparous species exhibiting a biannual life cycle with a lifespan ranging from 24 to 28 months. The recruitment was extremely high in 2010 and occurred between January and April but most recruits arrived at the end of March. Recruits are produced by two year old individuals, which die shortly after hatching. Female densities were low in winter. Females produced only one brood during their lifetime with an average brood size of 29. H. nirae secondary production was estimated at 9.66 gDW m−2 y−1 and the production to biomass ratio (P/B¯) was 2.26 y−1. Results are compared with previous published data on other Ampeliscid amphipods. The biannual life cycle appears unusual for most of Ampeliscidae except for the genus Haploops where a biannual life cycle appears characteristic of this genus. The secondary production calculated for H. nirae was high compared to other biannual Ampeliscidae species and constituted one of the highest production values calculated for an Ampeliscid.

Nephrops norvegicus is a burrowing decapod, found in the North Atlantic and Mediterranean Sea at depths of 10–1200 m, and currently the most valuable species taken by the commercial fishing industry in Scotland. It constructs and inhabits extensive burrow complexes in suitable muddy sediments. Owing to its variable emergence patterns, catch rates from traditional trawl surveys are not considered a good indicator of population size. Nephrops populations around Scotland are assessed using an underwater television (UWTV) survey method. Sediment samples are collected at the end of each UWTV deployment. This study focuses on two areas off the coast of Scotland and investigates the accuracy of the sediment maps used for assessment purposes, and the relationship between Nephrops burrow density and sediment composition, over the period 2002–2007. Nephrops have a stock-specific relationship with the sediment they inhabit, which retains the same form through fluctuations in population size.

Many benthic marine invertebrate species have a dispersive larval stage in their life histories. Larvae typically spend hours, weeks, or months developing in plankton before they become competent to settle and metamorphose. Recruitment to benthic populations depends on the numbers of competent larvae transported to sites and/or the interaction between larvae and the surface of substratum. While there is considerable evidence that on large spatial scales, the number of competent larvae transported to sites is determined primarily by hydrodynamics, success of larval settlement on small spatial scales is mediated by biotic and abiotic characteristics of substratum. Larvae of many marine polychaetes require specific cues to settle and metamorphose. Cues can originate from conspecific or congeneric individuals, microbial films, sympatric species, food items, or habitat. Larval settlement in an individual species can be controlled by a single cue or a mixture of cues. Larval settlement of multiple species can be mediated by a common cue or a mixture of cues. Although a variety of chemicals, including proteins, free fatty acids, polysaccharides, inorganic ions, and neurotransmitters, have been suggested as inducing larval settlement of marine polychaetes, few natural cues have been isolated and structurally identified.

Summary of interactions Source Type Confidence

Biotope classification High

High

Biological Traits Medium

High

Medium

Medium

Medium

Technical report by established agency

Describes community bioturbation potential for multiple selected species, quantifying the method and type of bioturbation.

Peer Reviewed Journal Article

Grey literature not all peer reviewed

Sensitivity of Abra albaPeer reviewed grey literature by industry experts

Information on key prey taxa of bivalves (Abra alba,


Describes the temporal variability of the Abra alba community


Biological Traits Abra nititdaNon peer reviewed grey literature by industry experts

High

Medium

Medium

Biological traits of Amphipods Published Book High

Biological traits Global database High

Biological traits of Terebellidae Published Book Medium

High

Medium

Additional information on the relationship of Ampelisca tenuicornis with other taxa (parasite)


Biological traits. Ampelisca migration patterns


Biological triats, amphipod sensitivity to sewages


Sensitivity of Amphiura filiformisPeer reviewed grey literature by industry experts

Biological traits Amphiura filiformis Peer Reviewed Journal Article

Medium

Medium

High

High

High

High



Temporal variability Amphiura filiformis Peer Reviewed Journal Article

Biological traits of Aphelochaeta marioniPeer reviewed grey literature by industry experts

Biological traits of Arenicola marinaPeer reviewed grey literature by industry experts

Temporal variability Arenicola marina Peer Reviewed Journal Article

Medium

High

Medium

Medium

High

High

Biological traits Arenicola marina Peer Reviewed Journal Article

Sensitivity of Brissopsis lyriferaPeer reviewed grey literature by industry experts

