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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 Sublittoral Rock Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0861) and accompany the report of the same title upon completion of the project. 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 four sections, 3a, 3b, 3c and 3d, 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 255 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed using both the TREx add-in for Excel and a manual species name search.Worksheet 3b indicates the process used to refine the full list of 255 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 N). The resulting list of 159 species identified is presented below the table.Worksheet 3c shows the 159 taxa identified split by taxonomic Order and indicative functional group. Data for the biological zone and energy level of the biotopes the species is listed in are also included.Worksheet 3d shows selected species for each rudimentary functional group. Species have been selected to ensure that the characterising species cover all variables of depth zone and energy level. A fill list of taxa identified for inclusion in the project is shown 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 4 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, and whether the trait was used in defining the ecological groups in Columns D and E. 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 X). 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 Y, 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

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 coarse sediment 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 D. A summary of the information taken from the source is included in Column E (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns F and G 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:

Alexander, D., Coates, D., Tillin, H., Tyler-Walters, H. 2015. Conceptual Ecological Modelling of Sublittoral Rock 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

BUILD STATUS:

Version Date Author0.1 17/11/14 DA0.2 05/03/15 DA0.3 12/03/16 DA1.0 23/03/15 DA

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Copy Version Issue Date

This spreadsheet forms part of the project "Conceptual Ecological Modelling of Sublittoral Rock Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0861) and accompany the report of the same title upon completion of the project. 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 four sections, 3a, 3b, 3c and 3d, 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 255 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed using both the TREx add-in for Excel and a manual species name search.Worksheet 3b indicates the process used to refine the full list of 255 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 N). The resulting list of 159 species identified is presented below

Worksheet 3c shows the 159 taxa identified split by taxonomic Order and indicative functional group. Data for the biological zone and energy level of the biotopes the species is listed in are also included.Worksheet 3d shows selected species for each rudimentary functional group. Species have been selected to ensure that the characterising species cover all variables of depth zone and energy level. A fill list of taxa identified for inclusion in the project is shown 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 4 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, and whether the trait was used in defining the ecological groups in Columns D and E. 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 X). 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 Y, 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 coarse sediment 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 D. A summary of the information taken from the source is included in Column E (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns F and G 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

Alexander, D., Coates, D., Tillin, H., Tyler-Walters, H. 2015. Conceptual Ecological Modelling of Sublittoral Rock 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 Committee

Reason/CommentsDraft version submitted for commentsDraft incorporating revisionsFinal draftFinal

Issued To

This spreadsheet forms part of the project "Conceptual Ecological Modelling of Sublittoral Rock Habitats to Inform Indicator Selection" (Project Ref: C14-0208-0861) and accompany the report of the same title upon completion of the project. 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

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

This worksheet is split into four sections, 3a, 3b, 3c and 3d, 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 255 taxa are listed. The second column indicates the results of a conservation status check performed on all the species listed using both the TREx add-in for

Worksheet 3b indicates the process used to refine the full list of 255 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 N). The resulting list of 159 species identified is presented below

Worksheet 3c shows the 159 taxa identified split by taxonomic Order and indicative functional group. Data for the biological zone and energy level

Worksheet 3d shows selected species for each rudimentary functional group. Species have been selected to ensure that the characterising species cover all variables of depth zone and energy level. A fill list of taxa identified for inclusion in the project is shown 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 4 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, and whether the trait was used in defining the ecological groups in Columns D and E. Definitions of each trait and a list of trait categories (for

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 X). 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 Y, 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 coarse sediment 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

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 D. A summary of the information taken from the source is included in Column E (which correlates with the information presented in Column J of the Interactions Matrix). The source type and confidence assessment are included as Columns F and G 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

Alexander, D., Coates, D., Tillin, H., Tyler-Walters, H. 2015. Conceptual Ecological Modelling of Sublittoral Rock Habitats to Inform Indicator Report No.].

Biotope Code MNCR CodeA3.1 IR.HIRA3.11 IR.HIR.KFaRA3.111 IR.HIR.KFaR.AlaA3.112 IR.HIR.KFaR.AlaAnCrSpA3.113 IR.HIR.KFaR.LhypFaA3.114 IR.HIR.KFaR.LhypParA3.115 IR.HIR.KFaR.LhypRA3.116 IR.HIR.KFaR.FoRA3.117 IR.HIR.KFaR.LhypRA3.12 IR.HIR.KSedA3.121 IR.HIR.KSed.SacA3.122 IR.HIR.KSed.LsacSacA3.123 IR.HIR.KSed.LsacChoRA3.124 IR.HIR.KSed.DesFilRA3.125 IR.HIR.KSed.XKScrRA3.126 IR.HIR.KSed.XKHalA3.127 IR.HIR.KSed.ProtAhnA3.2 IR.MIRA3.21 IR.MIR.KRA3.211 IR.MIR.KR.LdigA3.212 IR.MIR.KR.LhypTA3.213 IR.MIR.KR.LhypTXA3.214 IR.MIR.KR.LhypA3.215 IR.MIR.KR.XFoRA3.216 IR.MIR.KR.LhypVtA3.217 IR.MIR.KR.HiaSwA3.22 IR.MIR.KTA3.221 IR.MIR.KT.LdigTA3.222 IR.MIR.KT.XKTA3.223 IR.MIR.KT.XKTXA3.224 IR.MIR.KT.LsacTA3.225 IR.MIR.KT.FilRVSA3.3 IR.LIRA3.31 IR.LIR.KA3.311 IR.LIR.K.LhypLochA3.312 IR.LIR.K.LhypLsacA3.313 IR.LIR.K.LsacA3.314 IR.LIR.K.LhypCapeA3.315 IR.LIR.K.SarA4.1 CR.HCRA4.11 CR.HCR.FaTA4.111 CR.HCR.FaT.BalTubA4.112 CR.HCR.FaT.CTubA4.12 CR.HCR.DpSpA4.121 CR.HCR.DpSp.PhaAxiA4.13 CR.HCR.XFaA4.131 CR.HCR.XFa.ByErSpA4.132 CR.HCR.XFa.CvirCriA4.133 CR.HCR.XFa.SwiLgAsA4.134 CR.HCR.XFa.FluCoAs

A4.135 CR.HCR.XFa.SpNemAdiaA4.136 CR.HCR.XFa.SubCriTfA4.137 CR.HCR.XFa.FluHocuA4.138 CR.HCR.XFa.MolA4.139 CR.HCR.XFa.SpAnVtA4.2 CR.MCRA4.21 CR.MCR.EcCrA4.211 CR.MCR.EcCr.CarSwiA4.212 CR.MCR.EcCr.CarSpA4.213 CR.MCR.EcCr.UrtScrA4.214 CR.MCR.EcCr.FaAlCrA4.215 CR.MCR.EcCr.AdigVtA4.23 CR.MCR.SfRA4.231 CR.MCR.SfR.PidA4.232 CR.MCR.SfR.PolA4.233 CR.MCR.SfR.HiaA4.241 CR.MCR.CMus.CMytA4.242 CR.MCR.CMus.MdisA4.3 CR.LCRA4.31 CR.LCR.BrAsA4.311 CR.LCR.BrAs.AmenCioA4.312 CR.LCR.BrAs.LgAsSpA4.313 CR.LCR.BrAs.AntAsHA4.314 CR.LCR.BrAs.NeoPro

Biotope NameA3.1 : Atlantic and Mediterranean high energy infralittoral rockA3.11 : Kelp with cushion fauna and/or foliose red seaweeds

A3.116 : Foliose red seaweeds on exposed lower infralittoral rock

A3.12 : Sediment-affected or disturbed kelp and seaweed communities

A3.125 : Mixed kelps with scour-tolerant and opportunistic foliose red seaweeds on scoured or sand-covered infralittoral rock

A3.2 : Atlantic and Mediterranean moderate energy infralittoral rockA3.21 : Kelp and red seaweeds (moderate energy infralittoral rock)

A3.215 : Dense foliose red seaweeds on silty moderately exposed infralittoral rock

A3.22 : Kelp and seaweed communities in tide-swept sheltered conditionsA3.221 : Laminaria digitata, ascidians and bryozoans on tide-swept sublittoral fringe rockA3.222 : Mixed kelp with foliose red seaweeds, sponges and ascidians on sheltered tide-swept infralittoral rockA3.223 : Mixed kelp and red seaweeds on infralittoral boulders, cobbles and gravel in tidal rapidsA3.224 : Laminaria saccharina with foliose red seaweeds and ascidians on sheltered tide-swept infralittoral rockA3.225 : Filamentous red seaweeds, sponges and Balanus crenatus on tide-swept variable-salinity infralittoral rockA3.3 : Atlantic and Mediterranean low energy infralittoral rockA3.31 : Silted kelp on low energy infralittoral rock with full salinityA3.311 : Mixed Laminaria hyperborea and Laminaria ochroleuca forest on moderately exposed or sheltered infralittoral rockA3.312 : Mixed Laminaria hyperborea and Laminaria saccharina on sheltered infralittoral rockA3.313 : Laminaria saccharina on very sheltered infralittoral rockA3.314 : Silted cape-form Laminaria hyperborea on very sheltered infralittoral rockA3.315 : Sargassum muticum on shallow slightly tide-swept infralittoral mixed substrataA4.1 : Atlantic and Mediterranean high energy circalittoral rockA4.11 : Very tide-swept faunal communities on circalittoral rockA4.111 : Balanus crenatus and Tubularia indivisa on extremely tide-swept circalittoral rockA4.112 : Tubularia indivisa on tide-swept circalittoral rockA4.12 : Sponge communities on deep circalittoral rockA4.121 : Phakellia ventilabrum and axinellid sponges on deep, wave-exposed circalittoral rockA4.13 : Mixed faunal turf communities on circalittoral rockA4.131 : Bryozoan turf and erect sponges on tide-swept circalittoral rockA4.132 : Corynactis viridis and a mixed turf of crisiids, Bugula, Scrupocellaria, and Cellaria on moderately tide-swept exposed circalittoral rockA4.133 : Mixed turf of hydroids and large ascidians with Swiftia pallida and Caryophyllia smithii on weakly tide-swept circalittoral rockA4.134 : Flustra foliacea and colonial ascidians on tide-swept moderately wave-exposed circalittoral rock

A3.111 : Alaria esculenta on exposed sublittoral fringe bedrockA3.112 : Alaria esculenta forest with dense anemones and crustose sponges on extremely exposed infralittoral bedrockA3.113 : Laminaria hyperborea forest with a faunal cushion (sponges and polyclinids) and foliose red seaweeds on very exposed infralittoral rockA3.114 : Sparse Laminaria hyperborea and dense Paracentrotus lividus on exposed infralittoral limestoneA3.115 : Laminaria hyperborea with dense foliose red seaweeds on exposed infralittoral rock

A3.117 : Laminaria hyperborea and red seaweeds on exposed vertical rock

A3.121: Saccorhiza polyschides and other opportunistic kelps on disturbed upper infralittoral rockA3.122 : Laminaria saccharina and/or Saccorhiza polyschides on exposed infralittoral rockA3.123 : Laminaria saccharina, Chorda filum and dense red seaweeds on shallow unstable infralittoral boulders and cobblesA3.124 : Dense Desmarestia spp. with filamentous red seaweeds on exposed infralittoral cobbles, pebbles and bedrock

A3.126 : Halidrys siliquosa and mixed kelps on tide-swept infralittoral rock with coarse sedimentA3.127 : Polyides rotundus, Ahnfeltia plicata and Chondrus crispus on sand-covered infralittoral rock

A3.211 : Laminaria digitata on moderately exposed sublittoral fringe rockA3.212 : Laminaria hyperborea on tide-swept, infralittoral rockA3.213 : Laminaria hyperborea on tide-swept infralittoral mixed substrataA3.214 : Laminaria hyperborea and foliose red seaweeds on moderately exposed infralittoral rock

A3.216 : Laminaria hyperborea on moderately exposed vertical rockA3.217 : Hiatella arctica and seaweeds on vertical limestone / chalk

A4.135 : Sparse sponges, Nemertesia spp., and Alcyonidium diaphanum on circalittoral mixed substrataA4.136 : Suberites spp. with a mixed turf of crisiids and Bugula spp. on heavily silted moderately wave-exposed shallow circalittoral rockA4.137 : Flustra foliacea and Haliclona oculata with a rich faunal turf on tide-swept circalittoral mixed substrataA4.138 : Molgula manhattensis with a hydroid and bryozoan turf on tide-swept moderately wave-exposed circalittoral rockA4.139 : Sponges and anemones on vertical circalittoral bedrockA4.2 : Atlantic and Mediterranean moderate energy circalittoral rockA4.21 : Echinoderms and crustose communities on circalittoral rockA4.211 : Caryophyllia smithii and Swiftia pallida on circalittoral rockA4.212 : Caryophyllia smithii, sponges and crustose communities on wave-exposed circalittoral rockA4.213 : Urticina felina and sand-tolerant fauna on sand-scoured or covered circalittoral rockA4.214 : Faunal and algal crusts on exposed to moderately wave-exposed circalittoral rockA4.215 : Alcyonium digitatum and faunal crust communities on vertical circalittoral bedrockA4.23 : Communities on soft circalittoral rockA4.231 : Piddocks with a sparse associated fauna in sublittoral very soft chalk or clayA4.232 : Polydora sp. tubes on moderately exposed sublittoral soft rockA4.233 : Hiatella-bored vertical sublittoral limestone rockA4.241 : Mytilus edulis beds with hydroids and ascidians on tide-swept exposed to moderately wave-exposed circalittoral rockA4.242 : Musculus discors beds on moderately exposed circalittoral rockA4.3 : Atlantic and Mediterranean low energy circalittoral rockA4.31 : Brachiopod and ascidian communities on circalittoral rockA4.311 : Solitary ascidians, including Ascidia mentula and Ciona intestinalis, on wave-sheltered circalittoral rockA4.312 : Large solitary ascidians and erect sponges on wave-sheltered circalittoral rockA4.313 : Antedon spp., solitary ascidians and fine hydroids on sheltered circalittoral rockA4.314 : Neocrania anomala and Protanthea simplex on sheltered circalittoral rock

Substratum

Bedrock; stable boulders

Bedrock; boulders and cobblesBoulders, cobbles, pebbles and gravelBedrock & boulders

Bedrock, cobbles and pebbles with mobile sandBedrock and stable bouldersBedrock & boulders

Bedrock & boulders

Bedrock; boulders & cobbles

Bedrock; boulders and cobbles

Bedrock; boulders

Bedrock; boulders

Bedrock, boulder

Bedrock, boulders

Bedrock & boulders

Bedrock; very large boulders Bedrock Bedrock; massive boulders Bedrock Bedrock; boulders Bedrock; large boulders Bedrock; boulders Bedrock; unstable boulders and cobbles: often nearby coarse sediment Bedrock; boulders

Bedrock; boulders Bedrock, boulders or cobbles with coarse sediment

Bedrock; boulders

Boulders, cobbles, pebbles and gravel Bedrock; boulders

Bedrock Bedrock Bedrock, boulders and cobbles

Bedrock, boulders and cobbles Boulders, cobbles and gravel Boulders, bedrock and gravel

Bedrock, boulders and cobbles Bedrock, boulders, cobbles and mixed substrata Bedrock; boulders Bedrock; boulders

Bedrock & boulders Mixed substrata Bedrock;boulders Bedrock, boulders Bedrock, boulder, cobble

Bedrock Bedrock

Bedrock; boulder Bedrock; very large boulders Bedrock, boulder Bedrock, boulder, cobble

Bedrock

Bedrock, cobbles

Bedrock

Bedrock (soft)

Boulders; cobbles; pebbles Bedrock, boulders Cobble; pebble; mud Bedrock, cobble, sand

Bedrock;boulders Bedrock, boulders Bedrock, boulders Bedrock, boulders

Bedrock, boulders, cobbles

Bedrock Bedrock

Bedrock Bedrock, shells, pebbles, gravel Bedrock, boulders, cobbles Bedrock, boulders Bedrock, boulders Bedrock, boulders, cobbles Bedrock, boulders Bedrock, boulders Bedrock, boulders

Wave exposure

Extremely exposed, Very exposed, Exposed

ExposedExposed, Moderately exposed

Exposed, Moderately exposed, Sheltered

Moderately exposed

Sheltered, Very sheltered, Extremely sheltered

Sheltered, Very sheltered, Extremely sheltered

Extremely exposed, Very exposed, Exposed, Moderately exposed

Exposed, Moderately exposed, Sheltered


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

Moderately exposed Exposed, Moderately exposed Exposed, Moderately exposed, Sheltered Exposed, Moderately exposed, Sheltered

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

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

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

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

Very exposed, Exposed, Moderately exposed


Moderately exposed

Moderately exposed, Sheltered

Very sheltered, Extremely sheltered

Moderately exposed Moderately exposed Exposed

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

Sheltered, Very sheltered Sheltered, Very sheltered Moderately exposed, Sheltered, Very sheltered

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

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

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

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

Moderately strong (1-3 kn), Weak (>1 kn)


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

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


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


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

Moderately strong (1-3 kn), Weak (>1 kn)

Moderately strong (1-3 kn) Moderately strong (1-3 kn), Weak (>1 kn) Moderately strong (1-3 kn), Weak (>1 kn) Moderately strong (1-3 kn), Weak (>1 kn) Moderately strong (1-3 kn), 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)

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

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

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

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

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

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

Weak (>1 kn), Very weak (negligibleWeak (>1 kn), Very weak (negligible

Weak (>1 kn), Very weak (negligible)

Moderately strong (1-3 kn)

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

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

Moderately strong (1-3 kn), 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)

Zone Salinity

Full (30-35 ppt)

InfralittoralInfralittoralInfralittoralInfralittoralInfralittoralInfralittoral

Sublittoral fringe

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

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

Circalittoral - lower, Circalittoral - upper

Circalittoral - lower, Circalittoral - upper

Infralittoral, Sublittoral fringe Full (30-35 ppt) Infralittoral, Sublittoral fringe Sublittoral fringe Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral - upper Full (30-35 ppt) Infralittoral - upper Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral - lower Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral - upper, Sublittoral fringe Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral - upper Full (30-35 ppt) Infralittoral - upper, Sublittoral fringe Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt) Infralittoral Full (30-35 ppt)

Full (30-35 ppt) Infralittoral, Sublittoral fringe Full (30-35 ppt) Sublittoral fringe Full (30-35 ppt)

Full (30-35 ppt) Full (30-35 ppt) Full (30-35 ppt) Full (30-35 ppt) Full (30-35 ppt) Full (30-35 ppt)

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

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

Infralittoral, Sublittoral fringe Infralittoral - upper Full (30-35 ppt)

Full (30-35 ppt)

Infralittoral - upper Full (30-35 ppt) Infralittoral - upper, Sublittoral fringe Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt)

Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt)

Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt)

Full (30-35 ppt), Reduced (18-30ppt)

Circalittoral Circalittoral Full (30-35 ppt)

Full (30-35 ppt)Circalittoral - lower, Circalittoral - upperCircalittoral - lower, Circalittoral - upper

Circalittoral Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral - lower Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Infralittoral - lower, Circalittoral Full (30-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt), Variable (18-35 ppt) Circalittoral - lower, Circalittoral - upper Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral Circalittoral Full (30-35 ppt) Infralittoral - lower, Circalittoral Full (30-35 ppt) Circalittoral - upper Full (30-35 ppt) Circalittoral Full (30-35 ppt) Circalittoral, Infralittoral Full (30-35 ppt), Variable (18-35 ppt)

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

Circalittoral - upper Full (30-35 ppt), Reduced (18-30ppt) Full (30-35 ppt), Reduced (18-30ppt)

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

Depth Band Reference codeR346

0-5 m, 5-10 m, 10-20 m, 20-30 m, 30-50 m, Lower shore R346R346R346R346R346R346R346

0-5 m, 5-10 m, 10-20 m, 20-30 m R346R346R346R346

0-5 m R346R346R346R346

5-10 m R346R346R346R346R346R346

0-5 m, 5-10 m, 10-20 m R346R346R346R346R346R346R346R346

0-5 m R3460-5 m R346

R346R346R346R346

0-5 m, 5-10 m, 10-20 m, 20-30 m R346R346R346R346R346

0-5 m, 5-10 m, 10-20 m, 20-30 m R346R346

20-30 m, 30-50 m R34620-30 m, 30-50 m R346

R34610-20 m, 20-30 m R34610-20 m, 20-30 m R34610-20 m, 20-30 m, 30-50 m R346

R346

0-5 m, 5-10 m, 10-20 m, Lower shore

0-5 m, Lower shore 10-20 m, 20-30 m, 30-50 m 0-5 m, 5-10 m, 10-20 m 0-5 m, 5-10 m 0-5 m, 5-10 m, 10-20 m, 20-30 m 5-10 m, 10-20 m, 20-30 m

0-5 m, 5-10 m, 10-20 m 0-5 m, Lower shore 0-5 m, 5-10 m, 10-20 m, 20-30 m

5-10 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, 10-20 m, Lower shore 0-5 m, 5-10 m, 10-20 m, Lower shore 0-5 m, Lower shore 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 0-5 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, Lower shore 0-5 m, Lower shore 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 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 10-20 m, 20-30 m 10-20 m, 20-30 m