Relationships Brissopsis lyrifera Peer Reviewed Journal Article

Biological traits of Callianassa subterraneaNon peer reviewed grey literature by industry experts

Biological traits of Calocaris macandreae Peer Reviewed Journal Article

Biological traits Capitella capitataPeer reviewed grey literature by industry experts

Medium

Medium

Medium

High

High

Medium

Medium

Biological traits Capitella capitata Peer Reviewed Journal Article

Sensitivity Carcinus maenasNon peer reviewed grey literature by industry experts

Sensitivity Cerastoderma eduleNon peer reviewed grey literature by industry experts

Sensitivity Echinocardium cordatumPeer reviewed grey literature by industry experts

Biological traits Hediste diversicolorPeer reviewed grey literature by industry experts

Biological traits of Lagis koreniNon peer reviewed grey literature by industry experts

Biological traits of Macoma balthicaNon peer reviewed grey literature by industry experts

Biological Traits Grey Literature Medium

Biological Traits Medium

Medium

High

Medium

Medium

Medium

Medium

Non peer reviewed grey literature by industry experts

Biological traits of Mya truncataNon peer reviewed grey literature by industry experts

Sensitivity Nephrops norvegicusPeer reviewed grey literature by industry experts

Biological traits of Nephtys hombergiiNon peer reviewed grey literature by industry experts

Biological traits of Ocnus planciNon peer reviewed grey literature by industry experts

Biological traits of Owenia fusiformisNon peer reviewed grey literature by industry experts

Biological traits of Phornois muelleriNon peer reviewed grey literature by industry experts

Medium

Medium

Medium

Medium

Medium

High

Biological traits of Polydora ciliataNon peer reviewed grey literature by industry experts

Biological traits of Pgospio elegansNon peer reviewed grey literature by industry experts

Biological traits of Sagartiogeton undatusNon peer reviewed grey literature by industry experts

Biological traits of Scalibregma inflatumNon peer reviewed grey literature by industry experts

Biological traits of Virgularia mirabilisNon peer reviewed grey literature by industry experts

Predators and habitat of Ampelisca tenuicornis


Predators of Echinoidea High

Medium

High

Peer reviewed book High


Medium

High

Biological traits. Peer reviewed book High


Temporability Echinocardium cordatum Peer Reviewed Journal Article

Biological trait Calocaris macandreae (predators and parasites)


Biological traits of Calocaris macandreae

Biological traits of Echinoderms

Biological traits of Labidoplax mediaNon peer reviewed grey literature by industry experts

Predator of C. maenas Peer Reviewed Journal Article


High

High

Medium

Global database High



Feeding behavour for polychaete worms High


Biological traits Cirriformia tentaculata Peer Reviewed Journal Article

Sensitivity of Cirriformia tentaculata Peer Reviewed Journal Article

Biological traits of Euclymene oerstedii Peer Reviewed Journal Article

Cirriformia tentaculata - Feeding types

Ampharete lindstroemi - Feeding types


Goniada maculata - Depth range

Biological traits of Goniada maculata Medium

Biological traits of Lagis koreni Medium

Prey of flatfish Medium

Medium

High



Medium




Biological trait Magelona johnstoni Peer Reviewed Journal Article

Biological trait Magelona johnstoni Peer Reviewed Journal Article

Malacoceros fuliginosus - Feeding types

Maxmuelleria lankesteri - Feeding types

Biological traits of Maxmuelleria lankesteri Non peer reviewed grey literature by industry experts


Medium


Biological traits Microprotopus maculatus High


Medium

Medium

Biological traits of various bivalve speies. Peer reviewed book High

Medium

Mellina palmata- Feeding types

Biological traits Mellina palmata Peer Reviewed Journal Article

Microprotopus maculatus - Feeding types


Mya truncata - Feeding types

Biological traits Mya truncataNon peer reviewed grey literature by industry experts

Biological traits Mysella bidentata Peer Reviewed Journal Article

Mysella bidentata and Thyasira spp. in circalittoral muddy mixed sediment.