10-20 m, 20-30 m

5-10 m, 10-20 m, 20-30 m

5-10 m, 10-20 m

10-20 m, 20-30 m R346R346

0-5 m, 5-10 m, 10-20 m, 20-30 m R3460-5 m, 5-10 m, 10-20 m R346

R346R346R346R346R346

0-5 m, 10-20 m, 20-30 m, 30-50 m R346R346R346R346

5-10 m, 10-20 m R346R346R346

0-5 m, 5-10 m, 10-20 m, 20-30 m R346R346R346R346R346R346R346R346

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

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

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

0-5 m, 5-10 m, 10-20 m, Lower shore 0-5 m, 5-10 m, 10-20 m, Lower shore

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

Species UK Status/RarityAlcyonium digitatum Presumed nativeUrticina felina Presumed nativeSagartia elegans Presumed nativeCorynactis viridisPomatoceros triqueter Presumed nativeCalliostoma zizyphinum Presumed nativeAsterias rubens Presumed nativeEchinus esculentus Presumed native; IUCN: Lower risk(LR / nt)Botryllus schlosseri Presumed nativeCorallinaceaeCorallina officinalisCallophyllis laciniataPlocamium cartilagineumCryptopleura ramosaDelesseria sanguineaDictyota dichotomaAlaria esculentaLaminaria hyperboreaSemibalanus balanoides Presumed nativePatella vulgata Presumed nativeNucella lapillus Presumed native; OSPAR ListedMytilus edulis Presumed nativePalmaria palmataLithothamnion glacialeMastocarpus stellatusLaminaria digitataCliona celata Presumed nativeAnemonia viridis Presumed nativeParacentrotus lividus Presumed native; Nationally scarceSaccorhiza polyschidesHalichondria panicea Presumed nativeGibbula cineraria Presumed nativeDilsea carnosaKallymenia reniformisLaminaria saccharinaUlva lactucaObelia geniculata Presumed nativeMembranipora membranacea Presumed nativeNitophyllum punctatumPhycodrys rubensCutleria multifidaDesmarestia aculeataDesmarestia viridisLanice conchilega Presumed nativePagurus bernhardus Presumed nativeElectra pilosa Presumed nativeCystoclonium purpureumChondrus crispusPhyllophora pseudoceranoidesHeterosiphonia plumosa

Brongniartella byssoidesChondria dasyphyllaPolysiphonia elongataRhodomela confervoidesEctocarpaceaePunctariaCladostephus spongiosusChorda filumRhodophycotaBonnemaisonia asparagoidesLithothamnionHypoglossum hypoglossoidesDesmarestia ligulataLaminariaEnteromorphaBalanus crenatus Presumed nativeAglaophenia pluma Presumed nativeCancer pagurus Presumed nativeCalliblepharis ciliataPhyllophora crispaHalidrys siliquosaPorifera indet crustsTubularia indivisa Presumed nativePhellia gausapataAmphipodaMargarites helicinus Presumed nativeDidemnidaeBotrylloides leachi Presumed nativePachymatisma johnstonia Presumed nativeCaryophyllia smithii Presumed nativeNecora puber Presumed nativeClavelina lepadiformis Presumed nativeHalarachnion ligulatumMembranoptera alataScypha ciliata Presumed nativeEsperiopsis fucorum Presumed nativeDysidea fragilis Presumed nativeNemertesia antennina Presumed nativeMarthasterias glacialis Presumed nativeSchottera nicaeensisCrisiidaeErythroglossum laciniatumAhnfeltia plicataFurcellaria lumbricalisPolyides rotundusGracilaria gracilisCeramium nodulosumUlvaBryozoa indet crustsAntedon bifida Presumed nativeOphiothrix fragilis Presumed native

Dynamena pumila Presumed nativeLomentaria articulataFucus serratusCladophora rupestrisAlcyonidium diaphanum Presumed nativeCrossaster papposus Presumed nativeHenricia oculata Presumed nativeLithophyllumOdonthalia dentataCaprellidaeMolgula manhattensis Presumed nativeHalurus flosculosusFoliose red algaeHenriciaDercitus bucklandi Presumed nativeTethya aurantium Presumed nativeHydrozoaPolychaetaMaja squinadoHiatella arctica Presumed nativeBugulaAslia lefevreiMorchellium argus Presumed nativeAplidium punctum Presumed nativePolycarpa scuba Presumed nativeCarcinus maenas Presumed nativeSpirorbidaeAlcyonidium hirsutum Presumed nativeAscidiella scabra Presumed nativeDendrodoa grossularia Presumed nativeAscidiella aspersa Presumed nativeOphiocomina nigra Presumed nativeBonnemaisonia hamiferaRhodophycota indet.(non-calc.crusts)PoriferaMyxilla incrustans Presumed nativeSertularia argentea Presumed nativeMetridium senile Presumed nativeActinothoe sphyrodeta Presumed nativeFlustra foliacea Presumed nativeBryozoaPolymastia boletiformisSuberites carnosus Presumed nativeStelligera rigida Presumed nativeStelligera stuposa Presumed nativeAxinella dissimilis Presumed nativeAxinella infundibuliformisPhakellia ventilabrum Presumed native; Nationally scarceHemimycale columella Presumed nativeHaliclona viscosaSertularella gayi Presumed native

Nemertesia ramosa Presumed nativeAlcyonium glomeratum Presumed nativeEunicella verrucosa Presumed native; Protected in UKPorella compressa Presumed nativePentapora foliacea Presumed nativeParasmittina trispinosa Presumed nativeLuidia ciliaris Presumed nativeStichastrella rosea Presumed nativeHolothuria forskali Presumed nativeCiocalypta penicillus Presumed nativeCellepora pumicosa Presumed nativeAbietinaria abietina Presumed nativePododesmus patelliformis Presumed nativeOphiura albida Presumed nativeCiona intestinalis Presumed nativeAscidia mentula Presumed nativeBispira volutacornis Presumed nativePholas dactylus Presumed nativeSuberites ficus Presumed nativePolydoraHymeniacidon perleve Presumed nativeHaliclona oculata Presumed nativeHalecium halecinum Presumed nativeSarcodictyon roseum Presumed nativeInachus phalangium Presumed nativeLeucosolenia botryoides Presumed nativeSabellaria spinulosa Presumed nativeJanolus cristatus Presumed nativeBugula flabellata Presumed nativeBugula plumosa Presumed nativeHenricia sanguinolenta Presumed nativePolyclinum aurantium Presumed nativeHomarus gammarus Presumed nativeMusculus discors Presumed nativeDictyopteris membranaceaKirchenpaueria pinnata Presumed nativeProtanthea simplexMunida rugosa Presumed nativeNeocrania anomala Presumed nativeCorella parallelogramma Presumed nativeAscidia virginea Presumed nativeBougainvillia ramosa Presumed nativeSabella pavonina Presumed nativeBuccinum undatum Presumed nativeTerebratulina retusa Presumed nativePsammechinus miliaris Presumed nativePolycarpa pomaria Presumed nativePolymastia mamillarisMycale rotalisIophon hyndmaniRaspailia ramosa Presumed native

Haliclona fistulosaHaliclona simulansObelia dichotoma Presumed nativeProtula tubularia Presumed nativeTonicella marmorea Presumed nativeAntedon petasus Presumed nativePlumularia setacea Presumed nativeRhodophyllis divaricataChylocladia verticillataPterothamnion plumulaSargassum muticumAntithamnionCallithamnionGriffithsia devoniensisPolysiphonia fucoidesCladophoraSphaerococcus coronopifoliusPolyneura bonnemaisoniiLaminaria ochroleucaCystoseira baccataBryopsis plumosaScrupocellariaCellaria sinuosa Presumed nativeIophonopsis nigricansHaliclona urceolusHalopteris catharina Presumed nativePolyplumaria frutescens Presumed nativeAglaophenia tubuliferaSwiftia pallidaEucratea loricata Presumed nativeSecuriflustra securifrons Presumed nativeLeptometra celtica Presumed nativeDiazona violacea Presumed nativeHydrallmania falcata Presumed nativeCerianthus lloydii Presumed nativeEpizoanthus couchii Presumed nativeProstheceraeus vittatus Presumed nativeBugula turbinata Presumed nativeCellaria fistulosa Presumed nativeTubularia larynx Presumed nativeRhizocaulus verticillatus Presumed nativeHarmothoeTerebellidaePycnogonidaDyopedos porrectus Presumed nativePandalus montagui Presumed nativeCrisia eburnea Presumed nativeHalecium beanii Presumed nativeSertularella gaudichaudi Presumed nativeSagartia troglodytes Presumed nativePisidia longicornis Presumed native

Conopeum reticulum Presumed native

Biotopes to be considered:A3.11 : Kelp with cushion fauna and/or foliose red seaweeds

A3.116 : Foliose red seaweeds on exposed lower infralittoral rock

A3.12 : Sediment-affected or disturbed kelp and seaweed communities

A3.125 : Mixed kelps with scour-tolerant and opportunistic foliose red seaweeds on scoured or sand-covered infralittoral rock

A3.21 : Kelp and red seaweeds (moderate energy infralittoral rock)

A3.215 : Dense foliose red seaweeds on silty moderately exposed infralittoral rock

A3.22 : Kelp and seaweed communities in tide-swept sheltered conditionsA3.221 : Laminaria digitata, ascidians and bryozoans on tide-swept sublittoral fringe rockA3.222 : Mixed kelp with foliose red seaweeds, sponges and ascidians on sheltered tide-swept infralittoral rockA3.223 : Mixed kelp and red seaweeds on infralittoral boulders, cobbles and gravel in tidal rapidsA3.224 : Laminaria saccharina with foliose red seaweeds and ascidians on sheltered tide-swept infralittoral rockA3.225 : Filamentous red seaweeds, sponges and Balanus crenatus on tide-swept variable-salinity infralittoral rockA3.31 : Silted kelp on low energy infralittoral rock with full salinityA3.311 : Mixed Laminaria hyperborea and Laminaria ochroleuca forest on moderately exposed or sheltered infralittoral rockA3.312 : Mixed Laminaria hyperborea and Laminaria saccharina on sheltered infralittoral rockA3.313 : Laminaria saccharina on very sheltered infralittoral rockA3.314 : Silted cape-form Laminaria hyperborea on very sheltered infralittoral rockA3.315 : Sargassum muticum on shallow slightly tide-swept infralittoral mixed substrataA4.1 : Atlantic and Mediterranean high energy circalittoral rockA4.11 : Very tide-swept faunal communities on circalittoral rockA4.111 : Balanus crenatus and Tubularia indivisa on extremely tide-swept circalittoral rockA4.112 : Tubularia indivisa on tide-swept circalittoral rockA4.12 : Sponge communities on deep circalittoral rockA4.121 : Phakellia ventilabrum and axinellid sponges on deep, wave-exposed circalittoral rockA4.13 : Mixed faunal turf communities on circalittoral rockA4.131 : Bryozoan turf and erect sponges on tide-swept circalittoral rockA4.132 : Corynactis viridis and a mixed turf of crisiids, Bugula, Scrupocellaria, and Cellaria on moderately tide-swept exposed circalittoral rockA4.133 : Mixed turf of hydroids and large ascidians with Swiftia pallida and Caryophyllia smithii on weakly tide-swept circalittoral rockA4.134 : Flustra foliacea and colonial ascidians on tide-swept moderately wave-exposed circalittoral rockA4.135 : Sparse sponges, Nemertesia spp., and Alcyonidium diaphanum on circalittoral mixed substrataA4.137 : Flustra foliacea and Haliclona oculata with a rich faunal turf on tide-swept circalittoral mixed substrataA4.138 : Molgula manhattensis with a hydroid and bryozoan turf on tide-swept moderately wave-exposed circalittoral rockA4.139 : Sponges and anemones on vertical circalittoral bedrock

A3.111 : Alaria esculenta on exposed sublittoral fringe bedrockA3.112 : Alaria esculenta forest with dense anemones and crustose sponges on extremely exposed infralittoral bedrockA3.113 : Laminaria hyperborea forest with a faunal cushion (sponges and polyclinids) and foliose red seaweeds on very exposed infralittoral rockA3.115 : Laminaria hyperborea with dense foliose red seaweeds on exposed infralittoral rock

A3.117 : Laminaria hyperborea and red seaweeds on exposed vertical rock

A3.121: Saccorhiza polyschides and other opportunistic kelps on disturbed upper infralittoral rockA3.122 : Laminaria saccharina and/or Saccorhiza polyschides on exposed infralittoral rockA3.123 : Laminaria saccharina, Chorda filum and dense red seaweeds on shallow unstable infralittoral boulders and cobblesA3.124 : Dense Desmarestia spp. with filamentous red seaweeds on exposed infralittoral cobbles, pebbles and bedrock

A3.126 : Halidrys siliquosa and mixed kelps on tide-swept infralittoral rock with coarse sedimentA3.127 : Polyides rotundus, Ahnfeltia plicata and Chondrus crispus on sand-covered infralittoral rock

A3.211 : Laminaria digitata on moderately exposed sublittoral fringe rockA3.212 : Laminaria hyperborea on tide-swept, infralittoral rockA3.213 : Laminaria hyperborea on tide-swept infralittoral mixed substrataA3.214 : Laminaria hyperborea and foliose red seaweeds on moderately exposed infralittoral rock

A3.216 : Laminaria hyperborea on moderately exposed vertical rockA3.217 : Hiatella arctica and seaweeds on vertical limestone / chalk

A4.21 : Echinoderms and crustose communities on circalittoral rockA4.211 : Caryophyllia smithii and Swiftia pallida on circalittoral rockA4.212 : Caryophyllia smithii, sponges and crustose communities on wave-exposed circalittoral rockA4.213 : Urticina felina and sand-tolerant fauna on sand-scoured or covered circalittoral rockA4.214 : Faunal and algal crusts on exposed to moderately wave-exposed circalittoral rockA4.215 : Alcyonium digitatum and faunal crust communities on vertical circalittoral bedrockA4.23 : Communities on soft circalittoral rockA4.231 : Piddocks with a sparse associated fauna in sublittoral very soft chalk or clayA4.232 : Polydora sp. tubes on moderately exposed sublittoral soft rockA4.233 : Hiatella-bored vertical sublittoral limestone rockA4.241 : Mytilus edulis beds with hydroids and ascidians on tide-swept exposed to moderately wave-exposed circalittoral rockA4.242 : Musculus discors beds on moderately exposed circalittoral rockA4.31 : Brachiopod and ascidian communities on circalittoral rockA4.311 : Solitary ascidians, including Ascidia mentula and Ciona intestinalis, on wave-sheltered circalittoral rockA4.312 : Large solitary ascidians and erect sponges on wave-sheltered circalittoral rockA4.313 : Antedon spp., solitary ascidians and fine hydroids on sheltered circalittoral rockA4.314 : Neocrania anomala and Protanthea simplex on sheltered circalittoral rock

Fauna identified from above table selected for inclusion in project:Abietinaria abietinaActinothoe sphyrodetaAglaophenia plumaAglaophenia tubuliferaAhnfeltia plicataAlaria esculentaAlcyonidium diaphanumAlcyonium digitatumAlcyonium glomeratumAnemonia viridisAntedon bifidaAntedon petasusAntithamnion Aplidium punctumAscidia mentulaAscidia virgineaAscidiella aspersaAscidiella scabraAsterias rubensAxinella dissimilisAxinella infundibuliformisBalanus crenatusBispira volutacornisBonnemaisonia asparagoidesBonnemaisonia hamiferaBotryllus schlosseriBugula flabellataBugula plumosaCalliblepharis ciliataCalliostoma zizyphinumCallithamnion Callophyllis laciniata

Cancer pagurusCarcinus maenasCaryophilia smithiiCellaria Cellepora pumicosaCeramium nodolosumCerianthus lloydiiChondrus crispusChorda filumCiocalypta penicillusCiona intestinalisCladophora rupestrisClavelina lepadiformisCliona celataCorallina officinalisCorella parallelogrammaCorynactis virdisCrisia eburneaCrossaster papposusCryptopleura ramosaDelesseria sanguineaDesmarestia aculeataDesmarestia ligulataDesmarestia viridisDexamine spinosaDictyota dichotomaDilsea carnosaDynanema pumilaDyopedos porrectusDysidea fragilisEchinus esculentusElectra pilosaEnteromorpha intestinalisEpizoanthus couchiiEsperiopsis fucorumFlustra foliaceaFucus serratusGibbula cinerariaHalecium beaniiHalecium halecinumHalichondria paniceaHaliclona oculataHaliclona viscosaHalidrys siliquosaHarmothoe Henricia oculataHiatella arcticaHolothuria forskaliHomarus gammarusHydrallmania falcataHymeniacidon perleve

Hypoglossum hypoglossoidesIophonopsis nigricansJanolus cristatusKallymenia reniformisKirchenpaueria pinnataLaminaria digitataLaminaria hyperboreaLaminaria ochroleucaLaminaria saccharinaLanice conchilegaLeucosolenia Lithothamnion gracialeLuidia ciliarisMargarites helicinusMarthasterias glacialisMastocarpus stellatusMembranipora membranaceaMetridium senileMolgula manhattensisMorchellium argusMusculus discorsMytilus edulisMyxilla incrustansNecora puberNemertesia antenninaNemertesia ramosaNeocrania anomalaNucella lapillusObelia geniculataOphiocomina nigraOphiothrix fragilisPachymatisma johnstoniaPagurus bernhardusPalmaria palmataPandalus montagui Paracentrotus lividusParasmittina trispinosaPatella vulgataPentapora foliaceaPhakellia ventilabrumPhellia gausapataPholas dactylusPhycodrys rubensPlocamium cartilagineumPododesmus patelliformisPolyclinum aurantiumPolydora Polyides rotundusPomatoceros triqueterPorella compressaProtanthea simplex

Protula tubulariaRaspailia ramosaRhodomela confervoidesSabella pavoniaSaccorhiza polyschidesSagartia elegansSarcodictyon roseumSargassum muticumSargatia elegansSchottera nicaeensisScypha ciliataSecuriflustra securifronsSemibalanus balanoidesSertularella gaudichaudiSertularia argenteaStelligera stuposaSuberites carnosusSuberites ficusSwiftia pallidaTerebratulina retusaTethya aurantiumTubularia indivisaUlva lactucaUrticina felinaEunicella verrucosa

Title Species 1 Title Species 2 Title Species 3

Alaria esculentaAlaria esculentaLaminaria hyperboreaLaminaria hyperborea Foliose red seaweeds

Laminaria hyperborea

Laminaria saccharina Saccorhiza polyschidesChorda filum Laminaria saccharinaDesmarestia aculeata Desmarestia ligulata

Halidrys siliquosaChondrus crispus Polyides rotundus Ahnfeltia plicata

Laminaria digitataLaminaria hyperboreaLaminaria hyperboreaLaminaria hyperborea

Laminaria hyperboreaHiatella arctica

Laminaria digitata

Laminaria saccharina Laminaria hyperboreaLaminaria saccharina

Laminaria hyperborea Laminaria ochroleucaLaminaria hyperborea Laminaria saccharinaLaminaria saccharinaLaminaria hyperboreaSargassum muticum

Balanus crenatusTubularia indivisa

Phakellia ventilabrum

Corynactis viridis Bugula Cellaria Swiftia pallida Caryophilia smithiiFlustra folicaeaSuberites ficus Suberites carnosusHaliclona oculata Flustra foliaceaMolgula manhattensis

Caryophyllia smithii Swiftia pallidaCaryophyllia smithiiUrticina felina

Alcyonium digitatum

Polydora Hiatella arcticaMytilus edulisMusculus discors

Ciona intestinalis Ascidia mentula

Antedon bifida Antedon petasusNeocrania anomala Protanthea simplex

Important species 1 Important species 2 Important species 3Laminaria hyperborea Alaria esculentaSemibalanus balanoides Laminaria digitata Mastocarpus stellatusSagartia elegans Phellia gausapata Corynactis viridisDelesseria sanguinea Phycodrys rubens Corallina officinalis

Dilsea carnosa Hypoglossum hypoglossoides Schottera nicaeensisDelesseria sanguinea Cryptopleura ramosa Corynactis viridisLaminaria saccharina Saccorhiza polyschides Halidrys siliquosaSaccorhiza polyschides Laminaria saccharina Alaria esculentaDesmarestia viridis Plocamium cartilagineum Alcyonium digitataDesmarestia aculeata Chondrus crispusBonnemaisonia asparagoidesLaminaria saccharina Laminaria hyperborea Saccorhiza polyschidesDictyota dichotoma Aglaophenia pluma Botryllus schlosseriRhodomela confervoides Urticina felinaLaminaria digitata Laminaria hyperboreaPalmaria palmata Halichondria panicea Pomatoceros triqueterDictyota dichotoma Cryptopleura ramosa Botryllus schlosseriDictyota dichotoma Cryptopleura ramosa Membranipora membranaceaDelesseria sanguinea Phycodrys rubens Antedon bifidaPlocamium cartilagineum Calliblepharis ciliata Dysidea fragilisAlcyonium digitatum Antedon bifida Echinus esculentusCliona celata Dysidea fragilis Pachymatisma johnstoniaLaminaria digitata Laminaria hyperborea Laminaria saccharinaHalichondria panicea Chondrus crispus Corallina officinalisLaminaria hyperborea Halidrys siliquosa Dictyota dichotomaDictyota dichotoma Corallina officinalis Anemonia viridisDesmarestia ligulata Sargassum muticum Callophyllis laciniataCallithamnion Antithamnion Ceramium Laminaria hyperborea Laminaria saccharinaCallophyllis laciniata Dictyota dichotoma Corynactis viridisPlocamium cartilagineum Cryptopleura ramosa Dictyota dichotomaDelesseria sanguinea Phycodrys rubens Chorda filumPhycodrys rubens Delesseria sanguinea Bonnemaisonia hamiferaLaminaria saccharina Obelia geniculata Chorda filumPachymatisma johnstonia Halichondria panicea Esperiopsis fucorumTubularia indivisa Balanus crenatus Halichondria paniceaTubularia indivisa Halichondria panicea Sertularia argenteaBalanus crenatus Pachymatisma johnstonia Esperiopsis fucorumPhakellia ventilabrum Axinella infundibuliformis Axinella dissimilisAxinella infundibuliformis Axinella dissimilis Stelligera stuposaHalecium halecinum Nemertesia antennina Alcyonidium diaphanumAlcyonidium diaphanum Flustra foliacea Pentapora foliaceaCaryophyllia smithii Pachymatisma johnstonia Cliona celataClavelina lepadiformis Ascidia mentula Aglaophenia tubuliferaAlcyonidium diaphanum Bugula flabellata Alcyonium digitatumDysidea fragilis Scypha ciliata Nemertesia antenninaAlcyonidium diaphanum Crisia eburnea Urticina felinaNemertesia antennina Halecium beanii Hydrallmania falcataNemertesis antennina Tubularia indivisa Alcyonidium diaphanum

Parasmittina trispinosa Asterias rubens Ophiothrix fragilisParasmittina trispinosa Echinus esculentus Nemertesia antenninaAlcyonium digitatum Echinus esculentus Cliona celataCiocalypta penicillus Balanus crenatus Pomatoceros triqueterParasmittina trispinosa Echinus esculentus Alcyonium digitatumEchinus esculentus Parasmittina trispinosa Pomatoceros triqueterPholas dactylus Polydora Bispira volutacornisPholas dactylus Dysidea fragilis Suberites carnosusCliona celata Halichondria panicea Haliclona oculataNemertesia antennina Bugula plumosa Bugula flabellateAsterias rubens Urticina felina Necora puber Tethya aurantium Scypha ciliata Pachymatisma johnstoniaCiona intestinalis Ascidia mentula Ascidia virgineaEchinus esculentus Pomatoceros triqueter Caryophyllia smithiiAscidia mentula Aplidium punctum Corella parallelogrammaNeocrania anomala Pododesmus patelliformis Pomatoceros triqueterPomatoceros triqueter Terebratulina retusa Pododesmus patelliformis

Important species 4 Important species 5 Important species 6

Palmaria palmata Lithothamnion graciale Patella vulgataMargarites helicinus Cancer pagurusAlcyonium digitatum Sagartia elegans Corynactis viridis

Calliostoma zizyphinum Echinus esculentus Asterias rubens

Corallina officinalis Kallymenia reniformis Callophyllis laciniataPomatoceros triqueter

Desmarestia aculeata Balanus crenatusCancer pagurus Gibbula cineraria

Patella vulgata Gibbula cineraria Dynanema pumilaDelesseria sanguinea Obelia geniculata Electra pilosaClavelina lepadiformis Crossaster papposus Henricia oculataClavelina lepadiformis Pomatoceros triqueter Gibbula cinerariaHalichondria panicea Tubularia indivisa Sabellaria spinulosa

Necora puber Clavelina lepadiformis Calliostoma zizyphinum

Ulva lactuca Enteromorpha intestinalis Leucosolenia Chondrus crispus Balanus crenatus Pomatoceros triqueterPagurus bernhardus Ophiocomina nigra Ophiothrix fragilisUlva lactuca Halichondria panicea Esperiopsis fucorumCladophora Carcinus maenas Mytilus edulis

Caryophyllia smithiiClavelina lepadiformis Antedon bifida Obelia geniculataPomatoceros triqueter Carcinus maenas Pagurus bernhardusAscidiella aspersa Antedon bifida Asterias rubensDexamine spinosa Ceramium nodolosumTubularia indivisa Balanus crenatus Alcyonium digitatumMyxilla incrustans Alcyonium digitatum Sagartia elegansSagartia elegans Urticina felina Metridium senileAlcyonium digitatum Sagartia elegans Sertularia argenteaCaryophyllia smithii Corynactis virdis Holothuria forskaliCaryophyllia smithii Corynactis virdis Holothuria forskaliFlustra foliacea Scypha ciliata Pachymatisma johnstoniaNemertesia antennina Nemertesia ramosa Halecium halecinumNemertesia antennina Actinothoe sphyrodeta Pentapora foliaceaNemertesia antennina Pomatoceros triqueter Antedon petasusUrticina felina Nemertesia antennina Scypha ciliataHalecium halecinum Sertularia argentea Epizoanthus couchiiCerianthus lloydii Janolus cristatus Alcyonium digitatumSertularella gaudichaudi Flustra foliacea Alcyonidium diaphanumCliona celata Pachymatisma johnstonia Echinus esculentus

Echinus esculentus Nemertesia antennina Abietinaria abietinaAlcyonium glomeratum Alcyonium digitatumHaliclona viscosa Marthasterias glacialis Asterias rubensCellepora pumicosa Alcyonidium diaphanum Cliona celataAbietinaria abietina Caryophyllia smithii Pomatoceros triqueterCaryophyllia smithii Asterias rubens Ophiothrix fragilisCliona celata Suberites ficus Flustra foliaceaBispira volutacornis Necora puber Cancer pagurusHymeniacidon perleve Alcyonium digitatum Sarcodictyon roseumJanolus cristatus Alcyonidium diaphanum Flustra foliaceaCarcinus maenas Homarus gammarus Cancer pagurusDysidea fragilis Cliona celata Stelligera stuposaClavelina lepadiformis Neocrania anomala Pomatoceros triqueter

Ascidia virginea Botryllus schlosseri Esperiopsis fucorumProtula tubularia Ascidia mentula Kirchenpaueria pinnataSabella pavonina Ciona intestinalis Ophiothrix fragilis

Important species 7 Important species 8 Important species 9

Halichondria panicea Pachymatisma johnstonia

Ulva Dictyota dichotoma

Mastocarpus stellatus Fucus serratus Cladophora rupestrisPachymatisma johnstonia Calliostoma zizyphinum

Balanus crenatus

Scypha ciliata Electra pilosa Ascidiella scabraHalichondria panicea Anemonia viridis Gibbula cineraria

Balanus crenatus Pomatoceros triqueter

Echinus esculentus Asterias rubens Necora puberGibbula cineraria Antedon bifidaGibbula cineraria Obelia geniculata

Actinothoe sphyrodeta Esperiopsis fucorum Alcyonidium diaphanumAsterias rubens Nucella lapillusFlustra foliacea Cancer pagurus Calliostoma zizyphinumAlcyonium digitatum Pentapora foliacea Nemertesia antenninaAlcyonium digitatum Pentapora foliacea Nemertesia antenninaAlcyonium digitatum Urticina felina Sagartia elegansAlcyonium digitatum Raspailia ramosa Caryophylia smithiiMarthasterias glacialis Asterias rubens Necora puberAlcyonium digitatum Echinus esculentus Securiflustra securifronsBalanus crenatus Cancer pagurus Asterias rubensAsterias rubens Clavelina lepadiformis Alcyonium digitatumPandalus montagui Dyopedos porrectus Harmothoe Lanice conchilega Urticina felina Pagurus bernhardusAsterias rubens Marthasterias glacialis Henricia oculata

Luidia ciliaris Porella compressa Nemertesia antenninaAsterias rubensPododesmus patelliformisAntedon bifida Clavelina lepadiformis Ciona intestinalisAlcyonium digitatum Asterias rubens Mytilus edulis

Halecium halecinumCellapora pumicosa. Ophiothrix fragilis Asterias rubensSargatia elegans Balanus crenatusFlustra foliacea Bugula plumosaPododesmus patelliformis Caryophyllia smithii

Dysidea fragilis Suberites carnosus Caryophyllia smithiiCaryophyllia smithii Parasmittina trispinosa Echinus esculentus

Important species 10

Marthasterias glacialisCancer pagurusIophonopsis nigricans

Sabella pavoniaHolothuria forskali

Corynactis viridis

Ascidia mentulaNecora puber

Polyclinum aurantium

Echinus esculentusAsterias rubens

Condensed Species List OrderBonnemaisonia asparagoides BONNEMAISONIALESBonnemaisonia hamifera BONNEMAISONIALESPalmaria palmata PALMARIALESAhnfeltia plicata AHNFELTIALESCorallina officinalis CORALLINALESLithothamnion glaciale CORALLINALESCalliblepharis ciliata GIGARTINALESDilsea carnosa GIGARTINALESChondrus crispus GIGARTINALESCallophyllis laciniata GIGARTINALESMastocarpus stellatus GIGARTINALESSchottera nicaeensis GIGARTINALESKallymenia reniformis GIGARTINALESPolyides rotundus GIGARTINALESPlocamium cartilagineum PLOCAMIALESAntithamnion CERAMIALESCallithamnion CERAMIALESCeramium virgatum CERAMIALESCryptopleura ramosa CERAMIALESDelesseria sanguinea CERAMIALESHypoglossum hypoglossoides CERAMIALESPhycodrys rubens CERAMIALESRhodomela confervoides CERAMIALESDictyota dichotoma DICTYOTALESDesmarestia aculeata DESMARESTIALESDesmarestia viridis DESMARESTIALESDesmarestia ligulata DESMARESTIALESAlaria esculenta LAMINARIALESChorda filum LAMINARIALESLaminaria digitata LAMINARIALESLaminaria hyperborea LAMINARIALESSaccharina latissima LAMINARIALESSaccorhiza polyschides LAMINARIALESLaminaria ochroleuca LAMINARIALESHalidrys siliquosa FUCALESFucus serratus FUCALESSargassum muticum FUCALESEnteromorpha intestinalis ULVALESUlva lactuca ULVALESCladophora rupestris CLADOPHORALESLeucosolenia LEUCOSOLENIIDAScypha ciliata LEUCOSOLENIIDAPachymatisma johnstonia ASTROPHORIDATethya aurantium HADROMERIDASuberites carnosus HADROMERIDASuberites ficus HADROMERIDA

Stelligera stuposa HADROMERIDACliona celata HADROMERIDAAxinella dissimilis HALICHONDRIDAPhakellia ventilabrum HALICHONDRIDACiocalypta penicillus HALICHONDRIDAHalichondria panicea HALICHONDRIDAHymeniacidon perleve HALICHONDRIDAAxinella infundibuliformis HALICHONDRIDAEsperiopsis fucorum POECILOSCLERIDAMyxilla incrustans POECILOSCLERIDARaspailia ramosa POECILOSCLERIDAIophonopsis nigricans POECILOSCLERIDAHaliclona oculata HAPLOSCLERIDAHaliclona viscosa HAPLOSCLERIDADysidea fragilis DICTYOCERATIDATubularia indivisa CAPITATAHalecium beanii CONICAHalecium halecinum CONICAAbietinaria abietina CONICADynamena pumila CONICAHydrallmania falcata CONICASertularella gaudichaudi CONICASertularia argentea CONICAKirchenpaueria pinnata CONICANemertesia antennina CONICANemertesia ramosa CONICAAglaophenia pluma CONICAAglaophenia tubulifera CONICAObelia geniculata PROBOSCOIDASarcodictyon roseum STOLONIFERAAlcyonium digitatum ALCYONACEAAlcyonium glomeratum ALCYONACEASwiftia pallida ALCYONACEAEunicella verrucosa GORGONACEACaryophyllia smithii SCLERACTINIACerianthus lloydii CERIANTHARIAEpizoanthus couchii ZOANTHARIAAnemonia viridis ACTINIARIAUrticina felina ACTINIARIAMetridium senile ACTINIARIASagartia elegans ACTINIARIAActinothoe sphyrodeta ACTINIARIAPhellia gausapata ACTINIARIAProtanthea simplex ACTINIARIACorynactis viridis CORALIMORPHARIAHarmothoe PHYLLODOCIDAPolydora SPIONIDALanice conchilega TEREBELLIDA

Bispira volutacornis SABELLIDASabella pavonina SABELLIDAPomatoceros triqueter SABELLIDAProtula tubularia SABELLIDASemibalanus balanoides THORACICABalanus crenatus THORACICADexamine spinosa AMPHIPODADyopedos porrectus AMPHIPODAPandalus montagui DECAPODAHomarus gammarus DECAPODAPagurus bernhardus DECAPODACancer pagurus DECAPODANecora puber DECAPODACarcinus maenas DECAPODAPatella vulgata PATELLIDAMargarites helicinus VETIGASTROPODIDAGibbula cineraria VETIGASTROPODIDACalliostoma zizyphinum VETIGASTROPODIDANucella lapillus NEOGASTROPODAJanolus cristatus NUDIBRANCHIAMytilus edulis MYTILOIDAMusculus discors MYTILOIDAPododesmus patelliformis OSTREOIDAHiatella arctica MYOIDAPholas dactylus MYOIDANeocrania anomala CRANIIDATerebratulina retusa TEREBRATULIDACrisia eburnea CYCLOSTOMATIDAAlcyonidium diaphanum CTENOSTOMATIDAMembranipora membranacea CHEILOSTOMATIDAElectra pilosa CHEILOSTOMATIDAFlustra foliacea CHEILOSTOMATIDASecuriflustra securifrons CHEILOSTOMATIDABugula flabellata CHEILOSTOMATIDABugula plumosa CHEILOSTOMATIDACellaria CHEILOSTOMATIDAPorella compressa CHEILOSTOMATIDAPentapora fascialis CHEILOSTOMATIDAParasmittina trispinosa CHEILOSTOMATIDACellepora pumicosa CHEILOSTOMATIDAAntedon bifida COMATULIDAAntedon petasus COMATULIDALuidia ciliaris PAXILLOSIDACrossaster papposus VELATIDAHenricia oculata SPINULOSIDAAsterias rubens FORCIPULATIDAMarthasterias glacialis FORCIPULATIDAOphiothrix fragilis OPHIURIDA

Ophiocomina nigra OPHIURIDAEchinus esculentus ECHINOIDAHolothuria forskali ASPIDOCHIROTIDAClavelina lepadiformis ENTEROGONAPolyclinum aurantium ENTEROGONAMorchellium argus ENTEROGONAAplidium punctum ENTEROGONACiona intestinalis ENTEROGONACorella parallelogramma ENTEROGONAAscidiella aspersa ENTEROGONAAscidiella scabra ENTEROGONAAscidia mentula ENTEROGONAAscidia virginea ENTEROGONABotryllus schlosseri PLEUROGONAMolgula manhattensis PLEUROGONA

Indicative Group Biological Zone Energy LevelRed algae - filamentous Infralittoral H/LRed algae - filamentous Infralittoral M/LRed algae - foliose Infralittoral H/MRed algae - filamentous Infralittoral HRed algae - calcerous Infralittoral H/M/LRed algae - calcerous Infralittoral H/MRed algae - foliose Infralittoral H/MRed algae - foliose Infralittoral H/M/LRed algae - foliose Infralittoral H/M/LRed algae - foliose Infralittoral H/M/LRed algae - foliose Infralittoral H/MRed algae - foliose Infralittoral HRed algae - foliose Infralittoral H/MRed algae - foliose Infralittoral HRed algae - foliose Both H/M/LRed algae - foliose Infralittoral MRed algae - foliose Infralittoral MRed algae - foliose Infralittoral H/M/LRed algae - foliose Both H/M/LRed algae - foliose Both H/M/LRed algae - foliose Infralittoral H/MRed algae - foliose Infralittoral H/M/LRed algae - foliose Infralittoral HBrown algae - foliose Both H/M/LBrown algae - filamentous Infralittoral H/M/LBrown algae - foliose Infralittoral HBrown algae - foliose Infralittoral H/MKelp Infralittoral HKelp Infralittoral H/LKelp Infralittoral H/MKelp Infralittoral H/M/LKelp Infralittoral H/M/LKelp Infralittoral H/LKelp Infralittoral LBROWN algae - branched Infralittoral H/MBROWN algae - branched Infralittoral MBrown algae - branched Infralittoral M/LGreen algae - foliose Infralittoral H/MGreen algae - foliose Infralittoral H/M/LGreen algae - foliose Infralittoral MSponge Circalittoral H/MSponge Both H/MSponge Both H/M/LSponge Both H/M/LSponge Circalittoral H/M/LSponge Circalittoral M

Sponge Both H/M/LSponge - boring Both H/M/LSponge Circalittoral HSponge Circalittoral HSponge Circalittoral MSponge Both H/MSponge Both MSponge Circalittoral HSponge Both H/MSponge Circalittoral H/MSponge Circalittoral H/LSponge Circalittoral HSponge Circalittoral H/MSponge Circalittoral MSponge Both H/M/LHydroid Both H/MHydroid Circalittoral HHydroid Circalittoral H/M/LHydroid Circalittoral M/HHydroid Infralittoral MHydroid Circalittoral HHydroid Circalittoral HHydroid Circalittoral HHydroid Circalittoral LHydroid Both H/M/LHydroid Circalittoral H/M/LHydroid Infralittoral HHydroid Circalittoral HHydroid Infralittoral H/M/LSoft coral Circalittoral MSoft coral Both H/M/LSoft coral Circalittoral H/MSoft coral Circalittoral MSoft coral Circalittoral HHard coral Both H/M/LAnenome - burrowing Circalittoral HAnenome - colonial Circalittoral HAnenome Infralittoral M/LAnenome Both H/MAnenome Circalittoral H/M/LAnenome Both H/MAnenome Circalittoral HAnenome Infralittoral HAnenome Circalittoral LAnenome Circalittoral MPolychaete - scaleworm Circalittoral HPolychaete Circalittoral MPolychaete - tube building Both H/M

Polychaete - tube building Circalittoral MPolychaete - tube building Circalittoral H/LPolychaete - calcerous tube dwelling Both H/M/LPolychaete - calcerous tube dwelling Circalittoral LBarnacles Infralittoral HBarnacles Both H/MAmphipod Both H/M/LAmphipod Circalittoral HShrimp Circalittoral HLobster Circalittoral MHermit crab Both H/M/LCrab Both H/M/LCrab Both H/M/LCrab Both M/LGastropod Infralittoral H/MGastropod Infralittoral HGastropod Both H/M/LGastropod Both H/M/LGastropod - predatory Both H/MNudibranch Circalittoral H/MBivalve - attached Both H/MBivalve - attached Circalittoral MBivalve - attached Circalittoral M/LBivalve - boring Both H/MBivalve - boring Circalittoral LBivalve - attached Circalittoral LBivalve - attached Circalittoral H/LBryozoan - Cyclostomatous Circalittoral HBryozoan - Ctenostomatous Both H/MBryozoan - Cheilostomatous Infralittoral H/MBryozoan - Cheilostomatous Both H/MBryozoan - upright Circalittoral H/MBryozoan - upright Circalittoral HBryozoan - upright Circalittoral H/MBryozoan - upright Both H/MBryozoan - upright Circalittoral HBryozoan - upright Circalittoral H/MBryozoan - Cheilostomatous Circalittoral HBryozoan - Cheilostomatous Circalittoral H/M/LBryozoan - Cheilostomatous Circalittoral H/MFeather star Both H/M/LFeather star Circalittoral H/M/LStarfish Circalittoral H/MStarfish Both H/M/LStarfish Both H/M/LStarfish Both H/M/LStarfish Both H/M/LBrittle star Both H/M/L

Brittle star Both M/LUrchin Both H/M/LHolothurian Circalittoral H/MAscidian - colonial Both H/M/LAscidian - colonial Circalittoral H/MAscidian - colonial Both H/MAscidian - colonial Both M/LAscidian - solitary Both M/LAscidian - solitary Circalittoral H/LAscidian - solitary Infralittoral M/LAscidian - solitary Infralittoral MAscidian - solitary Both H/M/LAscidian - solitary Circalittoral LAscidian - colonial Both H/M/LAscidian - solitary Both H/M

Indicative Group Species 1 Species 2Red algae - filamentous Ahnfeltia plicataRed algae - calcerous Corallina officinalisRed algae - foliose Chondrus crispus Palmaria palmataBrown algae - filamentous Desmarestia aculeataKelp Laminaria hyperborea Laminaria ochroleucaBrown algae - branched Halidrys siliquosa Sargassum muticumGreen algae - foliose Ulva lactucaGreen algae - filamentous Cladophora rupestrisSponge - encrusting Halichondria panicea Myxilla incrustansSponge - erect Suberites carnosus Axinella dissimilisSponge - boring Cliona celataSoft coral Alcyonium digitatum Swiftia pallidaHard coral Caryophyllia smithiiHydroid Sertularia argentea Kirchenpaueria pinnataAnenome Urticina felina Phellia gausapataAnenome - burrowing Cerianthus lloydiiAnenome - colonial Epizoanthus couchiiPolychaete - scaleworm HarmothoePolychaete PolydoraPolychaete - tube building Sabella pavonina Lanice conchilegaPolychaete - calcerous tube dwelling Pomatoceros triquter Protula tubulariaBarnacles Balanus crenatusAmphipod Dyopedos porrectusShrimp Pandalus montaguiLobster Homarus gammarusHermit crab Pagurus bernhardusCrab Cancer pagurus Necora puberGastropod Gibbula cineraria Margarites helicinusGastropod - predatory Nucella lapillusNudibranch Janolus cristatusBivalve/bivalve like - attached Mytilus edulis Neocrania anomalaBivalve - boring Pholas dactylusFeather star Antedon bifida Antedon petasusStarfish Asterias rubens Luidia ciliarisBrittle star Ophiothrix fragilisUrchin Echinus esculentusHolothurian Holothuria forskaliAscidian - colonial Clavelina lepadiformis Polyclinum aurantiumAscidian - solitary Ascidia mentula Ascidiella aspersaBryozoan - Cyclostomatous Crisia eburneaBryozoan - Ctenostomatous Alcyonidium diaphanumBryozoan - Cheilostomatous Electra pilosa Membranipora membranaceaBryozoan - upright Flustra folicea Bugula plumosa