High


Medium

High


Feeding method Phoronis muelleri Medium

Biological traits - Photis longicaudata High

Nuculoma tenuis - Feeding types

Biological traits of Nuculoma tenuis Peer Reviewed Journal Article

Ocnus planci - Feeding types

Parasitism - Ocnus planci Peer Reviewed Journal Article

Biological trait information of various bivalve species.


Biological traits of Molluscs, Opisthobranchs



Biological traits - Polydora ciliata High


Rhodine gracilior - Feeding types Global database High

High



Biological traits of Phoronis muelleri Peer reviewed book High


Biological traits of Thysanocardia procera Peer reviewed book High

High


Medium


Pygospio elegans - Feeding types and depth range

Association with Rhodine gracilior Peer Reviewed Journal Article

Parasite of Sagartiogeton undatus

Biological traits of Anthozoa (Sagartiogeton undatus)

Thysanocardia procera - Feeding types

Biological traits - Tubificoides (pseudogaster)


Biological traits - Tubificoides (pseudogaster)

Biological traits of Magelona johnstoniNon peer reviewed grey literature by industry experts

Biological traits of Nuculoma tenuis Medium

Medium


High

High

High

High

High


Biological Traits - Ampharete lindstroemie Grey literature not all peer reviewed

Biological traits of Balanus crenatus

Senstivity of th urchin Echinus esculentusNon peer reviewed grey literature by industry experts

Biologuical traits of Echnius Technical report by established agency

Aquaculture potential and biological traits of Echinussp.


Echinus sp. and their proximinity to kelp bed.


Overview of marine ecosystem function and drivers



High

Medium

High

Medium

Medium

Biological traits of hermit crab Pagurus bernhardus

Pagarus bernhardus - Intolerant to substratum loss. England East Coast.


Pagarus bernhardus, ophiura albida - Favours areas of physical disturbance. Association of Polydora


Biologial traits. Epifaunal comunities on Pagurus shells.


Scoloplos armiger - Favours Nutrients. Wadden Sea.


Sensitivity of Scoloplos armiger, Nuculoma tenuis and Tubificoides - to physical disturbance. Scottish Loch. Mediomastus fragilis tolerance


Medium

High

High

High

High

Location preference of Scoloplosin feacal mounds of Arenicola marina.


Hesionura elongata, Spiophanes bombyx - Favours Substratum loss. SE England.


Spiophanes bombyx - Favours Substratum loss. North Sea.


Biologial trait of Spiophanes bombyxNon peer reviewed grey literature by industry experts

Sensitivity to trawl fisheries P. pellucidas and opportunisitic behaviour of E. cordatum and Brissopsis lyrifera


High

Medium

High

Medium

IUCN Threatened species Global database High

Medium

Sensitivity Nephtys hombergii to sedimentation rates


Sensitivity to trawl fisheries - Thysanocardia procera and Lagis koreni


Predators of Pygospio elegans, Scoloplos armiger and Tubificoides


Predators of bivalves (Abra alba, Macoma balthica, Mysella bidentata, Nuculoma tenuis)


Predators of Ampharete lindstroemi Peer Reviewed Journal Article

Medium

High

High

Medium

High

Predators of Mellina palmata Peer Reviewed Journal Article

Impacts of climate change on marine benthos. Response of species to climate change will be buffered by environmenatal tolerances, such as tidal stress, energy, sediments etc. Biotope distribution may change. Tolerant species may increase their migration northward, whilst northen species may reduce and decline in abundance.


Climate change will likely change species distribution. Increased rainfall may influence salinity.


Temperature affects on primary production and its limitations.


Uptake of atmospheric CO2 into the oceans reduces pH levels, enhancing ocean acidification


High

High

High

Medium

Medium

Ocean acidification will affect the reproduction and growth of Bivalve species such as Macoma balthica. As an ecosystem engineer, changes will affect coastal diversity and ecosystem functioning.


The major determinant of infaunal communitycomposition is sediment granulometry, with depth being of secondary importance. For epibenthos, depth is the major factor and thesediment composition seemed less significant.


Summary of influencing factors on biological production. Greatest production is noted in areas typified by shallow gravelly areas with high tidal stress and thermal homogeneity. Spatial heterogeneity of sediment granulometric variables occurred primarily between stations while those ofother variables (e.g., depth, stratification, and tidal bed stress) were more regional.