Project Species:Ahnfeltia plicataAlaria esculentaAlcyonidium diaphanumAlcyonium digitatumAnemonia viridis

Antedon bifidaAntedon petasusAscidia mentulaAscidiella aspersaAsterias rubensAxinella dissimilisBalanus crenatusBugula plumosaCancer pagurusCaryophyllia smithiiCerianthus lloydiiChondrus crispusCiona intestinalisCladophora rupestrisClavelina lepadiformisCliona celataCorallina officinalisCorella parallelogrammaCrisia eburneaDelesseria sanguineaDesmarestia aculeataDexamine spinosaDyopedos porrectusDysidea fragilisEchinus esculentusElectra pilosaEpizoanthus couchiiEunicella verrucosaFlustra foliceaGibbula cinerariaHalichondria paniceaHaliclona viscosaHalidrys siliquosaHarmothoeHolothuria forskaliHomarus gammarusJanolus cristatusKirchenpaueria pinnataLaminaria hyperboreaLaminaria ochroleucaLanice conchilegaLuidia ciliarisMargarites helicinusMembranipora membranaceaMetridium senileMytilus edulisMyxilla incrustansNecora puberNemertesia ramosaNeocrania anomalaNucella lapillus

Obelia geniculataOphiothrix fragilisPagurus bernhardusPalmaria palmataPandalus montaguiParasmittina trispinosaPhellia gausapataPholas dactylusPolyclinum aurantiumPolydoraPolyides rotundusProtula tubulariaSabella pavoninaSargassum muticumSertularia argenteaSpirobranchus triquiterSuberites carnosusSwiftia pallidaUlva lactucaUrticina felina

Species 3 Species 4

Delesseria sanguinea

Alaria esculenta

Dysidea fragilisHaliclona viscosa

Eunicella verrucosa

Obelia geniculata Nemertesia ramosaAnemonia viridis Metridium senile

Corella parallelogramma

Parasmittina trispinosa

Species Trait Included in ProjectGrowth form YESBioturbator? NOFragility NOHeight NOAdult dispersal potential NOSociability NOToxic/poisonous NOFlexibility NOGrowth rate NODependency NODistribution YESBiogeographic range NOMigratory NOBiological zone NOWave exposure preferences NOReproduction 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 YESKey prey taxa YESTemporal variable population YESConnectivity to other habitats/species YES

RationaleFundamental trait will inform modelNot relevant to rock habitatNot relevant to ecological model when considering unimpacted environmentNot relevant to ecological model - factor covered by other traitsTrait considered above scope of ecological modelsNot relevant to ecological model - no bearing on ecological function/driving factorsNot relevant to ecological model - no bearing on ecological function/driving factorsNot relevant to ecological model - no bearing on ecological function/driving factorsNot relevant to ecological modelNot relevant to ecological model/covered by other traitsDistribution may influence spatial variation within model.Covered by other traitsNot relevant to ecological modelWill be either circalittoral or infralittoralEnergy preference will be covered by tidal stream preferenceNot 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 modelTrait considered above scope of ecological modelsTrait considered above scope of ecological modelsFundamental trait will inform modelWill inform ecosystem functionWill inform temporal aspects of modelFundamental trait will inform model

Included in Ecological Group DeterminationNONONONONONONONONONONONONONONONONONONONONONONONONONONONONONONONONONO

RationaleCovered by other traitsNot relevant in rock habitatTrait has no bearing on ecological groups within scope of projectCovered by other traitsTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectCovered by other traitsCovered by other traitsTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of projectTrait has no bearing on ecological groups within scope of project

SourceMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMarLINMESLMESLMESLMESL

Species (Biotope name) Species (WORMS name)

Ahnfeltia plicata Ahnfeltia plicata

Ahnfeltia plicata

Alaria esculentaAlaria esculenta

Alaria esculenta

Alaria esculenta

Alaria esculenta

Alaria esculentaAlcyonidium diaphanum Alcyonidium diaphanumAlcyonidium diaphanum

Alcyonidium diaphanum



Alcyonidium diaphanumAlcyonidium diaphanumAlcyonidium diaphanum

Alcyonium digitatumAlcyonium digitatum

Alcyonium digitatum

Alcyonium digitatum

Alcyonium digitatumAlcyonium digitatum

Anemonia viridisAnemonia viridis

Anemonia viridis

Anemonia viridis

Anemonia viridis

Anemonia viridis

Antedon bifidaAntedon bifida

Antedon bifida

Antedon bifida

Antedon petasus Antedon petasusAntedon petasusAntedon petasus

Antedon petasus

Antedon petasus

Antedon petasusAscidia mentula Ascidia mentulaAscidia mentula

Ascidia mentula

Ascidia mentula

Ascidia mentulaAscidia mentulaAscidia mentula

Ascidia mentula

Ascidia mentula

Ascidiella aspersaAscidiella aspersa

Ascidiella aspersa

Ascidiella aspersaAscidiella aspersa

Asterias rubensAsterias rubens

Asterias rubens

Asterias rubens

Asterias rubens

Axinella dissimilisAxinella dissimilis

Axinella dissimilis

Axinella dissimilis

Balanus crenatusBalanus cretanus

Balanus crenatus

Balanus crenatus

Balanus crenatusBalanus crenatusBugula plumosa Bugula plumosaBugula plumosaBugula plumosaBugula plumosaCancer pagurus Cancer pagurusCancer pagurus

Cancer pagurus

Caryophyllia smithii Caryophyllia smithiiCaryophyllia smithii

Caryophyllia smithii

Cerianthus lloydiiCerianthus lloydii

Cerianthus lloydii

Cerianthus lloydii

Cerianthus lloydii

Chondrus crispus Chondrus crispus

Chondrus crispus

Ciona intestinalisCiona intestinalis

Ciona intestinalis

Ciona intestinalis

Ciona intestinalis

Cladophora rupestrisCladophora rupestris

Cladophora rupestris

Clavelina lepadiformisClavelina lepadiformis

Clavelina lepadiformis



Cliona celataCliona celata

Cliona celata

Corallina officinalisCorallina officinalis

Corallina officinalis

Corella parallelogramma Corella parallelogrammaCorella parallelogrammaCorella parallelogramma

Corella parallelogramma

Corella parallelogrammaCrisia eburnea Crisia eburneaCrisia eburneaCrisia eburnea

Crisia eburnea

Crisia eburnea

-

Delesseria sanguineaDelesseria sanguinea

Delesseria sanguinea

Desmarestia aculeata Desmarestia aculeataDesmarestia aculeata

Desmarestia aculeata

Desmarestia aculeata

Dexamine spinosa Dexamine spinosaDexamine spinosaDexamine spinosaDexamine spinosa

Dexamine spinosaDexamine spinosaDyopedos porrectus Dyopedos porrectusDyopedos porrectusDyopedos porrectusDyopedos porrectus

Dyopedos porrectus

Dysidea fragilis Dysidea fragilisDysidea fragilisDysidea fragilis

Dysidea fragilis

Dysidea fragilisDysidea fragilisDysidea fragilisDysidea fragilis

Echinus esculentusEchinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentus

Echinus esculentusEchinus esculentus

Electra pilosaElectra pilosa

Electra pilosaElectra pilosa

Electra pilosa

Epizoanthus couchiiEpizoanthus couchii


Epizoanthus couchii


Eunicella verrucosaEunicella verrucosa

Eunicella verrucosa

Eunicella verrucosa

Flustra foliacea Flustra foliaceaFlustra foliaceaFlustra foliacea

Flustra foliacea

Flustra foliaceaGibbula cineraria Gibbula cinerariaGibbula cinerariaGibbula cinerariaGibbula cinerariaGibbula cineraria

Gibbula cineraria

Gibbula cinerariaGibbula cineraria

Halichondria paniceaHalichondria panicea

Halichondria panicea

Haliclona viscosa

Haliclona (Rhizoniera) viscosaHaliclona viscosaHaliclona viscosa

Haliclona viscosaHaliclona viscosa

Halidrys siliquosaHalidrys siliquosa

Halidrys siliquosa

Harmothoe HarmothoeHarmothoeHarmothoeHarmothoeHolothuria forskali Holothuria forskali

Holothuria forskali

Holothuria forskali

Holothuria forskaliHolothuria forskaliHolothuria forskali

Homarus gammarusHomarus gammarus

Homarus gammarus

Homarus gammarus

Janolus cristatus Janolus cristatusJanolus cristatusJanolus cristatus

Kirchenpaueria pinnataKirchenpaueria pinnata

Kirchenpaueria pinnataKirchenpaueria pinnata

Kirchenpaueria pinnata

Laminaria hyperborea Laminaria hyperborea

Laminaria hyperborea

Laminaria hyperboreaLaminaria ochroleuca Laminaria ochroleucaLaminaria ochroleucaLaminaria ochroleuca

Lanice conchilega Lanice conchilegaLanice conchilega

Lanice conchilegaLanice conchilega

Lanice conchilega

Lanice conchilegaLanice conchilegaLuidia ciliaris Luidia ciliarisLuidia ciliarisMargarites helicinus Margarites helicinusMargarites helicinusMargarites helicinusMargarites helicinusMargarites helicinusMembranipora membranacea Membranipora membranaceaMembranipora membranaceaMembranipora membranacea

Membranipora membranaceaMembranipora membranaceaMembranipora membranaceaMetridium senile Metridium senileMetridium senile

Metridium senile

Mytilus edulis Mytilus edulisMytilus EdulisMytilus EdulisMytilus edulis

Mytilus edulis

Mytilus edulisMyxilla incrustans Myxilla incrustans

Myxilla incrustansMyxilla incrustans

Necora puber Necora puberNecora puber

Nemertesia ramosa Nemertesia ramosa

Nemertesia ramosa

Neocrania anomala Novocrania anomala

Neocrania anomala

Neocrania anomala

Nucella lapillus Nucella lapillusNucella lapillus

Nucella lapillus

Obelia geniculata Obelia geniculataObelia geniculataObelia geniculataObelia geniculataObelia geniculata

Obelia geniculataOphiothrix fragilis Ophiothrix fragilis

Ophiothrix fragilisOphiothrix fragilis

Ophiothrix fragilis

Ophiothrix fragilis

Pagurus bernhardus Pagurus bernhardus

Pagurus bernhardusPagurus bernhardusPagurus bernhardusPagurus bernhardusPagurus bernhardus

Palmaria palmata Palmaria palmataPalmaria palmata

Pandalus montagui Pandalus montaguiPandalus montagui

Pandalus montaguiPandalus montaguiPandalus montaguiPandalus montaguiParasmittina trispinosa Parasmittina trispinosaParasmittina trispinosa

Parasmittina trispinosaParasmittina trispinosa

Phellia gausapata Phellia gausapataPhellia gausapataPhellia gausapataPhellia gausapataPholas dactylus Pholas dactylus

Pholas dactylusPholas dactylus

Polyclinum aurantium Polyclinum aurantiumPolyclinum aurantiumPolyclinum aurantiumPolyclinum aurantiumPolyclinum aurantiumPolyclinum aurantiumPolydora PolydoraPolydora

Polydora

PolydoraPolyides rotundus Polyides rotundusPolyides rotundusProtula tubularia Protula tubulariaProtula tubulariaProtula tubulariaProtula tubularia

Protula tubulariaProtula tubulariaProtula tubularia

Protula tubulariaSabella pavonina Sabella pavonina

Sabella pavoninaSabella pavoninaSargassum muticum Sargassum muticum

Sargassum muticumSargassum muticumSargassum muticum

Sertularia argentea Sertularia argentea

Sertularia argenteaSertularia argenteaSertularia argentea

Sertularia argenteaSertularia argenteaSertularia argenteaSertularia argentea

Spirobranchus triquiter Spirobranchus triquiterSpirobranchus triquiter

Spirobranchus triquiterSpirobranchus triquiter

Spirobranchus triquiter

Spirobranchus triquiter

Suberites carnosus Suberites carnosus

Suberites carnosusSuberites carnosusSuberites carnosus

Suberites carnosus

Swiftia pallida Swiftia pallida

Swiftia pallida

Swiftia pallidaSwiftia pallidaSwiftia pallida

Ulva lactuca Ulva lactucaUlva lactucaUlva lactucaUrticina felina Urticina felina

Urticina felina

Species Confusion Growth form

Foliose;Turf;Crustose soft

Straplike / Ribbonlike;Foliose;Forest

Alcyonidum gelatinosum Crustose soft;Shrub;Dendroid

Digitate

Radially symmetrical

Pinnate;Radial

Pinnate;Radial

Gelatinous Cylindrical

Cylindrical

Radial;Stellate

Flabellate;Digitate

Conical

TurfSpiral Plumes

Articulate

Cylindrical; Radial

Vermiform unsegmented;Cylindrical

Tur

Cylindrical

Tall, slim, cylindrical

Shrub

Cylindrical

Bivalved

Articulate; Crustose hard; Pinnate;Turf

Cylindrical; somewhat more rectangular in outline and laterally flattened.

Dendroid

Turf;Foliose

Arborescent / Arbuscular

Articulate

Articulate

Crustose soft;Massive;Cushion

Globose

Crustose hard

Polyp

Arborescent / Arbuscular

Turf;Foliose;Crustose hard

Conical

Cushion

Foliose

Vermiform segmented

Cylindrical

Articulate

Oval, Flattened

Pinnate

Variable depending on microhabitat, but typically from spreading cushions with mamillate oscular chimneys, to massive lobose forms with tall, rugose, volcanolike oscular chimneys.

Forest;Straplike / Ribbonlike;Foliose

Forest, Foliose

Tubicolous;Vermiform segmented

Tubicolous

Radial;Stellate

Turbinate

Encrusting on alge, lacyEncrusting Mesh

Encrusting Lacy Mat

Radial);Globose

Bivalved

Cushion

Articulate

Pinnate

Bivalved

Bivalved

Turbinate

Pinnate: Zig/Zag unbranched stems arising from a basal rhizome

Radial;Stellate

Articulate

Crustose soft;Foliose

Articulate

Flat, Nodular

Radial; Cylindrical

Bivalved

cushion

Tubicolous;Vermiform segmented

Veriform, Segmented, Tubicolous

Dichotomous

Tubicolous; Vermiform segmented

Tubiculous; Segmented

Tubiculous

Pinnate: Regularly alternating branches which are long and stringy

Arborescent / ArbuscularPomatoceros triquiter Vermiform segmented;Tubicolous

Tubicolous

Crustose Soft, Cushion

Variable; typically spheroid and anchored to the substratum by a short stalk. It may also be encrusting or massive and lobed.

In the British Isles it typically assumes a hollow contractile, massive figshape, attached by a stalk to the substrate.

Arborescent / Arbuscular flabellate

Blade Shape Highly Variable

Globose

Mobility/movement

Permanent attachment




Permanent Attachment


Crawler

Crawl and Swim. Spend most of their time temprarily attached.

Temporary Attachment

Temporary Attachment

Attached

Attached

Crawler


Rather than being attached at its base like most sea squirts, it is joined on by a part of one side.


Permanent attachmentAttachedCrawler


Attached

Burrower


Temporary attachment; Permanent Attachment


AttachedAttached

Attached

Attached



Crawler

Crawler, Swimmer


Crawler


Encrusting, Attached

Crawler


AttachedEncrusting, Attached


CrawlerCrawler

Crawler

Crawler, can also swim backwards rapidly. Crawler

Attached



Swimmer;Crawler;Burrower

Crawler;Burrower;Swimmer

Crawler



Temporary attachment

Temporary attachment;Permanent attachment


Swimmer;Crawler




Crawler


Attached

Crawler

Crawler

Attached

Crawler

Encrusting, Attached

Attached

Burrower


AttachedBurrower

Burrower

AttachedAttached

Attached

Attached

Attached

Attached





Attached


Attached

Temporary attachment;Crawler

Typical food types

Phytoautotroph

Phytoautotroph

Suspended organic particulates and plankton

Phytoplankton & Zooplankton

Fish; Mollusks; Zooplankton; Phytoplankton; detritus

Particulate matter such as detritus and plankton.

Plankton and detritus


Detritus

Bivalves, polychaetes, small crustaceans, other echinoderms and carrion.

Can be fearsome predators of bivalve molluscs such as mussels and clams.

Suspended particulate matter

Zooplankton and other organic particles of a suitable size, such as detritus and phytoplankton.


A variety of live molluscs and crustaceans as well as carrion.Molluscs, e.g. mussels and whelks

Zooplankton, organic particulates

Zooplankton

Phytoautotroph

Seston

Phytoautotroph

Suspended detritus and plankton

Bacteria, phytoplankton, detritus, and dissolved organic matter (DOM).

Phytoautotroph

Phytoplankton and organic detritus

Small microorganisms, including diatoms and other unicellular algae.

Phytoautotroph

Phytoautotroph

Seston, detritus

Seston, faecal pellets

Fine suspended organic matter.

Phytoplankton, algal spores.

Dead organic material, mixed diet including plant and animal material, plankton and detritus.

Recorded feeding on; worms, barnacles (e.g. Balanus spp.), hydroids, tunicates, bryozoans (e.g. Membranipora spp.), macroalgae (e.g. Laminaria spp.), bottom material and detritus (reviewed by Lawrence 1975).

Suspended organic particles and zooplankton

Phytoplankton, detritus and dissolved organic matter.

Seaweeds, detritus

Phytoplankton

Detritus, plankton, bacteria

Phytoautotroph

Small invertebrates, seagrass, diatoms

Silty deposits

Unselective

A variety of live molluscs and crustaceans as well as carrion.

Variety of bottomliving animals alive or dead. Erect tufted bryozoans such as Bugula and Cellaria


Phytoautotroph

Phytoautotroph

Detritus

Echinoderms, polychaetes, molluscs, crustaceans and probably carrionMainly on other Echinoderms such as brittle stars, starfish and urchins.

Kelp Fronds


Zooplankton but also larger prey. (See additional information.)

Bacteria, phytoplankton, detritus, and dissolved organic matter (DOM).

Detritus

Fish, prawns, other crabs. All sorts of bottomliving animals including worms and molluscs. Some populations have been found to eat large quantities of seaweed.


Seston

seston

Barnacles and musselsMulluscs and small invertebrates

Invertebrates such as crustaceans and molluscs and also carrion or even brown algae. More likely to be predatorial at greater depths.


PhytoplanktonPOM, POC, PON

Organic detritus, carrion

Phytoautotroph

Detritus


Small Organisms

Suspended organic particles

Suspended organic particles

Detritus

detritus

PhytoautotrophPhytoautotroph

Detritus, plankton, bacteria


Phytoautotroph

PhytoplanktonSuspended organic particles and zooplankton



Suspended matter including plankton

Phytoautotroph

Shore crab is its ordinary prey, but it feeds on limpets and other mollusca and nereids, schrimps and Echinus miliaris.

Feeding method

Photoautotroph

Photoautotroph

Active suspension feeder

Active suspension feeder; Predator

Predator

Passive suspension feeder


Siphon, filter feeder

Siphon, filter feeder

Filter Feeder

Scavenger;Predator

Predator


Passive suspension feeder;Active suspension feeder

Passive Suspension Feeder / Active Suspension Feeder

Suspension feeder

Predator

Passive suspension feeder;Predator

Passive suspension feeder;Predator

Photoautotroph


Photoautotroph


Passive suspension feeder; Active suspension feeder

Photoautotroph


Filter FeederSiphon, filter feeder

Suspension feeder; active

Photoautotroph

Photoautotroph

Deposit Feeder, Grazer

Active suspension feeder; Mucus Threads



Omnivore, predator, scavenger

Grazer (grains/particles);Grazer (fronds/blades);Grazer (surface/substratum)

Grazer


Filter Feeder

Photoautotroph

Predator/Scavenger; Raptorial feeding; active carnivores

Deposit feeder

Scavenger;Omnivore

Grazer

Filter feeder

Photoautotroph

Photoautotroph

Passive suspension feeder;Active suspension feeder;Surface deposit feeder;Subsurface deposit feeder

Predator;Scavenger

Deposit Feeder, GrazerFilter feeder

Filter Feeder




Filter Feeder

Omnivore;Herbivore;Predator;ScavengerFish, prawns, other crabs. All sorts of bottomliving animals including worms and molluscs. Some populations have been found to eat large quantities of seaweed.




Predator;Scavenger Active Carnivore

Filter Feeder

Suspension feeder

Passive suspension feeder;Active suspension feeder;Surface deposit feeder;Subsurface deposit feeder

Arms are raised into the passing current where tube feet filters out suspended food which is then passed along the arm to the central mouth.

Surface deposit feeder;Predator;Active suspension feeder

Photoautotroph

Scavenger

Filter Feeder

Active Suspension Feeder


Suspension feederSuspension feeder

Active suspension feeder;Surface deposit feeder

Active suspension feeder;Surface deposit feeder

Photoautotroph

Filter Feeder

Filter Feeder

Photoautotroph

Filter Feeder


Active suspension feederFilter Feeder

Passive Suspension Feeder

Photoautotroph

Predator

Environmental position Habitat

Epilithic Attached

Epifloral Attached

Epifaunal;Epilithic Attached

Colonial

Epifaunal; Epilithic Erect

AttachedEpifaunal; Epilithic

Attached; Sedentry

Epibenthic;Epifaunal;Epilithic Free Living

AttachedEpifaunal

attached

Epifaunal

Attached

Epibethic Free Living

Epibenthic;Epilithic Erect

Epifaunal;Epilithic Attached

EpibenthicAttached

Epibenthic Free living

Epifaunal Attached

Infaunal Tubiculous

Epilithic Attached

Epifaunal Attached

Epilithic;Epiphytic Attached

Epibenthic;Epifaunal;Epilithic Attached

Epifaunal; Infaunal; Epibenthic; Epilithic Attached

Epifloral Attached

Solitary

Epifaunal; epibenthicAttached

Epiphytic; Epilithic attached

Attached

Epifloral Attached

Epifloral Attached

Epifaunal

epibenthic; Epifaunal Free living

Free Living

Epibenthic; EpifaunalEpilithic AttachedEpilithic

Encrusting

Epifaunal Free living

Epifaunal;Epilithic;Epiphytic;Epizoic;Epibenthic Attached

Colonial, Attached

epibenthic; Epifaunal attached

Epibenthic;Epifaunal Attached

Colonial

Epibenthic;Epifaunal;Epilithic Attached

Free living

Epibenthic, Epifaunal, Epizoic

Free Living

Epiphytic; Epilithic AttachedEncrusting

AttachedEpilithic; Epifaunal; epibenthic

Epilithic;Epifloral Attached


Free Living

Epibenthic; Epifaunal

Demersal;Epifaunal;Epibenthic Free living

Epifaunal Free Living

Colonial, AttachedEpilithic;Epifaunal

Colonial, Attached



Infaunal Tubiculous

Infaunal Tubiculous

Epifaunal;Infaunal Free living

Epibenthc; Epifaunal; Epilithic free living

Colonial, Attachedcolonial

Epiphytic colonial

Epilithic;Epifaunal Erect

Epilithic Attached

Epilithic Attached

Epifaunal;Epibenthic Free living

Epifaunal;Epibenthic Attached

Epifaunal Attached

Epifaunal attached

Epifaunal Free living

Attached

Epiphytic Colonial


Epiphytic usually growing on the surface of kelp such as laminaria hyperborea

Epifaunal;Epibenthic Free living

Epifaunal

Epilithic;Epiphytic Attached


Epifaunal/Epibenthic Attached, colonial

Epifaunal/Epibenthic Attached, individual

Lithotomous Burrow dwelling

Epibenthic/Epifaunal ColonialEpibenthic;Epilithic Tubiculous

Epibenthic; epilithic

Epilithic attachedEpilithic attached

Epibenthic attached

attached

Infaunal Attached

Epilithic Attached

Epilithic attached

Epifaunal

AttachedEpibenthic;Epifaunal;Epilithic

Tubiculous

Epibenthic/Epifaunal Attachedepilithic

Epibenthic/Epifaunal Colonial

Epilithic Attached

Epibenthic;Epifaunal Attached

Size

Medium(1120 cm)

Large(>50cm)

MediumLarge (2150cm)

Can be up to 30cm

Medium (1120cm)

up to 20cm diameter, 8cm height

Medium (1120cm)

Each Arm 710cm length

70mm across, 180mm tentacle span

Smallmedium (10cm)510cm

130 mm

Mediumlarge(2150cm)

Medium(1120 cm)

Small(12cm)

Tufts up to 8cm height

Mediumlarge(2150cm)

Small(12cm)

Medium(1120 cm)

Mediumlarge(2150cm)

Medium(1120 cm)

Medium(1120 cm)

Small(12cm)

Medium (1120cm)

Medium(1120 cm)

Smallmedium(310cm)

Small Medium (35cm)Small(12cm)

Mediumlarge(2150cm)

Large (1m+)Small(12cm)

Small(12cm)Very small(<1cm)

6.5mm

Medium(1120 cm)

Individual zooids measure 0.1 X 0.25 mm on average

Medium(1120 cm)

Smallmedium(310cm)

Variable, up to 12mm

Mediumlarge(2150cm)

Medium(1120 cm)

Small (12cm)

Small(12cm)

Medium(1120 cm)

Large(>50cm)

Smallmedium(310cm)

MediumLarge (25cm)

Mediumlarge(2150cm)

SmallMedium (7.5cm)

7080mm height, 15mm width

Spread of up to 30cm, and height to 5cm.

typically 36cm. May be up to 10cm.

Large(>50cm)

Large, up to 2m

1.5m

Mediumlarge(2150cm)

Large(>50cm)

Small(12cm)

68mm1030cm

Mediumlarge(2150cm)

Medium(1120 cm)

20cm width. 5cm Height

Smallmedium(310cm)

Medium(1120 cm)

Small(12cm)

0.2315mm

Smallmedium(310cm)

37cm

0.512cm

Medium(1120 cm)

Smallmedium(310cm)

Large(>50cm)

up to 10cm, generally 45cm.

up to 20cm

Smallmedium (6cm)

50mm tall, 30mm across

Medium(1120 cm)

30mm acrossSmallmedium(310cm)

up to 30mm long

20cm20cmVery small(<1cm)

0.30.5cm

300mm L, 4mm W.

300mm Long

Over 2m

Frond over 1m

Medium (1120cm)

Small(12cm)

150mm

A few cm high and across

720cm

30cm

up to 100cm

Medium(1120 cm)

Distribution

Occurs on all coasts of Britain and Ireland. There is a paucity of records from south east England, reflecting a lack of suitable substrata.

Found on all British and irish CoastsAll UK Coasts

Abundantly distributed in Brtish Isles. Dense populations in the Greater Thames estuary and Northern Bristol Channel.

All coasts of UK. Particularly abundant in the Southern North Sea (Thames estuary)

Found on all British and irish Coasts

Found around the Shetland Isles, Orkney and east coast of Scotland, south to Flamborough Head in England. Its distribution continues along the south west of England and the west coasts of England, Wales, Scotland and Ireland including the Isle of Man.

Southern and Western Coasts of Great Britain

Britain and Ireland; excluding SE coast of england.

Common around the british coast, except the south east

Generally a northern species in Britain and Ireland. Common in the northern part of the Irish Sea and inshore in the west coast of Scotland.

Present on the western coasts of Britain from Portsmouth all around to the west coast of Scotland where it extends northwards to just south of Cape Wrath. Absent from the east basin of the Irish Sea.

Generally distributed around British Isles; else where from wester Norway southwards to the mediterranean and black sea. Generally distributed around the british isles

Common on all British coasts but appears to be more common on south and west coasts


Present at a few sites around Mull, the south west of England and the western extremities of Wales. In Ireland there are records from the south east, the south west, and along the Atlantic coast round to the north east.

Found on all British and irish Coasts, and offshore in the North Sea and Celtic Sea

Extremely common on all UK coastsThroughout the British Isles and Ireland but recorded predominantly in the south


Shetland, NE England, SW England, Wales, Ireland and NW Scotland

Recorded all around the coast of Britain, except on the east coast between the Tees estuary and Harwich. It may be locally abundant in some places

Widely distributed on rocky shores on all British and Irish coasts.

Found on all British and Irish Coasts

Found on all British and Irish Coasts on suitable substrata

Found on all British and Irish Coasts on suitable substrata

Found on all British Coasts


All Bristish Coasts

Recorded from all coasts of the British Isles. However, records from the east coasts are sparse, presumably due to the lack of suitable substrata.

Generally distributed around the UK

All British Coasts

Recorded from Northumberland, North to shetland, from the Clyde Sea, Anglesey, and the Bristol Channel.

This species is widely distributed around the coasts of Britain and Ireland but rare on the north east coast of Scotland.

Common on most coasts of the British Isles but absent from most of east coast of England, the eastern English Channel and some parts of north Wales

All around the coast of the UK

Common on the west coast of Scotland.

Common on all coasts of the British Isles, although under recorded on parts of the east coast

Common around the western and southern coasts of Britain it remains unrecorded in the North sea. Epizoanthus couchii also been recorded in southern Ireland.

Common on all rocky coasts of Britain and Ireland

Common on all british and Irish Coasts

Recorded northwards to north Pembrokeshire and eastwards to Portland Bill in Britain. Common in parts of south Devon and Cornwall and at Lundy. Present on the south and west coasts of Ireland but common only in Galway and Donegal Bays.

All British coasts


Widely distributed and fairly common in the Britain and Ireland

Not ResearchedFound on all coasts of the British Isles

Common on south and west coasts of Britain and Ireland.

South west and west britain.


Recorded from most British and Irish coasts except the southeast, reaching its northern distribution limit around the Shetland Isles

Common around all British and Irish Coasts

Widely distributed throughout british isles

Recently known from W. coast of Ireland; Northern Ireland (Antrim; Strangford Lough); W. coast of Scotland (Mull; Coll; Tiree; Oban; Lewis); S.W. Britain (Salcombe, Devon, Eddystone, Hard Deeps; Scilly Isles; Lundy).

Found on most coasts of Britain and Ireland. Scarce along the south east coast of Britain due to a lack of suitable substrata

South west England, and north to lundy. Absent from ireland.

South west England including Lundy, the Isles of Scilly, south Devon and Cornwall.

Lanice conchilega is found around all coasts of Britain and Ireland.

Found on all British coasts except the southern North Sea.

Northern Species recorded from the shetlands to as far south as Yorkshir and Dublin

All British and Irish Coasts

All coasts of Britain and Ireland

All British and Irish coasts.

Recently known from many sitesa round the british isles.

Very common all around the coast of the British Isles, with large commercial beds in the Wash, Morecambe Bay, Conway Bay and the estuaries of southwest England, north Wales, and west Scotland

Found on all British and Irish coasts.

Widely distributed round all British and Irish coasts.

Common on all rocky coasts of Britain and Ireland

From the Firth of Clyde up the west coast of Scotland including the Hebrides, Shetland, the south coast of England and the Isle of Man. In Ireland along the south coast, the northwest and the northeastFrom the Firth of Clyde up the west coast of Scotland including the Hebrides, Shetland, the south coast of England and the Isle of Man. In Ireland along the south coast, the northwest and the northeast

Common around all British and Irish Coasts

Widespread around the biritsh isles

All British and Irish coasts, except for the east coast of Scotland, around the Humber Estuary and north East Anglia and south Kent coast

Found on all British and Irish coasts

North Sea

Generally distributed throughout Britain and Ireland, but apparently absent from significant stretches of coast in eastern England.

Recorded around the coasts of Britain and East and South coast of Ireland.

Common off all british and irish coasts

NE scotland and NW ireland

S and W coast of England

All Great Britain, most common in southwest

Found on all britsh and irish coastsGenerally distributed around the British Isles

Widespread across the southern and western coasts of britain and ireland

Widely distributed and locally abundant around all british coasts

Pholas dactylus occurs in Britain from Kent along the south and southwest coasts including south Wales, Anglesey and Solway. Also recorded from several sites on the east coasts of Yorkshire and Northumbria and southwest Ireland

All British Coasts

Widespread and common throughout the British Isles

Common in the UK especially in the southern north sea.

Common and widespread on all coasts

Common and widespread on all coasts

Widesprad along atlantic facing coasts of britain and IrelandWidesprad along atlantic facing coasts of britain and Ireland

A common species on all coasts of the British Isles.

South coast, North cornish coast and lundy, Pembrokeshire and the menai strait, isles of man. Clyde and loch ryan in scotland and widely distributed in ireland.

South coast to the Isles of Scilly and along the north Cornish coast to Lundy. Populations have also been recorded in Strangford Lough, Northern Ireland and Loch Ryan and the Firth of Clyde, Scotland.

West coast of scotland and the southwest irish coast

Found on all but the most exposed rocky shores, probably under recorded in Ireland.

Throughout UK

Found on all coasts of the British Isles

Depth range

Lower shore to 22 m

08 m

Extreme Low Water to 100mLow in the intertidal, doesn't say deepest range (offshore)

Extreme low water to 100m

Low Water (Springs) to 50m

Shallow Waters; Bathymetric range extending from extreme low water spring tides to 100m.

up to 20m rare below 1012m

Extreme Low Water Springs to 450m

20 several hundered m

20100m

Present mainly in the intertidal especially in pools from midtide downwards but not extending far into the sublittoral.

Low shore to 200m

Lower intertidal and sublittorally to 80m

Intertidal to depths of 650 m.

0 100+

Insufficient information

From LWN to 200mLittoral and sub littoral down to >50m

050mFrom intertidal down to 100mAll depths

>1000m10100m

To at least 100 m depth.

mid eulittoral, exceptionally to 24 m

Lower shore to at least 500 m

Intertidal to a few metres sublittorally.

Sublittoral to 50 m

0150m

0 18m

0200m

2045

Lower eulittoral to at least 30 m

Lower shore to 15m

rocky substrata on the lower rocky shore to 15m.

Lower intertidal and sublittorally to 60m

15700m

Cryptic lowlittoral and sublittoral

Low shore shallow water to >25m

0 1200 m

040m (Sometimes Deeper)

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

Low shore to 10m

Low water to 100m

mid depth (10m) to deep water.

4m to at least 50m

most common at depths greater than 10m

Sublittoral 1200

Lower shore to 130m

sublittoral to 100m

Intertidal to ca 569 m

Infralittoral to 50m

0 44.5m

Intertidal to 4 m

25000m

Shallow sublittoral to 300m; occasionally on shores at extreme low water

LWST to 60 m.

Shallow sublittoral

Max 40m

Intertidal to about 100m, occasionally up to 350m

Extreme low water to 47m although usually to ca 30m

Low Water to 25m

Intertidal to 1700m

From 4 m to 400 m depth.

From the lower shore to the sublittoral

Lower shore to shallow water

Lower shore to considerable depths.

Intertidal to approx. 5m

Low water down to 400m

Found down to 70 m depth.

10500 m

151500

0 40 m

35230m

Depths higher than 30m abundance was low.

From MTL to 140 m, seldom down to 500 m

Intertidally to 470m, locally abundant to depths of 27m

To a depth of 20m

5230m

shallow sublittoral to continental shelf

0183m

Rocky shore to 15m

32Lower shore to 30m

To 35 m

041mLow shore to 100m

2170mmid shore to 24mMid shore 12m

Lower shore to 100m

>25m

Shallow subtidal and rockpools on the shore. 05m

Shallow waters

subtidal species

up to 70m

Down to 108m

Has been found in Brackish localities. Found in shallow pools on sand and mud flats. Estimates to 100m depth.

1560m

15m

Down to at least 100m

Shallow water distribution limited to west coast of scotland and the kenmare river in SW ireland because of deep water upwellings.

Occurs in deep waters offshore in ireland to a depth of 600m

Substratum preference

Bedrock;Cobbles;Pebbles;Coarse clean sand

Bedrock;Large to very large boulders;Small boulders;Cobbles;Pebbles;Artificial (e.g. metal/wood/concrete)

Bedrock;Large to very large boulders;Small boulders;Cobbles Rocky shores, shelly sands, coarse grounds

Large to very large boulders (2); Small Boulders (3); Bedrock (1); Artificial (e.g. metal/wood/concrete) (39); Caves (46); Overhangs (48); Cobbles (4)

Rocks, Eel grass and Kelp

Algae;Bedrock;Large to very large boulders

Cirri used to anchor the feather star to the seabed, kelp or sponges.

Hard Substratum; wrecks; kelp and sessile animals

Prefers brightly lit, seasweed rich areas of rocky reefs in shallow water. Can also be found in pools on the shore and on kelp fronds or on ell grass strands.

Hard Substratum; wrecks; kelp and sessile animalsRock, Stones, Shells.Vertical or overhanging surfaces, wrecks and rock faces. Many individuals are usually found together in suitable habitats.

Clumps may cover a large area of muddy seabed, with the squirts attached to pebbles or shells within the mud.

Can establish on soft muds but is commonly found associated with hard subtrates, like pier pylons, rocks and docks

Gravel / shingle;Coarse clean sand ;Bedrock

Bedrock;Large to very large boulders

Bedrock;Artificial (e.g. metal/wood/concrete);Cobbles;Gravel / shingle;Large to very large boulders;Pebbles;Small boulders

Stones. Epiphytic on Crustacean and mollusc shellsrocks or other hard surfaces, especially on harbour walls and pier pilings

This species can be found in small groups or as a 'turf' on rocks or other hard surfaces, especially on harbour walls and pier pilings.Cobbles;Gravel / shingle;Pebbles;Sandy mud;Muddy sand;Fine clean sand;Coarse clean sand;Bedrock;Muddy gravelVariety of habitats

Bedrock; Large to very large boulders; Cobbles; Other Species

Rock, also found on a variety of substrata such as sand patches in weak tidal current areas.

Muddy gravel;Coarse clean sand);Fine clean sand ;Sandy mud ;Mud;Mixed;Muddy sand;Gravel / shingle

Bedrock;Large to very large boulders;Small boulders;Rockpools

Other species;Algae;Artificial (e.g. metal/wood/concrete);Bedrock;Large to very large boulders);Small boulders

Bedrock;Small boulders;Cobbles;Algae

Artificial (e.g. metal/wood/concrete);Bedrock;Cobbles;Large to very large boulders ;Overhangs;Small boulders;Pebbles

Soft Rock, Limestone

Artificial (e.g. metal/wood/concrete);Bedrock;Large to very large boulders;Rockpools;Crevices / fissures

Rocks, Shells and Stones

Bedrock;Large to very large boulders; small boulders; Gravel/Shingle; Muddy Gravel;Coarse clean sand; Fine clean sand; Sandy Mud; Muddy Sand; Mud; Mixed; Biogenic Reef; Artifical; Rockpools; Under boulders

seagrasses, drift algae, oyster reef, dock, pilings, breakwaters and manmade debris.

Bedrock;Large to very large boulders;Small boulders;Rockpools

Rocky Substrata

Among laminaria holdfasts, and other algae, in Zostera and on sand or mud.

Bedrock;Large to very large boulders

Rocks, Stones or Seaweed.

Encrusting on rocks and shells.

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

Bedrock;Large to very large boulders;Small boulders;Cobbles;Algae;Other species;Artificial (e.g. metal/wood/concrete);Seagrass;Under boulders;Caves;Overhangs

Artificial (e.g. metal/wood/concrete);Bedrock;Large to very large boulders

Bedrock;Large to very large boulders;Mixed;Small boulders;Cobbles

Rocky shores, on weed and under stones. Sometimes found in rock pools.

sheltered rocky shores, underneath stones and seaweeds

Bedrock;Cobbles;Large to very large boulders;Small boulders

mainly occurs on vertical rock faces, in silt free areasVertical and horizontal rock

Bedrock;Large to very large boulders;Small boulders;Rockpools;Cobbles

Algae;Other species;Gravel / shingle;Crevices / fissures

Rock, rocks and stones, rarely sediment

Rocky sublittoral habitats with considerable wave exposure

Bedrock;Crevices / fissures;Caves;Large to very large boulders

Sheltered rocky coasts

Restricted to hard substrata

Variety of habitats

Found on rock and a wide variety of other substrata including molluscan and crustacean shells, macroalgae, seagrasses and artificial wooden strucutures

Bedrock;Large to very large boulders;Cobbles; Pebbles;Artificial (e.g. metal/wood/concrete)

Bedrock; BouldersRock

Coarse clean sand;Sandy mud;Fine clean sand;Muddy sand

Coarse clean sand;Fine clean sand;Muddy sand;Gravelley sand

under stones, in pools and on fucus and laminaria and smaller algae.

typically found on marine algae e.g. laminaria digitata and laminaria hyperborea

Bedrock;Large to very large boulders;Biogenic reef;Artificial (e.g. metal/wood/concrete;Caves;Overhangs

stones, shells, sand and mud.

Bedrock;Large to very large boulders;Sandy mud; Muddy gravel;Rockpools;Biogenic reef;Artificial (e.g. metal/wood/concrete);Mixed;Muddy sand;Small boulders;Under boulders;Caves;Crevices / fissures

Bedrock;Large to very large boulders;Pebbles;Gravel / shingle;Rockpools;Under boulders very common on rocky and stony seabeds, can also be found on sandy and muddy bottoms.