Environmental factors affecting secondary production. Filter feeders have the highest production as do bivalves. Onmivores, predators and arthropods also have high production. Production and P/B ratios were negatively related to water depth and positively related to water temperature


Habitat specific information, relationship between depth and light and primary producers, characterising taxa.


High

Medium

Published Book

High

High

Review of benthic hypoxia High

Published BookHigh

High

Light attenuation is greatly affected by suspended particulate matter and depth.


Effects of depth on light attenuation and the subsequent affect on primary production.

Biological textbook - industry experts. Sources form peer reviewed journals

Nutrient cycling, coupled benthic-pekagic ecosystem.

Overview of marine ecosystem function and drivers



Sediment stability and water flows, influence of microbial activity and infauna (deposit feeders and tube builders)

Impacts of benthic fauna on sediment stability.


High

High

Medium

High

High

Medium

Published Book High

Impact of the presence of deposit feeding bivalves (C. edule) on the water-flow and sediment stability


Stabilising and destabelising effects of fauna on sediments.


Processes concerning waves and water currents


Bioturbation of A. filiformis and implications on the O2 fluxes at the sediment-water interface.


Bioturbation and biogeochemical fluxes of three ecosystem engineers (Abra alba, Lanice conchilega and Nephtys sp.)


Currents play a significant role in the distribution of nutrients and organic carbon recycled by or produced by benthic fauna


Respiration rates in coastal benthic communities

Medium

High

Sediment stability and water flows High

Grey literature Medium

High

Predatory polychaetes Medium

Connections between bioturbation and nitrogen cycling.


Effect of ocean acidication on oxygen and nitrogen cycles



Sediment stability, geology of cohesive mud and biological effects.

Overview of carbon flow within fine sediment benthic communities and higher trophic levels. Indicates relative importance of deposit feeders, bacteria and detritus, over suspension feeders and onmivores.



Bioengineers High

High

Ecology of brittle star beds Peer reviewed report High

Medium

Medium

Medium


Predator-prey Nephtys hombergii Peer Reviewed Journal Article

Role of brittlestars in transfer of particles from water column to benthos. Predation on brittlestars by higher trophic levels.


Deposit-feeding sea cucumbers and their role for mineralisation processes and bacterial activity


Bioturbating Echinocardium and its effects on nutrient cycling and primary production


Predators of Holothurians (sea cucumbers) High

Medium

High

Species distribution Global database Medium

Bioturbation of Carcinus maenas High

Medium

High


Biological traits Magelona johnstoniNon peer reviewed grey literature by industry experts

Bioengineerin activities and their affect on nematode density and diversity



Ecosystem function of hermit crabs, driving influence of DO on Pagurus populations.


Actiniaria provision of habitat, predators and food providers. Not only for amphipods but also fish and crustaceans.


High

Medium

High

High

Medium

High

Anenomes play an important role in benthic–pelagic coupling as part of the benthic suspension feeding community (Sebens and Paine, 1978), transferring energy to the benthos from the water column and releasing metabolites, gametes, and offspring back into the water column. "Their ecological success is undoubtedly facilitated by their propensity for engaging in symbiotic relationships with other animals, including hermit crabs, molluscs, and clown fish."


Influence of ocean acdification to the larval development of sea urchin


Burowing effects of Calocaris macandreae Peer Reviewed Journal Article

Turbidity has an effect on suspension filter feeding bivalve populations. Bivalves increase biodeposition rates, and have a top-down control on phytoplankton.


Sediment stability and its influence on deposit and suspension feeding organisms


Tube dwellers and their association with the the benthic community


High

Medium

Medium

Bioturbation and ecosystem related effects High

Biodeposition and bioturbation High

The effects of tube dwelling polychaetes on the surrounding environment


The effects of tube dwelling Phoronid on the surrounding environment


Impacts of biogenic structures of L.conchilega and Mellina cristata on the ecosystem.