Bedrock;Large to very large boulders;Small boulders;Cobbles;Pebbles;Gravel / shingle;Maerl

Other species;Bedrock;Large to very large boulders;Small boulders

bedrock, large to very large boulder, other species, small boulders

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

Kelp

Bedrock;Large to very large boulders;Small boulders;Cobbles;Pebbles;Gravel / shingle;Maerl;Muddy gravel;Under boulders;Crevices / fissures ;Other species (see additional information)

Bedrock;Large to very large boulders;Rockpools;Under boulders;Coarse clean sand;Fine clean sand;Gravelley sand

Bedrock ;Algae;Large to very large boulders

All Sediment Types

Shell and Rock, usually on vertical rock or fixed boulder surfaces

Rock faces or in pools and clefts

typically occurs on rocks in the kelp zone

Bedrock;Artificial (e.g. metal/wood/concrete)

Hard substrataBedrock;Mud

Rock

Stones and rocks

Found on stones in sand and mud, at and below low water.

Most common habitat is one of soft sediment. Rock and boulders, cobbles and pebbles, sand and gravel. Muddy Sand, Seagrass

Hard substrata

Hard substrata

Sandy bottoms, pebbles, shells or other hard substrate and occasionally on algal stems.

Bedrock;Large to very large boulders;Small boulders

Circalittoral Rock, often vertical surfacesBedrock, cobbles, large to very large boulders, pebbles, small boulders.

found attached to horizontal rock surfaces but is also found in muddy places attached to shells or stones buried in the mud

This is a common species in rocky habitats which are subject to strong water movement from either tidal streams or wave action. It is most abundant in narrows and rapids with tidal streams of more than 3 knots, where it may form large clumps

Artificial (e.g. metal, wood, concrete), Bedrock, Cobbles, Crevices / fissures, Gravel / shingle, Large to very large boulders, Pebbles, Small boulders

flat sloping rock and occasionally boulders

Stones, Rocks, Algae; fully marine and brackish; mostly on the shore but can be subtidal.

Other species (see additional information);Bedrock;Large to very large boulders;Small boulders;Crevices / fissures

Tidal stream preferences

Moderately Strong (13 kn);Weak (<1 kn)

Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn)

The slightest amount of current or tide tends to push the bunches almost flat to the seabed on which they are attached.

Strong (36 kn); Moderately Strong (13kn)

This species has previously been recorded within current exposed rocky channels, including areas subjected to gravel abrasion. Within the entrance of poole harbour A.diaphanum peaks in abundance within the deeper, more current scoured section of the crosschannel profile.

More abundant in areas of moving water

High water flow rate reflected by the presence of characteristic biota Alcyonium digitatum and flustra foliacea.

Stong Current and high wave exposure

Moderately Strong (13 kn)

Feather stars are most nitcable when they form large aggregations; these are most common in current swept areas not exposed to heavy surf.

Preferring positions with a strong current.

Moderate Current


Weak

Low energy habitats like estuaries, harbours and semienclosed embayments to protect them from strong current and tidal forces

Strong (36 kn);Moderately Strong (13 kn)

Very Strong (>6 kn);Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn);Very Weak (negligible)

Moderate/Strong; wave exposed areas where water flow rate is potentially high. The species has an upright branching growth form protruding up in to water column. Large increases in water flow rate may interfere with the posture of the animal, cause physical damage. Reductions in water flow rate may affect feeding efficiency.

Most common in areas with considerable turbidity and sand scourModerately Strong (13 kn);Weak (<1 kn)

Weak

Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn);Very Weak (negligible)

Moderate/Low ; Eleftheriou & Basford (1983) observed Cerianthus lloydii feeding under a wide range of hydrodynamic conditions which showed a great degree of adaptation to the prevailing conditions. Therefore, the species can tolerate some increase in water flow rate however, if water flow increases to strong then Cerianthus lloydii will be unable to feed and if such an increase lasted for a year the species would probably die.

Moderately Strong (13 kn);Weak (<1 kn); Very Weak (negligible)


Strong (36 kn); Moderately Strong (13 kn); Weak (<1kn); Very weak (negligable)

Moderately Strong (13 kn);Weak (<1 kn);Very Weak (negligible)

Seems to like a fair amount of water movement and occurs in places with quite strong currents or swell.

Moderate: the water currents they generate are probably localized so that they are still dependant on water flow to supply adequate food.

Weak/Low; Clavelina lepadiformis thrives in areas where there is very little, if any, water movement (for instance, Abereiddy Quarry, Pembrokeshire (Hiscock & Hoare, 1975)) as it is an active suspension feeder. Naranjo et al. (1996) found that the species was dominant in a low rate of water renewal, excess silting and high suspended solid concentrations.


Strong Currents

Moderate/Strong current

Weak, Very Weak, Moderately StrongStrong and Moderately Strong



Moderate with some sediment scour


Very Strong (>6 kn); Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn)

High water flow rate reflected by the presence of characteristic biota Alcyonium digitatum and flustra foliacea.

Very Strong (>6 kn) ;Strong (36 kn) ;Moderately Strong (13 kn); Weak (<1 kn)

Considerable water movement, e.g. gullies and/or areas exposed to wave action and tidal currents.Strong currents with low turbidity


Strong; Moderately Strong

Moderately Strong, Weak, Very Weak

Weak/Moderate

Thought to be restricted to calm water due to its fragility. Under clean and calm water.

Very Weak, Weak, Moderate, Moderately Strong (Slight to Moderate)

Weak/Moderate ; reduced flow rates may increase the abundance of the grazer Gibbula cineraria as strong water flow rates are likely to dislodge these animals from the blades of the kelp and other plants. Not only will this increase potential damage to the kelp plants and red seaweeds, the grazing activities of this species have also been shown to have a deleterious affect on the postsettlement establishment of larvae of ctenostome and cheilostome bryozoans, hydroids and ascidians (Turner & Todd, 1991).


Strong, Mod strong, Weak Very Weak


Water Velocity needs to be considered when assessing clearance activity of Lanice in an environment.

Weak

Moderate; Moderately Strong: Prolific growth is favoured by fast flowing water which provides food and oxygen.

Very Strong (>6 kn);Strong (36 kn);Moderately Strong (13 kn) Marked preference for areas of stong water flow.


Moderately strong (13 kn), Weak (>1 kn), Very weak (negligible)Moderately strong (13 kn), Weak (>1 kn), Very weak (negligible)


Moderately Strong (13 kn);Weak (<1 kn); Very Weak (negligible)


Very Strong (>6 kn);Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn)

Weak, Moderate, Strong



Strong (36 kn), Moderately strong (13 kn) Weak Moderately strong (13 kn), Weak (>1 kn), Very weak (negligible)

Moderate, Moderately Strong; Decreases in water flow rates are likely to cause a reduction of food availability for feather stars, and may affect growth rates. If this persists over time populations will likely be lost through death or migration to higher flow areas which offer better feeding (Hill, 2008). Effects of increases in water flow rates will depend on the magnitude of change with small increases likely to be tolerated, although feather stars are not able to tolerate very strong currents (>6 knots) and will likely be washed away (Hill, 2008).

Occurs where wave turbulence is strong

Weak to moderately strong

Strong; Moderate; Weak

Weak, Moderate, Strong

weak; moderate/strong

Strong (36 kn), Moderately strong (13 kn), Weak (>1 kn)Weak (>1 kn), Very weak (negligible

Weak (>1 kn), Very weak (negligible)Moderately strong (13 kn), Weak (>1 kn), Very weak (negligible)

Significant tidal currents

Moderately Strong. *Generally Intertidal rock pools on exposed/mod. Exposed shores.


Usually in clean water where currents are not strongWeak (<1kn)

This is a common species in rocky habitats which are subject to strong water movement from either tidal streams or wave action. It is most abundant in narrows and rapids with tidal streams of more than 3 knots, where it may form large clumps

Weak; very sheltered conditions

Very Strong (>6 kn);Strong (36 kn);Moderately Strong (13 kn);Weak (<1 kn)

Stong/ Moderate : Sea fans are found in strong tidal streams but most likely retract their polyps when current velocity gets too high for the polyps to retain food (Hiscock pers. comm.), leading to a reduction in viability. Tidal streams exert a steady pull on the colonies and are therefore likely to detach only very weakly attached colonies

Physiographic preferences

Open coast;Strait / sound;Enclosed coast / Embayment

Open coast;Sealoch

Outer Estuary

Open coast; Offshore Seabed; Enclosed Coast/Embayment

Open Coast; Offshore Seabed

Open Coast

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

Sheltered or moderately exposed habitats


Open Coast, Estuaries, Offshore seabed

Open Coast, Estuaries, Offshore seabed


Open coast; Offshore seabed

Open coast;Offshore seabed;Strait / sound;Sealoch;Ria / Voe;Estuary;Enclosed coast / Embayment

Open coast; Offshore Seabed; Enclosed Coast/Embayment

Offshore seabed;Strait / sound

Offshore Seabed

Offshore Seabed

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

The greratest abundance of C.smithii has been found on vertical rock faces in exposed conditions.

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

Open coast ;Offshore seabed;Strait / sound;Sealoch;Ria / Voe;Estuary;Enclosed coast / Embayment

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

Open Coast; Offshore Seabed;

Open Coast; Offshore Seabed;

Open Coast; Offshore Seabed; Strait/sound;Sealoch; Ria/Voe; Enclosed Coast/Embayment

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

Strait / sound;Ria / Voe;Enclosed coast / Embayment

Open Coast

Open Coast, Offshore seabed,




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

Open coast;Offshore seabed;Strait / sound

Open coast;Offshore seabed;Strait / sound;Sealoch;Ria / Voe

Open Coast; Enclosed Coast/Embayment

Open Coast


Open Coast, Offshore seabed, Estuaries, enclosed coast

Open Coast, Offshore Seabed

Sheltered Coast

Open Coast; Enclosed Coast/Embayment, sea lochs

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

Enclosed coast / Embayment (9);Offshore seabed (2);Strait / sound (3);Open coast (1)

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

Moderately exposed sitesOpen Coast, Enclosed coast embayment

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

Open coast;Offshore seabed

Open coast

Open Coast

Offshore seabed;Strait / sound;Ria / Voe

Open Coast; Wave surged vertical rock; in surge gullies and caves.

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

Open Coast; Wave surged gullies or caves; vertical (and overhanging rock)

Open coast;Offshore seabed;Sealoch

Open coast;Offshore seabed;Sealoch

Open coast;Offshore seabed;Strait / sound;Sealoch;Ria / Voe;Estuary;Enclosed coast / Embayment

Open coast;Offshore seabed;Sealoch;Ria / Voe;Estuary;Enclosed coast / Embayment

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

Conditions of moderate wave exposure

Open coast;Offshore seabed; Strait / sound


Open CoastOffshore SeabedOfshore Seabed


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

Open coast

Very exposed


Open Coast; Offshore SeabedOpen Coast; Offshore Seabed

Sheltered to wave exposed condition

Open CoastOpen Coast; Sea lochs; Enclosed coast;

Open Coast; Wave sheltered fjordic sealochsOpen Coast

Open Coast; Offshore Seabed; Strait/sound; Estuary; Isolated saline water (lagoon); Enclosed coast/Embayment.

Open Coast; Offshore Seabed; Estuaries

Does not grow on intertidal rock as it has a low tolerance to emersionOpen Coast; Enclosed CoastOpen Coast; Enclosed Coast

Open Coast; Estuaries

Open coast;Enclosed coast / Embayment

Open Coast, Offshore seabed,

Offshore seabed

Open Coast; Estuaries

Open coast ;Offshore seabed;Strait / sound;Sealoch;Ria / Voe;Estuary

Salinity preference Lifespan

610 years

Full (3040 psu) 35 years

610 yrs

Full (3040) psu

Variable (1840 psu); Full (3040 psu) 2150 yrs

Full (3040 psu);Variable (1840 psu);Reduced (1830 psu)

33.13835.352

20100 years

Full, Variable

Full (3040 psu)

Full (3040 psu)

Long lived

Full, Variable

Full (3040 psu);Variable (1840 psu) 610 yrs

Full (3040 psu)

Will tolerate salinities as low as 18 PSU and is therefore often found in estuaries

12 yrsFull (3040 psu);Reduced (1830 psu);Variable (1840 psu);Low (<18 psu)

Full (3040 psu)Variable (1840 psu);Full (3040 psu) 2150 years

1120 yearsFull

1120 years

Full

Full (3040 psu);Variable (1840 psu) 610 years

12 years


Full (3040 psu);Reduced (1830 psu);Variable (1840 psu)

Full (3040 psu);Low (<18 psu);Reduced (1830 psu);Variable (1840 psu)

Full (3040 PSU) 51100 years

Variable (1840 psu);Full (3040 psu)

Full

Variable; (27.16536.446PPS)

610 years

Full(35ppt)

Varied (1837)

Variable (1840 psu);Full (3040 psu);Reduced (1830 psu)

Full


Full (3040 psu)

Full (35)



Full (3435)

35 years

Full(35.008 35.363)

Full (3040 psu);Variable (1840 psu)

35 yearsFull, Variable (1739pps)

Full (3035ppt)


50 yearsFull

Variable (1830), Reduced, Full

Variable (1840 psu);Full (3040 psu);Reduced (1830 psu)


Full

Full (3040 psu);Variable (1840 psu)


Full

Full


2150 years

Full (3035ppt)Full (3035ppt)

Reduced (1830 psu);Variable (1840 psu);Full (3040 psu)

Full (3040 psu) Variable (1840 psu) 610 years

Full; Variable. *determined from physiographic pref. <1 year




27.16538.605

610 yearsLow (<18 psu);Full (3040 psu);Variable (1840 psu);Reduced (1830 psu)


Full (3040 psu)

Full (3035 ppt) FullFull

34 years

Full (31.23536.325)

Full (35.33335.339)


FullFull34.67035.363

12 years

Full, Variable, low

tolerant of low salinites

FullFull

Full (3035 ppt), Variable (1835 ppt)

Reduced (1830ppt), Variable (1835 ppt), Low (<18ppt)

Full, Variable

Tolerant of estuarine conditionsFullFull

Tolerant of estuarine conditions

Full, Variable


Full (3040), Variable (1840)

1120 years

Full (3468)

Variable; tolerates brackish communities.

2150 yearsVariable (1840 psu);Reduced (1830 psu);Full (3040 psu); Low (<18 psu)

Sensitivity to change

Some species, such as the kelp Alaria esculenta, were badly impacted by

Low Moderate Sensitivity

High intolerance to substratum loss; Displacement; synthetic compound contamination and hydrocarbon contamination.

Removal of the substratum would also remove the entire population of Ahnfeltia plicata growing on it. Intolerance is therefore assessed as high. Recoverability is recorded as high (see additional information below) but may be delayed if the hydrodynamic regime does not allow a supply of spores from distant populations.

The holdfast of Ahnfeltia plicata is a thin crust which grows epilithically (Dickinson, 1963). It is unlikely that the holdfast, or the encrusting tetrasporophyte phase, would survive removal from the substratum and be able to reattach to a new substratum. Intolerance is therefore assessed as high. Recoverability is recorded as high (see additional information below).

Moderate/Low Sensitivity

High Intolerances to Substratum Loss; Displacement; Chanes in Oxtgenation

Alcyonium digitatum is permanently attached to rocky substratum. Removal of the substratum would also remove this species within the area under consideration and mortality is judged to be high.Recoverability, see additional information below.

Alcyonium digitatum is likely to be highly intolerant of displacement. The species is permanently attached to the substratum and once displaced does not have the ability to reestablish its attachment. Recoverability, see additional information below.

Alcyonium digitatum mainly inhabits environments in which the oxygen concentration usually exceeds 5 ml l1 and respiration is aerobic. Assimilation of oxygen occurs simply by diffusion through the epidermis of exposed tissues and transport to tissues is facilitated by hydroplasmic flow and ciliary activity (Hickson, 1901). It is likely that Alcyonium digitatum would be highly intolerant of a period of hypoxia so an intolerance assessment of high is recorded. Recoverability, see additional information below.

take place and recovery should be possible within five years see additional information.

The species is subtidal, only occurring very rarely at extreme low water springs, so is likely to be intolerant of desiccation. Therefore, an intolerance of high is reported. See additional information for recovery.

The species is subtidal, although some populations are found at very low water springs, so an increase in emergence is likely to result in desiccation to which the species is highly intolerant. Therefore, an intolerance of high is reported. For recovery see additional information.

The species occurs in areas where there is fast current flow suitable for passive suspension feeding. In a series of unpublished experiments by Mr M. Hannan (cited in Hiscock, 1983) Antedon bifida was able to maintain its grip on a concrete substratum in currents up to 90cm/s in a flume. Thus, the clawlike cirri enable the species to survive in moderately strong currents and dense populations are found in areas where water flow is likely to be between 1 and 3 knots during maximal tidal flow. However, if water currents increased to very strong (>6 knots) from moderately strong for a period of a year it is likely that populations would not survive. La Touche (1978) observed that Antedon bifida was unable to maintain its arms in a vertical feeding position in all but the weakest of currents. However, the extremely flexible and opportunistic nature of the species arms and pinnules to adapt in other ways and in several types of water movement. Although dense beds of featherstars are likely to buffer water currents to some degree feeding is likely to be difficult and many individuals are likely to be washed away. Intolerance is therefore recorded as high. For recovery see additional information.

The species occurs in areas where there is fast current flow suitable for passive suspension feeding so a decrease in water flow is likely to reduce food supplies. Thus, growth and fecundity will be impaired and over the period of the year individuals are likely to die or move away to higher flow areas. Thus, intolerance of the species is considered to be high. For recovery see additional information.

Antedon bifida is not found in areas subject to wave exposure and so is likely to be intolerant. If subjected to wave exposure, the species will probably be unable to feed and may be damaged by wave action. Populations are unlikely to survive such a increase for a period of a year so intolerance is reported to be high. On return to normal conditions recovery should be possible within five years

Antedon bifida is likely to be intolerant of abrasion as individuals would probably be killed or damaged by a force equivalent to a scallop dredge dragged across them. The species can regenerate body parts even when most arms and part of the disc have been lost so most damaged individuals are likely to recover. For recovery see additional information below.

Echinoderms are considered to be stenohaline animals that lack the ability to osmo and ionregulate (Stickle & Diehl, 1987). The inability of echinoderms to osmoregulate extracellularly causes body fluid volume to decrease when individuals are transferred to higher external salinity. Over the period of a year, populations are unlikely to survive increased salinity. Echinoderm larvae have a narrow range of salinity tolerance and will develop abnormally and die if exposed to increased salinity. The species is therefore likely to be intolerant of an increase in salinity for a period of a year and an intolerance of high is reported.

Echinoderms are considered to be stenohaline animals that lack the ability to osmo and ionregulate (Stickle & Diehl, 1987). The inability of echinoderms to osmoregulate extracellularly causes body fluid volume to increase when individuals are transferred to lower external salinity probably resulting in the death of animals. Echinoderm larvae have a narrow range of salinity tolerance and will develop abnormally and die if exposed to reduced salinity. The species is therefore likely to be intolerant of a decrease in salinity for a period of a year and an intolerance of high is reported.

Asterias rubens is benthic and although able to migrate by crawling up to a distance of 150200 m per month or 57 m daily (Dare, 1982), it is not considered sufficiently mobile to avoid the physical removal of the substratum to which it is attached and a high intolerance rank is given. Recoverability, see additional information.

Asterias rubens has a large surface area : volume ratio and is relatively soft bodied so if stranded and continually exposed to air and sunshine for one hour would be prone to desiccation. Large numbers of dead starfish have been recorded after displacement and stranding upon the shore presumably from desiccation (see wave exposure) However, when migrating inshore to feed Asterias rubens seeks the damp underside of substrata which offer a degree of protection from direct sunlight and drying wind.

The geographic range of Asterias rubens illustrates that the species is tolerant of a range of temperatures and probably becomes locally adapted. The following observations of the response of Asterias rubens to changes in temperature have been reported: The response of Asterias rubens to prolonged exposure to unusually high temperatures is arm shedding (autotomy) then death (Schäfer, 1972).Starfish have also been found dead in isolated rock pools during prolonged emersion in calm hot weather, the suspected cause of death being increased water temperature (references in Lawrence, 1995).Feeding activity of Asterias rubens may also be disrupted by temperature changes. Feeding activity of Asterias rubens declines during cold winters and hot summers. For instance, Anger et al. (1977), described how turbulence within the water column caused destruction of a thermocline and allowed the temperature of the bottom water to increase to 1618°C causing a decrease in the feeding activity of Asterias rubens. The disruption of feeding activity has implications for recruitment. Nutrient reserves in the pyloric caeca are an important source of energy for the process of gametogenesis and therefore the acquisition of food in the summer preceding spawning is an important factor in determining fecundity (Jangoux & Van Impe, 1977; Oudejans et al. 1979).

Echinoderms are stenohaline owing to the lack of an excretory organ and a poor ability to osmo and ionregulate. The inability of echinoderms to osmoregulate extracellularly causes body fluid volume to increase or decrease when individuals are transferred to lower or higher external salinity respectively, e.g. a sudden inflow of river water into an inshore coastal area caused mass mortality of the conspecific species Asterias vulgaris at Prince Edward Island, Canada (Smith, 1940, in Lawrence, 1995).

Asterias rubens is an aerobic organism and oxygen uptake is by the tube feet and across the body wall. Hypoxic conditions can occur within the habitat of Asterias rubens owing to current changes, thermal stratification of the overlying water column and eutrophication. Theede et al., (1969) undertook experiments on the survival capacity of marine bottom invertebrates in oxygen deficient and hydrogen sulphide (H2S) containing seawater at 10°C. The duration of exposure survived by 50% of Asterias rubens exposed to oxygen deficient seawater (0.15 ml O2l1 = 0.2 mg 02l1) was 84 hours. In the presence of H2S (50 mg Na2S.9H2O L1) the resistance of Asterias rubens to oxygen deficient water dropped to 67 hours, thus it is likely that this species would not survive a week of hypoxic conditions.

High Sensitivity.

High intolerances to substratum loss; dessication; increase in emergence regime; displacement and change in salinity.

The species is permanently attached to the substratum. Substratum loss would result in the entire population. No information is available regarding the reproduction or dispersal abilities of this species so no assessment of recoverability can be made. Most sponges however, tend to be slow growing and long lived.

The species inhabits the lower infralittoral and circalittoral zones and as such is never exposed to the air. Desiccation, although unlikely, would probably result in death. No information is available regarding the reproduction or dispersal abilities of this species so no assessment of recoverability can be made. Most sponges however, tend to be slow growing and long lived.

The species inhabits the lower infralittoral and circalittoral zones and as such is never subject to an emergence regime. Emergence, although unlikely, would probably result in death. No information is available regarding the reproduction or dispersal abilities of this species so no assessment of recoverability can be made. Most sponges however, tend to be slow growing and long lived.

The species is permanently attached to the substratum. It is unlikely to be able to reform this attachment if displaced. No information is available regarding the reproduction or dispersal abilities of this species so no assessment of recoverability can be made. Most sponges however, tend to be slow growing and long lived.

The species inhabits open coasts and offshore areas where salinity is likely to be full. Decreases in salinity will result in conditions outside the preferred range for the species, probably causing death.

and 11 mm diameter can withstand 17 hours and 40 hours of aerial exposure respectively. Similarly, Barnes et al. (1963) recorded that Balanus crenatus had a mean survival time of 14.4 hours in dry air. An increase in the period of desiccation would therefore lead to a depression in the upper limit of the species distribution. A decrease in the period of desiccation could lead to an extension of Balanus crenatus up the shore. The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily colonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore recovery is predicted to be high.

Balanus crenatus is vulnerable to desiccation upon aerial exposure. The shell plates are more permeable than other littoral barnacles, therefore it loses water and dies quicker. Foster (1971) recorded that adults of 6 mm and 11 mm diameter can withstand 17 hours and 40 hours of aerial exposure respectively. An increase in the period of emergence would lead to a depression in the upper limit of the species distribution. A decrease in the period of emersion could lead to an extension of Balanus crenatus up the shore. The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily colonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore, recovery is predicted to be high.

Balanus crenatus is a boreal species, and is intolerant of increases in water temperature. In Queens Dock, Swansea where the water was on average 10 °C higher than average due to the effects of a condenser effluent, Balanus crenatus was replaced by the subtropical barnacle Balanus amphitrite. After the water temperature cooled Balanus crenatus returned (Naylor, 1965). It has a peak rate of cirral beating at 20 °C and all spontaneous activity ceases at about 25 °C (Southward, 1955). The species is more tolerant of lower temperatures. Balanus crenatus was unaffected during the severe winter of 196263, when average temperatures were 5 to 6 °C below normal (Crisp, 1964). The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily colonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore recovery is predicted to be high.

Balanus crenatus is permanently attached to the substratum and could not survive if it was removed. However, the species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily colonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994) so recovery is predicted to be high.

Barnacles have a low resilience to chemicals such as dispersants, dependant on the concentration and type of chemical involved (Holt et al., 1995). They are less intolerant than some species (e.g. Patella vulgata) to dispersants (Southward & Southward, 1978) and Balanus crenatus was the dominant species on pier pilings at a site subject to urban sewage pollution (Jakola & Gulliksen, 1987). Hoare & Hiscock (1974) found that Balanus crenatus survived near to an acidified halogenated effluent discharge where many other species were killed, suggesting a high tolerance to chemical contamination. Little information is available on the impact of endocrine disrupters on adult barnacles. Holt et al. (1995) concluded that barnacles are fairly sensitive to chemical pollution, therefore intolerance is reported as high. The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily recolonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore, recovery is predicted to be high.

Balanus crenatus respires anaerobically so it can withstand some decrease in oxygen levels. When placed in wet nitrogen, where oxygen stress is maximal and desiccation stress is minimal, Balanus crenatus has a mean survival time of 3.2 days (Barnes et al., 1963). It is therefore predicted that the species would not survive low oxygen levels for a week, so intolerance is reported as high. The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily recolonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore recovery is predicted to be high.

Not Sensitive/Low

Tolerant species with some Low intolerances. Only high intolerance was to synthetic compound contamination.

High Intolerances to substratum loss; displacement; synthetic compound contamination.

Chondrus crispus lives permanently attached to the substratum (Dixon & Irvine, 1977) and therefore the entire population would be removed if the substratum were to be lost. Intolerance is therefore recorded as high. Recovery of Chondrus crispus was monitored after a rocky shore was totally denuded by ice scour in Nova Scotia, Canada (Minchinton et al., 1997). Recovery to original biomass was achieved in 5 years (see additional information below). Recoverability is therefore recorded as high.

No information was found concerning displacement of Chondrus crispus. It seems unlikely that the holdfast could remain in situ for long enough on a rocky shore to reestablish a bond with the substratum. Intolerance is therefore assessed as high, though the decision is made with very low confidence. In the absence of holdfasts for regeneration, recovery is likely to take up to 5 years (see additional information below) so recoverability is recorded as high.

semelparous and annual to iteroparous and living 23 years depending on depth and salinity (in Sweden at least). Reproduction (in Plymouth) is recorded as occurring all year round.

Emergent epifauna are thought to be particularly vulnerable to damage from passing fishing gear (Jennings & Kaiser, 1998). Damage to emergent epifauna was the first sign of damage from scallop dredging on horse mussel beds (see Modiolus modiolus) (Service & Magorrian, 1997; Magorrian & Service, 1998; Service 1998). However, while several species of upright hydroids and bryozoans were adversely affected by bottom fishing, some species increased in abundance after fishing disturbance either due to their ability to rapidly colonize space (e.g. Nemertesia sp.) and/or their ability to recover from fragments or budding (e.g. small ascidians, especially Ascidiella spp. and Alcyonium digitatum) (Bradshaw et al., 2000; 2002). Ciona intestinalis is a large ascidian, with a soft, retractile body. Physical disturbance by a passing scallop dredge is likely to cause physical damage and death. Therefore, an intolerance of high has been recorded.

Adults are sessile and so cannot contribute to recovery through active immigration but the species is widespread. Rafting by adults attached to floating objects or shipping may form an important mechanism for recolonization. Dispersal through attachment to ships is believed to be the main reason behind the widespread global distribution. Otherwise, dispersal is mediated by the larval stage. Larval recruitment from other populations may be restricted by the larvae being retained near the adults in mucus threads. Settling time of the larva is quite short usually a few hours so dispersal may be limited. Reproductive frequency and longevity varies from semelparous and annual to iteroparous and living 23 years depending on depth and salinity (in Sweden at least). Reproduction (in Plymouth) is recorded as occurring all year round. Therefore, recoverability has been recorded as high.

Although the species is permanently attached to the substratum, there is some capability for reattachment. However, only after prolonged contact with the substratum and adults are likely to be lost from vertical substrata, or as a result of water flow. The species is widespread. Adults are sessile and so cannot contribute to recovery through active immigration. Local recovery may be facilitated by the retention of larvae in mucus string close to the parent adults. Settling time of the larva is quite short usually a few hours so dispersal may be limited. No information is available regarding the fecundity of this species. Reproductive frequency and longevity varies from semelparous and annual to iteroparous and living 23 years depending on depth and salinity (In Sweden at least). Reproduction (in Plymouth) is recorded as occurring all year round.

Low Sensitivity

High intolerances to substratum loss and displacement.

Cladophora rupestris forms a permanent attachment to substrata, so would be intolerant of substratum loss. Intolerance has been assessed to be high. Recoverability has been assessed to be very high (see additional information below).

Cladophora rupestris forms a permanent attachment to solid substrata. It is likely to be intolerant of displacement as, once removed, mature plants are unable to reattach. Intolerance has been assessed to be high. The species has a considerable ability for recovery. For instance, after the Torrey Canyon tanker oil spill in mid March 1967, recolonization by sporelings of Ulva and Cladophora had occurred by the end of April. Recoverability has been assessed to be very high.


High Intolerances to substratum loss; smothering; dessication; increase in emergence regime; abrasion and physical disturbance and displacement.

Clavelina lepadiformis is permanently attached to the substratum. Removal of substratum will result in loss of the population. Adults are unlikely to be able to reattach, therefore recovery depends upon dispersal of larvae during the breeding season. Intolerance to substratum loss is assessed as high. Recoverability is likely to be moderate (see Additional Information below).

Clavelina lepadiformis reaches up to 20 mm in height and often colonizes vertical surfaces and overhangs. Smothering by 5 cm depth of sediment will completely cover the majority of the population, with only those colonizing overhangs and vertical surfaces not being affected. Therefore, intolerance is assessed as high. Recoverability is likely to be moderate (see Additional Information below).

The sea squirt is a delicate animal, so exposure to desiccating influences for one hour will probably result in death of a proportion of the population. Therefore, intolerance is assessed as high. Recoverability is likely to be moderate (see Additional Information below).

Clavelina lepadiformis is a sublittoral species. Changes in the emergence regime, where the lower shore becomes exposed for longer, are likely to result in a proportion of the population dying. Intolerance to the emergence regime is assessed as high. Recoverability is likely to be moderate (see Additional Information below).

The light bulb tunicate is permanently attached to the substratum and is unable to move out of the way from abrasive objects. The body of the species is soft and delicate, so abrasion is likely to cause physical damage and possibly death. Intolerance is assessed as high. Recoverability is likely to be moderate (see additional information below).

After displacement, adults may not be able to reattach to the substratum. Intolerance is assessed as high. Recoverability is likely to be moderate (see Additional Information section below).

substances within 1 week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh 1977). New fronds of Corallina officinalis appeared on sterilised plots within six months and 10 percent cover was reached with 12 months (Littler & Kauker 1984).

Finely branched fronds or cushionlike turfs may hold water, reducing desiccation stress. Padilla (1984) noted that finely branched Corallina vancouveriensis held more water than coarsely branched or crustose corallines and survived on emergent substrata around tidepools. This effect is less marked in Corallina officinalis (Wiedemann pers. comm.). Corallina officinalis inhabits damp or wet gullies and rock pools and does not inhabit the upper shore, suggesting that it is intolerant of desiccation. Fronds are highly intolerant of desiccation and do not recover from a 15 percent water loss, which might occur within 40 45 minutes during a spring tide in summer (Wiedemann 1994). An abrupt increase in temperature of 10 deg C caused by the hot, dry 'Santa Ana' winds (between January and February) in Santa Cruz, California resulted in die back of several species of algae exposed at low tide (Seapy & Littler, 1984). Although fronds of Corallina spp. dramatically declined, summer regrowth resulted in dense cover by the following October, suggesting that the crustose bases survived. Severe damage was noted in Corallina officinalis as a result of desiccation during unusually hot and sunny weather in summer 1983 (an increase of between 4.8 and 8.5 deg C) (Hawkins & Hartnoll 1985). Hawkins & Hartnoll (1985) found that Corallina officinalis and encrusting corallines often die when their protective canopy of other algal species is removed. Therefore, this species is likely to be highly intolerant of increased desiccation, equivalent to being raised one level on the shore.Corallina officinalis settled on artificial substances within 1 week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh 1977). New fronds of Corallina officinalis appeared on sterilised plots within six months and 10 percent cover was reached with 12 months (Littler & Kauker 1984). In experimental plots, up to 15 percent cover of Corallina officinalis fronds returned within 3 months after removal of fronds and all other epiflora/fauna (Littler & Kauker 1984). Littler & Kauker (1984) suggested that the crustose base was more resistant of desiccation or heating than fronds. Although new bases may recruit and develop quickly the formation of new fronds from these bases and recovery of original cover may take longer, however, the population is likely to recover within 5 years.

the substratum. Norton (1992) noted that algal spore dispersal is probably determined by currents and turbulent deposition. However, red algae produce large numbers of spores that may settle close to the adult especially where currents are reduced by an algal turf or in kelp forests. It is likely that this species could recolonize an area from adjacent populations within a short period of time in ideal conditions but that recolonization from distant populations would probably take longer.

Delesseria sanguinea is normally subtidal and unlikely to be exposed to desiccation except at extreme low tides or in exposed rock pools. Strong insolation of 20mW per square cm (comparable to mid day on a cloudless December in the UK) causes significant reduction in photosynthesis in only a few hours (1 4 hrs) (Drew, 1983). A variety of subtidal red algae can survive aerial exposure for 14 hrs only at humidities of 100 percent (Kain & Norton, 1990). Spores and germlings are highly intolerant of desiccation due to there relatively high surface to area ratios. Subtidal algae are more intolerant of than intertidal algae to desiccation.

If Delesseria sanguinea is removed from the substratum it can not reattach itself and it is therefore highly intolerant. Kain (1975) examined recolonization of cleared concrete blocks in a subtidal kelp forest. Red algae colonized blocks within 26 weeks in the shallow subtidal (0.8 m) and 33 weeks at 4.4 m. Delesseria sanguinea was noted within 41 weeks (8 months) at 4.4 m in one group of blocks and within 5659 days in blocks cleared at two monthly intervals during winter months, but not in subsequent samples (Kain, 1975). This suggests that Delesseria sanguinea can recolonize areas, but is directly dependent on its reproductive season and spore availability. Rhodophyceae have non flagellate, and nonmotile spores that stick on contact with the substratum. Norton (1992) noted that algal spore dispersal is probably determined by currents and turbulent deposition. However, red algae produce large numbers of spores that may settle close to the adult especially where currents are reduced by an algal turf or in kelp forests. It is likely that this species could recolonize an area from adjacent populations within a short period of time in ideal conditions but that recolonization from distant populations would probably take longer.

The effects of deoxygenation in plants has been little studied. Since plants produce oxygen they may be considered relatively insensitive. However, a study of the effects of anaerobiosis (no oxygen) on some marine algae concluded that Delesseria sanguinea was very intolerant of anaerobic conditions; at 15 deg C death occurs within 24hrs and no recovery takes place although specimens survived at 5 deg C. (Hammer, 1972).

Low Sensitivity

High Intolerances to substratum loss; synthetic compound contamination; heavy metal contamination and hydrocarbon contamination

Sea urchins are slow moving and unlikely to escape removal of their substratum. However, a proportion of the population would probably survive removal of algal substratum. Investigation of the effects of algal destruction on populations of Strongylocentrotus droebachiensis suggested that populations of urchins do not migrate away from or starve in areas devoid of kelp, presumably because they are able to feed on alternative prey. Areas lacking algae were dominated by young urchins up to 4 years after removal of the kelp suggesting that kelp barrens afforded improved recruitment (Lang & Mann 1978), presumably because of the lack of suspension feeding organisms associated with kelp beds. The presence of coralline algae in 'urchin barrens' may encourage larval metamorphosis in echinoids (Pearce & Scheibling 1990).

Low Sensitivity

High intolerances to substratum loss; smothering and displacement.

Removal of the substratum, be it shell, rock, or macroalgae will result in removal of the attached colonies of Electra pilosa. Therefore, an intolerance of high has been recorded. Recoverability is likely to be very high (see additional information below).

Smothering by 5 cm of sediment is likely to prevent feeding, and hence growth and reproduction, as well as respiration. In addition, associated sediment abrasion may remove the bryozoan colonies or the macroalgae on which they grow. A layer of sediment will probably also interfere with larval settlement. Therefore, an intolerance of high has been recorded. Recoverability has been assessed as very high (see additional information below).

Removal of a colony from its substratum would probably be fatal, and encrusting bryozoa are not known to be able to reattach. Therefore, an intolerance of high has been recorded. Recoverability is likely to be very high.

Moderate Sensitivity.

High intolerance to substratum loss; displacement; decrease in salinity and changes in oxygenation.

Sea fans are attached to the substratum and would not survive substratum loss. Therefore, intolerance is high. Recruitment would have to occur from nearby populations as larval dispersal is likely to be poor. Settlement appears to be sporadic and may not occur for several years. Growth to large colonies will take in excess of 10 years and replacement of very large colonies in excess of 25 years.

Colonies that are detached are likely to survive for some time lying on the seabed but will be little able to feed and will most likely be smothered. Colonies cannot reattach and are therefore doomed. Not all colonies are likely to be killed in a displacement event and recovery to a population structure similar to before mortality is likely to be a few years.

Sea fans live in fully saline conditions in the open sea. It seems likely that lowered salinity will kill colonies. Recovery will depend on recruitment which is likely to be very slow if surviving colonies are distant and even partial recovery may take more than ten years. Also, colonies grow slowly and the establishment of populations with large individuals will take many years.

There was no information available concerning the effects of hypoxia for Eunicella verrucosa. However, as a species that lives in fully oxygenated waters in conditions of flowing waters, it is expected that they might have a high intolerance to decreased oxygen levels. Recovery will depend on recruitment which is likely to be very slow if surviving colonies are distant and even partial recovery may take more than ten years. Also, colonies grow slowly and the establishment of populations with large individuals will take many years.

High intolerances to substratum loss; displacement; synthetic compound contamination; decrease in salinity and extraction of other species.

Removal of the substratum would result in the removal of the Flustra foliacea population and its associated fauna. Recovery will depend on recruitment from other populations and is assessed as high (see additional information below).

Colonies of Flustra foliacea that are displaced while attached to their substratum, e.g. shell debris or rocks will probably survive if moved to a suitable habitat and not crushed in the process. But if removed from its substratum, Flustra foliacea colonies cannot reattach and will probably be washed to deep water or be deposited on the strand line and die. Therefore, an intolerance of high has been recorded, with a recoverability of high (see additional information below).

Ryland (1970) stated that, with a few exceptions, the Gymnolaemata were fairly stenohaline and restricted to full salinity (ca 35 psu) and noted that reduced salinities result in an impoverished bryozoan fauna. Similarly, Dyrynda (1994) noted that Flustra foliacea and Alcyonidium diaphanum were probably restricted to the vicinity of the Poole Harbour entrance by their intolerance to reduced salinity. Although, protected from extreme changes in salinity due to their subtidal habitat, the introduction of freshwater, or hyposaline effluents may adversely affect Flustra foliacea colonies. Therefore, an intolerance of high has been recorded, and recoverability assessed as high (see additional information below).

Flustra foliacea may occur on coarse grounds used for fishing of other marine species, e.g. scallops. Mobile fishing gear, such as scallop dredges and beam trawls result in physical disturbance to the sediment surface. Emergent epifauna have been shown to be particularly intolerant of physical disturbance, depending on intensity (see abrasion) and emergent fauna, together with shells, and rocks form part of bycatch. Therefore, an intolerance of high and a recoverably of high have been recorded (see additional information below).

Moderate Sensitivity

High Intolerances to substratum loss; increased water flow rate; increased wave exposure; displacement and introduction of non native species.

The morphology of the stipe and blade vary with water flow rate. In wave exposed areas, for example, Laminaria hyperborea develops a long and flexible stipe and this is probably a functional adaptation to strong water movement (Sjøtun, 1998). In addition, the lamina becomes narrower and thinner in strong currents (Sjøtun & Fredriksen, 1995). However, the stipe of Laminaria hyperborea is relatively stiff and can snap in strong currents. It is usually absent from areas of high wave action or strong currents although in Norway it can do well in rapids (J. Jones, pers. comm.).

Birkett et al. (1998) suggest that kelp are stenothermal (intolerant of temperature change) and that upper and lower lethal limits for kelp would be between 12 °C above or below the normal temperature tolerances. The optimum temperature for the development of Laminaria hyperborea gametophytes and young sporophytes is between 1017 °C (Kain, 1971). Above 17 °C, gamete survival is reduced (Kain, 1971) and gametogenesis is inhibited at 21 °C in this species (tom Dieck, 1992). Given its distribution in the North Atlantic this species is likely to be tolerant of low temperatures. This species is likely to be intolerant of change in temperature equivalent to either benchmark outside its normal range. The temperature tolerances of the gametophyte stages are different to those of the adult. Gametophytic development has been observed at 0 °C although development is slow and suggests that 0 °C is close to the lowest temperature allowing vegetative development of the primary cells (Sjøtun & Schoschina, 2002).

Laminaria hyperborea is unable to survive where wave action is extreme because of its large frond area attached to a stiff stipe which is liable to snap. Wave action depresses the upper limit of populations to several metres below low water. In Norway, for example, the upper and lower limits of Laminaria hyperborea are raised from 5 to 0 m and 32 to 26 m from exposed to sheltered sites respectively (Kain, 1971b). It is absent from Rockall possibly due to extreme exposure and strong currents or geographical isolation. Older and larger plants, especially if the holdfasts are weakened by feeding by Helcion pellucidum, are most intolerant of wave action and populations affected by wave action have a reduced age range. As wave exposure increases Laminaria hyperborea is outcompeted by Laminaria digitata or Alaria esculenta. In a study in Norway (Sjøtun et al., 1993) Laminaria hyperborea from the most wave exposed site (in Finnmark) exhibited the lowest annual biological productivity per plant. Furthermore, of the four most exposed sites, three of them corresponded with the lowest mean standing crops (fresh weights). Therefore, Laminaria hyperborea is probably highly intolerant of increases in exposure at the benchmark level. It could benefit from decreases in wave exposure, possibly extending its upper limit up the shore, however this would only happen if its upper limit was depressed below the lowest astronomical tide (LAT) as it is highly intolerant of emergence (J. Jones, pers. comm.).