Medium

High

Medium

Medium

Secondary production of Ampeliscids Medium

Medium

Burrowing thalassinidea and the effects on microbial activity


Burrowing Thalassinidea and the ecosystem effects (Microbial activity, nutrient cyling)


Primary production of microphytobenthos in interdal systems


Ampliscids and their structure on the benthic community



Sediment composition and suspension caused by wave energy


Burrow system Nephrops norvegicus High

Various physical and chemical interactions

Medium

Larval settlement of polychaetes Medium




Confidence assessment of quality for individual evidence sources

Quality Requirement

High

Peer reviewed

Or grey literature reports by established agencies

MediumOr expert opinion where feature described is well known/obvious pathway

LowOr no methods adopted and informed through expert judgement

Confidence assessment of applicability for individual evidence sources

Applicability Requirement

High

Study based on UK data

MediumStudy based in UK but uses proxies for CEM component listed

Or study not based in UK but based on exact feature and CEM component listed

LowStudy not based on UK data

Or study based on proxies for feature listed and proxies for CEM component listed

Overall confidence of individual evidence sources based on both quality and applicability

Overall Source ConfidenceApplicability Score

Low Medium High

Quality Score

Low Low Low Low

Medium Low Medium Medium

High Low Medium High

Individual Source Confidence

Does not fulfil ‘high’ requirement but methods are fully described, are considered fit for purpose and to a suitable level of detail

Does not fulfil ‘medium’ requirement for level of detail and fitness for purpose but methods are described

Individual Source Confidence

Or study based on exact feature listed (species, biotope or habitat) and exact CEM component listed (e.g. energy at the seabed)

Quality RequirementPeer reviewed

Or grey literature reports by established agencies

Or expert opinion where feature described is well known/obvious pathway

Or no methods adopted and informed through expert judgement

Applicability RequirementStudy based on UK data

Study based in UK but uses proxies for CEM component listed

Or study not based in UK but based on exact feature and CEM component listed

Study not based on UK data

Or study based on proxies for feature listed and proxies for CEM component listed

Does not fulfil ‘high’ requirement but methods are fully described, are considered fit for

Does not fulfil ‘medium’ requirement for level of detail and fitness for purpose but

Or study based on exact feature listed (species, biotope or habitat) and exact CEM

Table 1. Species Trait Definitions

Species Trait Definitions

Mobility/Movement

Typical Foods Types

Depth Range

Tidal Stream Preference

Physiographic preferences

Salinity preferenceLifespan

Sensitivity To Change

Tolerance to anoxic conditions

Temporal Variable Population

Relationships to other taxa

Table 2. Species Trait Standard Categories (MarLIN, 2006)

Standardised Species Trait CategoriesGrowth FormBoringCrustose hardCrustose softFlaccidMassiveCushionTurfFolioseShrub

Growth Form

Feeding Method

Bioturbator

Environmental Position

Habitat

Size

Distribution

Substratum Preference

Key Prey Taxa

Connectivity to other habitats /species

Arborescent / ArbuscularForestAlgal gravelAccretionMatFaunal bedsRadialStellateWhiplikeStraplike / RibbonlikeFiliform / FilamentousVermiform unsegmentedVermiform segmentedVermiform annulatedDigitate

Lanceolate

PenicillatePinnateCapitate / ClubbedClathrateReticulateFunnel shapedDendroidFlabellateTubicolousMedusiform / MedusoidCylindricalGloboseBullate / SaccateArticulateBivalvedTurbinatePisciformInsufficient InformationNot RelevantNot researchedConicalTadpole

Table 3. Model Level Definitions

Model Level

1. Regional to Global Drivers

5. Output Processes

Table 4. Model Component Definitions

DRIVING INFLUENCES1. Regional to Global DriversPropagule SupplyGeologyDepthWave ExposureWater CurrentsClimate2. Water Column ProcessesPrimary Production

Suspended Sediment

Light Attenuation

Water Chemistry & Temperature

Dissolved Oxygen

3. Local Processes/Inputs at the Seabed

Recruitment

Food Sources

- Plankton

- POM

- Detritus - Phyto-benthos - Carrion - Living Prey - MicrobesSeabed Mobility4. Habitat and Biological Assemblage

2. Water Column Processes

3. Local Processes/Inputs at the Seabed 4. Habitat and Biological Assemblage

6. Local Ecosystem Functions

7. Regional to Global Ecosystem Functions

Tube Building Fauna

Burrowing Fauna

Suspension and Deposit feeding Infauna

Echinoderms and Sessile Epifauna

OUTPUTS5. Output Processes

Supply of Propagules

Bioengineering

Bio-deposition

Secondary ProductionBioturbation6. Local Ecosystem Functions

Nutrient Cycling

Food ResourcesBiogeochemical Cycling

Sediment Stability

Microbial Activity Enhancement

Habitat Provision

7. Regional to Global Ecosystem FunctionsExport of Biodiversity

Export of Organic Matter

Biodiversity Enhancement

Biotope Maintenance

* note that some model components may be grouped together under a composite heading in Worksheet 7

Table 5. Direction of Interactions

Direction of Interaction

Positive

Negative

Mobile Epifauna, Predators and Scavengers

Feedback

Table 6. Magnitude of Interactions

Magnitude of Interaction

Low

Medium

High

Table 1. Species Trait Definitions

Species Trait DefinitionsThe adult body structure of an organism

The processes by which organisms consume their foodThe food which the organisms typically consume

How the organism exists within the environment

The geographical occurrence of individuals around UKThe range of depths the organism is found inThe sediment type that the organisms are likely to favourFavoured location in relation to tidal currents

The typical range of physiographic conditions the organism favoursThe typical range of salinity environments the organism favoursThe length of time the organism is likely to live naturally

An organisms connection and influence to other species or habitatsA species association with another taxa

Table 2. Species Trait Standard Categories (MarLIN, 2006)

Standardised Species Trait CategoriesMobility / MovementSwimmerCrawlerBurrowerDrifterTemporary attachmentPermanent attachmentInsufficient informationNot relevantField unsearched

The organisms ability and method of locomoting through its environment

The method and extent to which an organism is a bioturbator (if applicable)The position within a habitat the organism occupies in relation to the seabed

Average total length of an adult. In the case of worms this is the length from the prostomium to the pygidium, in crabs this is carapace length and in anemones this is the diameter of the disc

Organisms inability to recover from environmental and anthropogenic stressorsThe extent to which the organism is tolerant of sediments deprived of oxygenThe extent to which the organism is regarded as a key food source for other organisms The variation in an organisms population in relation to environmental conditions or annual seasonal fluxes

Temporary attachment / CrawlerCrawler / SwimmerSwimmer / BurrowerSwimmer / Crawler / BurrowerSwimmer / Burrower / Temporary AttachmentCrawler / Burrower

Table 3. Model Level Definitions

DefinitionHigh level influencing inputs to the habitat which drive processes and shape the habitat at a large-scale, e.g. water currents, climate etc. These are largely physical drivers which impact on the water column profile

The characterising fauna and sediment which typifies the habitat

Table 4. Model Component Definitions

DRIVING INFLUENCES1. Regional to Global Drivers

Supply of larvae, spores and/or body fragmentsUnderlying rock or substratum Distance between water surface and sea bedHydraulic wave actionMovement of water masses by tides and/or wind Short term weather and long-term meteorological conditions.

2. Water Column ProcessesThe production of new organic substances through photosynthesis

The penetration of light in the water column

The dissolved oxygen concentration in the water column above the seabed3. Local Processes/Inputs at the Seabed

Types of food ingested by the fauna of the habitat

Organic waste and debris contained within seabed sedimentsPlants and algae attached to the seabedDead and decaying animal fleshLive prey items such as benthic infauna or interstitial faunaMicroorganisms such as bacteria, diatoms and protozoaMovement of sediment on the seabed

4. Habitat and Biological Assemblage

Processes and inputs within the water column which feed into local seabed inputs and processesLocalised inputs and processes to the ecosystem which directly relate to the characterising fauna of the habitat

The specific environmental, chemical and physical processes performed by the biological components of the habitatThe functions resulting from the output processes of the habitat which are applicable on a local scale, whether close to the seabed or within the water columnEcosystem functions which occur as a result of the local processes and functions performed by the biota of the habitat at a regional to global scale.

Particles of sediment which have become elevated from the seabed and are being kept suspended by turbulence within the water column

The chemical and physical characteristics and composition of the water column. This parameter is inclusive of properties such as dissolved oxygen and nutrients

The process by which juvenile organisms join the adult population. Combines settlement and any mortality

Microscopic plants and animals which inhabit the water column (for the purposes of this study, phytoplankton and zooplankton have been grouped together)(Particulate Organic Matter) Non-living material derived from organic sources within the water column

Mobile scavenging and predatory crabs, polychaetes and molluscs

OUTPUTS5. Output Processes

Amount of biomass created as a direct result of consumptionSediment re-working by marine fauna

6. Local Ecosystem Functions

The growth of prey items as a food resource for other organismsThe cycling of organic carbon, nitrogen and oxygen

7. Regional to Global Ecosystem FunctionsExport of biodiversity, including propagules, outside of the habitat

* note that some model components may be grouped together under a composite heading in Worksheet 7

Table 5. Direction of Interactions

Definition

Tubicolous fauna which construct and live in tubes made from sedimentary material on the surface of the seabedBurrowing soft- (e.g. Polychaetes) and hard- bodied (Crustaceans) fauna living within the sedimentsVery small to medium sized suspension and deposit feeding bivalves and smaller infauna

(Sub-) surface urchins, free-living interface suspension/deposit feeding brittle stars and sea cucumbers and permanently attached (sessile) larger, longer-lived surface fauna

The production and transportation of larvae, spores or body fragments capable of regenerationFaunal modification of the natural habitat, e.g. tube building, burrow creation etcThe process by which filter feeding organisms capture particulate matter from the water column and deposit into the sediments

Cycling of organic and inorganic nutrients that involves processing into a different chemical form

Cohesion of sediments into a stable form more resistant to disturbanceEnhanced growth and activity of microbial organisms (e.g. bacteria, diatoms and protozoa) within the sedimentProvision of living space for other organisms through surface attachment of increased habitat complexity

Export of organic material outside of the habitat, such as food sources etc. Enhancements in biodiversity resulting from increased sediment stability and habitat provisionMaintenance of the habitat through sustained production and sediment stability

The CEM component being considered has a positive/enhancing influence on the component it is linked to, e.g. the presence of bioturbation in a habitat links to enhanced biogeochemical cycling.

The CEM component being considered has a negative/destabilising influence on the component it is linked to, e.g. the presence of bioturbation in a habitat links to reduced sediment stability.

Table 6. Magnitude of Interactions

Requirement

The CEM component being considered has an influencing effect on a higher level driver, e.g. the local ecosystem function ‘nutrient cycling’ feeds back to the ‘water chemistry and temperature’ in the water column.

Low level of connection or influence between ecosystem components. Removal of the link would likely not lead to significant changes in the ecosystem. Some degree of connection or influence between ecosystem components. Removal of the connection or influence may lead to changes in the ecosystem.Strong connection or influence between ecosystem components. Removal of the connection or influence would lead to large changes in the ecosystem.

Standardised Species Trait CategoriesFeeding MethodPhotoautotrophActive Suspension FeederPassive Suspension FeederSurface Deposit FeederSub-surface Deposit FeederOmnivoreHerbivoreScavengerSymbiont contribution

PlanktotrophChemoautotrophPredatorInterface feederGrazer (grains/particles)Grazer (fronds/blades)Grazer (surface/substratum)DetritivoreNot RelevantField unresearchedInsufficient informationPredator / ScavengerSurface Deposit Feeder / Sub-surface Deposit FeederSurface Deposit Feeder / DetritivorePassive Suspension Feeder / Active Suspension Feeder

Predator / DetritivoreSurface Deposit Feeder / Predator / Active Suspension FeederSurface Deposit Feeder / Sub-surface Deposit Feeder / ScavengerGrazer (grains/particles/fronds/substratum)

Passive Suspension Feeder / Active Suspension Feeder / Surface Deposit Feeder / Sub-surface Deposit Feeder

Standardised Species Trait CategoriesEnvironmental Position Habitat SizeEpifaunal Attached Very Small (<1cm)Epifloral Bed forming Small (1-2cm)Infaunal Burrow dwelling Small-Medium (3-10cm)Interstitial Ectoparasitic Medium (11-20cm)Demersal Encrusting Medium-Large (21-50cm)Pelagic Erect Large (>50cm)Epibenthic Free livingEpilithic Reef buildingEpiphytic Tubicolous

EpizoicNeustonicPleustonicLithotomousHyperbenthicInsufficient informationNot relevant

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