Laminaria hyperborea cannot reattach once removed and would be swept away. Experimental clearance experiments (Kain, 1979) in Isle of Man showed that Laminaria hyperborea out competed other opportunistic species (e.g. Alaria esculenta, Saccorhiza polyschides and Desmarestia spp.) and returned to near control levels of biomass within 3 years at 0.8m but that recovery was slower at 4.4m. However, Kain (1979) noted that grazing would slow recovery since, even though they did not prevent spore settlement, few sporophytes survived after 1 year in the presence of Echinus esculentus. These experiments did not remove the gametophyte 'seed' bank. Research on harvested populations of Laminaria hyperborea in Norway suggests that kelp forest biomass returned to preharvesting levels after 12 years, but that the plants were mainly small (1m) and that the age structure of the population was shifted towards younger plants. Sivertsen (1991, cited in Birkett et al., 1998) showed that kelp populations stabilize after about 45 year postharvesting. After 4 years post harvesting, kelps had only two thirds of their preharvesting canopy height. Regrowth was due primarily to growth of viable juveniles after harvesting. Current advice in Norway suggest that kelp forest should be left for 710 years after harvesting for the kelp biomass and nonkelp species to recover (Birkett et al., 1998b). Therefore, recovery is dependant on the depth (light availability) and grazing. However, given the potentially large number of spores and gametophytes it is likely that recolonization would occur rapidly and sporophytes may grow up to 0.94 cm /day under optimal conditions.

Low/Moderate Sensitivity

High intolerance to substratum loss; decrease in temperature and wave exposure.

Lanice conchilega lives in the sediment, a loss of substratum would cause a loss of the population. Therefore an intolerance of high has been recorded. A recoverability of high has been recorded (see additional information below).

Lanice conchilega is a species that is intolerant of low temperatures (Beukema, 1990). An intertidal population of Lanice conchilega, in the northern Wadden Sea, was wiped out during a severe winter of 1995/96 (Strasser & Pielouth 2001). The population took 3 years to fully recover, as there was low recruitment for the first 2 years. Crisp (1964) found there to be mortality of Lanice conchilega between the tidemarks but not at lower levels. After other hard winters Lanice conchilega recovered fast due to a good larval supply. Intolerance has, therefore, been recorded as high. Recoverability has been recorded as moderate (see additional information below).

Rees et al. (1977) found that only 1% of the Lanice conchilega population in Colwyn Bay apparently survived after winter storms. An increase in wave exposure is likely to lead to a high mortality of Lanice conchilega, therefore, intolerance has been recorded as high. Recoverability has been recorded as high (see additional information below).

Low/Not Sensitive

High intolerance to substratum loss; increase in emergence regime; extraction of this species and extraction of other species.

Adult Metridium senile live attached to the substratum and will be removed with it. Recolonization is likely to be rapid and occur within two or three years, although full recovery of dominance may take in excess of five years (see additional information below). Larvae of Metridium senile settle readily from the plankton and are known to colonize new structures such as wrecks and jetty piles within two to three years (K. Hiscock, own observations). Growth rate is rapid and a large size is reached in wellfed individuals within about nine months (see Bucklin, 1985). It is also possible that migration might occur from nearby adult populations as is believed to occur in recolonizing previously deoxygenated areas (Wahl, 1985). Therefore an intolerance of high and a recoverability of high has been recorded.

Metridium senile do not generally occur in the intertidal but may be present under piers and under overhangs, i.e. in shaded locations. In the case of an increase in emergence, migration is likely. Metridium senile is capable of detaching and floating away when conditions become undesirable. For instance, Wahl (1984, 1985) noted that in anoxic conditions, anemones detached from the substratum and drifted away. Although individual anemones are therefore unlikely to be killed by increase in emergence, they will be lost from a particular location and therefore intolerance is described as high.

If the species were to be extracted, mortality would occur.

Metridium senile sometimes occur, as small individuals, on kelp (Laminaria hyperborea) stipes. Kelp is extracted in some countries. Where attached individuals are collected, demise is certain.

Very Low Sensitivity

High Intolerance to substratum loss

Removal of the substratum, be it rock or sediment, will entail removal of the entire population and its associated community. Therefore, an intolerance of high has been recorded.Recovery may occur rapidly through good annual recruitment. However, examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. Recoverability will depend on recolonization by movement of young or juvenile Mytilus edulis from high shore or filamentous algal populations or recruitment of larvae settling directly on the new substratum. A single good recruitment event may return the population to prior levels within 1 5 years, and an intolerance of high has been recorded. However, recoverability may be protracted in some circumstances (see additional information below).


High Intolerance to substratum loss; dessication;displacement; Increase in Emergence regime and increase in wave exposure.

This species is permanently fixed to the substratum so substratum loss would cause death. See information on recoverability below.

The species is entirely subtidal and typically found below 10 m unless in very sheltered areas. Exposure to desiccating influences will probably cause death. See information on recoverability below.

The colonies of this species are permanently attached either to the substratum or to other colonies. On displacement individual colonies would be unable to reattach and therefore an intolerance of high has been recorded. See information on recoverability below.

The species is entirely subtidal and typically found below 10m unless in very sheltered areas. Emergence for an hour will probably cause death. See information on recoverability below.

The species is intolerant of high wave exposure and so is only found in sheltered areas. Increases in wave exposure above the preferred limits is likely to cause death, either through physical damage or prevention of feeding. See information on recoverability below.

High intolerance to substratum loss; smothering; increase in water flow rate; increase in wave exposure and displacement.

The adults are permanently cemented to the substratum so substratum loss would result in the death of the population. Adults are permanently attached to the substratum so no adult immigration is possible. No information is available about fecundity. Dispersal ability is not considered to be that great although in many locations there are nearby populations (particularly the west coast of Scotland). Reproduction occurs annually and over an extended period of time.

The dorsal valve of the shell can be clamped down and low oxygen concentrations can be tolerated for a few days. However smothering by sediment for a month will prevent feeding and restrict oxygen concentrations for considerably longer and will probably cause death. Adults are permanently attached to the substratum so no adult immigration is possible. No information is available about fecundity. Dispersal ability is not considered to be that great although in many locations there are nearby populations (particularly the west coast of Scotland). Reproduction occurs annually and over an extended period of time.

The species is found in waters with a maximum velocity of 23 knots. Increases above this level would probably cause death. Decreases in water flow rate are unlikely to have any effect as feeding currents are generated by the animal itself. Adults are permanently attached to the substratum so no adult immigration is possible. No information is available about fecundity. Dispersal ability is not considered to be that great although in many locations there are nearby populations (particularly the west coast of Scotland). Reproduction occurs annually and over an extended period of time.

This species is not very tolerant of wave exposure being generally found in sheltered locations like fjords and sea lochs or in deeper water. Increases in wave exposure above moderately exposed would probably cause death. Adults are permanently attached to the substratum so no adult immigration is possible. No information is available about fecundity. Dispersal ability is not considered to be that great although in many locations there are nearby populations (particularly the west coast of Scotland). Reproduction occurs annually and over an extended period of time.

Neocrania anomala is permanently attached to the substratum. If removed, the attachment cannot be reformed. Once detached, the brachiopod can then be moved around by water currents into unsuitable orientations or habitat and will probably cause death. Adults are permanently attached to the substratum so no adult immigration is possible. No information is available about fecundity. Dispersal ability is not considered to be that great although in many locations there are nearby populations (particularly the west coast of Scotland). Reproduction occurs annually and over an extended period of time.

Not sensitive/Low Sensitivity

High Intolerance to substratum loss.

Removal of the substratum would cause removal of adult and juvenile dog whelks together with their egg capsules. Therefore, an intolerance of high has been recorded. Given their poor dispersal ability, recruitment from other populations is likely to be slow, therefore a recoverability of low has been recorded.

Intolerant to substratum loss. 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.

High intolerances to substratum loss and smothering.

Ophiothrix fragilis is an epibenthic species so substratum loss would result in mortality. Breeding occurs annually and there may be multiple recruitment phases (Davoult et al., 1990). The larvae of this species can disperse over considerable distances in areas such as the English Channel where there are strong water flow rates (Davoult et al., 1990). With water that may move several kilometres per day due to residual flow (e.g. see Pingree & Maddock, 1977) and a larval duration of 26 days, the larvae can disperse up to 70100 km and establish populations elsewhere. Adults, although mobile, are not highly active. Some immigration of adults from nearby populations may be possible. Longevity estimates vary from 9 months (Davoult et al., 1990) to over 10 years (Gage, 1990). Reproductive capability may be reached in 610 months depending on time of recruitment (Davoult et al., 1990).

Although Ophiothrix fragilis is an epibenthic crawling species, it has a low level of locomotory activity and lacks muscular development. Smothering by 5 cm of sediment would probably cause death as it is unlikely that the brittle star would be able to burrow out of the covering material. Breeding occurs annually and there may be multiple recruitment phases (Davoult et al., 1990). The larvae of this species can disperse over considerable distances in areas such as the English Channel where there are strong water flow rates (Davoult et al., 1990). With water that may move several kilometres per day due to residual flow (e.g. see Pingree & Maddock, 1977) and a larval duration of 26 days, the larvae can disperse up to 70100 km and establish populations elsewhere. Adults, although mobile, are not highly active. Some immigration of adults from nearby populations may be possible. Longevity estimates vary from 9 months (Davoult et al., 1990) to over 10 years (Gage, 1990). Reproductive capability may be reached in 610 months depending on time of recruitment (Davoult et al., 1990).

High intolerance to substraum loss; displacement.

Palmaria palmata will be removed with the substratum, unable to reattach and it is therefore highly intolerant of substratum loss. There is little information on recruitment or recolonization rates. However, in kelp canopy removal experiments in the Isle of Man, Hawkins & Harkin (1985) observed a rapid increase in the number of Palmaria palmata sporelings and the species came to dominate cleared plots within five months. Rhodophyceae have non flagellate, and nonmotile spores that stick on contact with the substratum. Norton (1992) noted that algal spore dispersal is probably determined by currents and turbulent deposition. However, red algae produce large numbers of spores that may settle close to the adult especially where currents are reduced by an algal turf or in kelp forests. It is likely that this species could recolonize an area from adjacent populations within a short period of time in ideal conditions. However, since the dispersal range of spores is limited because the female does not release carpospores and needs to be close to the adult male population, recolonization from distant populations would probably take much longer.

If Palmaria palmata is removed from the substratum it is unable to reattach and so intolerance is very high. However, it is likely that this species could recolonize an area from adjacent populations within a short period of time in ideal conditions. In kelp canopy removal experiments in the Isle of Man, for example, Hawkins & Harkin (1985) observed a rapid increase in the number of Palmaria palmata sporelings on bare rock and the species came to dominate cleared plots within five months. Recolonization from distant populations would probably take longer, however, because dispersal distances are limited, with spores sinking and attaching close to adult plants.

Pholas dactylus lives permanently in a burrow excavated in soft rock, peat or similar substrata. Substratum loss will result in the death of the animal because when removed from its burrow and placed on the surface, it cannot excavate a new chamber (Barnes, 1980) and will be at risk from desiccation and predation. Provided a similar substratum remains and there is larval availability, recolonization is likely to occur and so recovery within five years should be possible, though maybe not to previous abundance.

Intolerance to removal from the substratum and displacement from original position onto a suitable substratum is high because Pholas dactylus cannot excavate a new chamber (Barnes, 1980) and so will die from predation or desiccation. Provided a suitable substratum remains and there is larval availability (the species spawns throughout the summer), recolonization is likely to occur and so recovery within five years should be possible, though maybe not to previous abundance.

Although no information on the specific effects of chemicals on Pholas dactylus was found TBT has been found to be toxic to many adult bivalves. Reports of reductions in numbers of bivalves in estuaries with high pleasure craft activity, have provided evidence of the high toxicity of TBT to bivalves (Beaumont et al., 1989). Laboratory toxicity trials have demonstrated that growth in oysters is inhibited by TBT (Waldock & Thain, 1983). In microcosm studies Beaumont et al. (1989) demonstrated that levels of 12µg/l TBT can rapidly kill adult bivalves in their natural habitat. For example, all Cerastoderma edule individuals died within two weeks at 13µg/l TBT concentrations and 80% died after 17 weeks at a TBT concentration of 0.060.17µg/l and Scrobicularia plana (Beaumont et al., 1989). Cerastoderma edule was found to be more intolerant of TBT than Scrobicularia plana in toxicity trials and was thought to be a reflection of the mode of feeding of the two species with filter feeding being a more direct route delivering a higher burden of the toxic material to the animal. Therefore, as a filter feeding bivalve Pholas dactylus it is likely that this species is also highly intolerant of TBT. Pholas dactylus spawns for several months every year, so when normal conditions resume rapid recolonization by the pelagic larvae is likely.

Low Sensitivity/Not Sensitive

High intolerance to substratum loss.

Removal of the substratum, perhaps by dredging, would result in the loss of Polydora ciliata tubes and hence the loss of the animals so intolerance is assessed as high. However, if some individuals remain rapid recolonization is possible because the species is capable of tube building throughout its life. Polydora ciliata of all ages that were removed from their tubes on many occasions, all built new tubes (Daro & Polk, 1973). Recovery is likely to be high because the larvae of Polydora ciliata are planktonic and capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.

Intolerant to physical disturbance long term changes from scallop dredging.

As it is an epifaunal species it would be significantly impacted by a lose of sediment. Also would be intolerant to increased salinity and smothering (as wouldn't be able to feed)

Low Sensitivity/Not Sensitive

High intolerances to substratum loss and smothering

Pomatoceros triqueter is attached permanently to rocks, boulders or shingle. Removal of substratum will remove calcareous tubes and animals contained in them. Intolerance is assessed as high. Recoverability is likely to be high (see additional information below).

Smothering with a 5 cm layer of sediment would completely cover the tubes of Pomatoceros triqueter that usually lie flat against the surface of the rock. It is also likely that too much sediment on the surface of rocks or shells would prevent settlement of larvae and impair the long term survival of populations. Intolerance has been assessed to be high. Recoverability is likely to be high (see additional information below).

Pomatoceros lives for up to 4yrs & matures at 4 months. The worm is hermaphrodite but the male & female gametes are separate at any one time. Spawning is at a maximum from MarchApril although breeding can occur throughout the year. Fertilisation is external & planktotrophic larvae then spend 3 weeks in the water column in the summer or as much as 2 months in the winter. The early maturation & long larval phase suggests that this genus has a strong recoverability potential.

Swiftia pallida would be removed by removal of the substratum, therefore intolerance has been assessed as high. Recovery would depend on the proximity of viable adults in the locality, as larval distribution is limited. If an entire population were removed, then recovery is unlikely. Hence recoverability is assessed as very low, resulting in a very high sensitivity assessment.

Swiftia pallida colonies grow attached to hard substrata, often perpendicular to the current. Displacement is likely to impair feeding and cause abrasion and mortality, as colonies are unable to reattach to the substrata following displacement. Therefore intolerance to this factor is assessed as high, and recovery very low, resulting in a very high sensitivity value.

No evidence was found on the tolerance of Swiftia pallida to a change in salinity. However the species occurs in fully saline conditions, so is assessed as highly intolerant to a decrease in salinity. Recovery would be very low due to poor larval recruitment and slow growth, therefore Swiftia pallida has been assessed as highly sensitive to a decrease in salinity.

No information was found on the effects of hypoxia on Swiftia pallida. However, the species lives in fully oxygenated conditions with high water flow levels, so it is expected that the seafan would be highly intolerant of decreased oxygen levels. Recovery will depend on recruitment, but is unlikely if surviving colonies are distant due to poor larval dispersal. Therefore recoverability has been assessed as very low, resulting in a very high sensitivity value.

Seafans are vulnerable to damage by mobile fishing gear, particularly in areas where they grow on low lying reefs adjacent to scallop beds (Tinsley, 2006). Trawling is likely to remove or severely damage Swiftia pallida. Eno et al. (1996) suggested that Eunicella verrucosa was "remarkably resilient" to impact from lobster pots. They found that some seafan colonies returned to an upright position immediately after impact, while others were permanently bent, which would reduce feeing efficiency. However Tinsley (2006) observed flattened seafans which had continued growing, with new growth being aligned perpendicular to the current, so clearly even colonies of Eunicella verrucosa which are damaged in this manner can continue to survive. Despite the likelihood that Swiftia pallida is resilient to static fishing, it has been assessed as highly intolerant due to the effect of dredges and trawls. Recovery of dislodged seafans is unlikely, and these colonies will probably die. Settlement of new colonies is dependant on recruitment, and larval supply, which is low, therefore recoverability is assessed as very low, resulting in a very high sensitivity value.

Low/Moderate Sensitivity

High intolerance to substratum loss

Urticina felina anemones adhere strongly to the substratum. Substratum loss would result in mortality. Some individuals might, however, be left behind as they typically live in fissures. Recruitment to replace lost individuals is likely to be slow.

Key prey taxa

Important food source for Tritonia hombergi and simnia patula

The sea slug, Acanthodoris pilosa, eats this species of bryozoan and lays its white egg ribbons on the fingers.

Lacked important predators

It is a source of food for Nucella lapillus in tidal sounds. Balanus crenatus is also grazed by Echinus esculentus and fish species probably nip its cirri.

Preyed upon by certain species of sea slugs.

Balanus is a barnacle which grows attached to hard surfaces such as boulders, shells & stones. The adults are <1cm in width & feed by capture of zooplankton with modified thoracic limbs. Barnacles are vulnerable to dredging & to deposition of large quantities of sediment mobilised by the dredging process. They are also susceptible to sedimentation that prevents location of suitable substrata for settlement by the larvae.

Balanus has a lifespan of 34yrs & is hermaphroditic. Female fecundity is up 8000 eggs and there may be several broods per year. Fertilisation is internal by an adjacent barnacle & large numbers of fertilised eggs are released as planktotrophic nauplius larvae into the plankton. The larvae are about 0.5mm in length & go through several stages of development before metamorphosing into a cypris larva. This is an active siteselection phase when the cypris is able to delay metamorphosis until a suitable substrate such as a rock or stone is located. This genus has a high dispersal & colonisation potential. Growth rates observed in this genus are fast, reaching sexual maturity in months & full adult biomass within a year. Full recovery of both population density & biomass is therefore achieved rapidly.

Important food source for copepodsAsterias Rubens

Cladophora rupestris forms an important habitat and food resource for juvenile isopods and amphipods that are a major component of fish diets (Jansson, 1967).

Callionymus maculatus, Trisopterus minutus, Gadus morhua

Is a main prey for Anarhichas lupus (wolf fish)

Electra pilosa is the preferred prey for several species of sea slugs, Limacia clavigera, Polycera quadrilineata, Onchidoris muricata and Adalaria proxima. Adalaria proxima is almost entirely dependant on Electra pilosa and its veligers will only metamorphose in the presence of Electra pilosa (Ryland, 1976; Picton & Morrow, 1994). Bryozoans, including Electra pilosa are also preyed on by pycnogonids (sea spiders) such as Pycnogonum littorale or Achelia spp. and sea urchins such as Echinus esculentus and Psammechinus miliaris (Ryland, 1976).

In some locations a tiny species of sea slug lives on the fans and feedson the polyps. The slugs coloration and shape match those of the polyps so may be difficult to spot.

Flustra foliacea is the preferred prey of the pycnogonid Achelia echinata, whose feeding behaviour is described by Wyer & King, 1973 and Ryland, 1976. In addition, Flustra foliacea is preyed on by sea urchins such as Echinus esculentus and Psammechinus miliaris (Ryland, 1976) and nudibranchs, especially the dorid Crimora papillata (Picton & Morrow, 1994).

The nudibranchs Doto dunnei and Eubranchus vittatus are often found feeding on this hydroid.

Polycera quadrilineata and Polycera faroensisSeveral species of sea slug, such as Onchidoris muricata

The primitive, shelled, tectibranch sea slug, Acteon tornatilis and wading birds.

can be pulled open by starfish,, broken into by crabs and drilled into by dogwhelks. They can even be attacked from inside

Nucella Lapillus

The sea slug Aeolidia papillosa and the sea spider Pycnogonum littorale feed on this and other sea anemones. It is possible that several fish species may feed on Metridium senile.

Asterias Rubens. Asterias preying on natural mussel beds indicated that Asterias detaches the majority ofmussel prey before ingestion

Preyed upon by crabs, dragonets callionymus lyra ad pleuronectes platessa

Temporal variable population

Balanus crenatus is also grazed by Echinus esculentus and fish species probably nip its cirri.

can be pulled open by starfish,, broken into by crabs and drilled into by dogwhelks. They can even be attacked from inside

Growth only occurs in summer

Connectivity to other habitats/species

Alaria esculenta forest can have an encrusting fauna of the mussel Mytulis edulis and semibalanus balanoides. Laminaria digitata can also be a part of the canopy

Another factor that might promote mussel recruitment is algal cover. In a previous study, we observed the extension of the shallow mussel zone associated with recruitment by the kelp Alaria esculenta (Gaymer et al., 2001a). The abrasion by macroalgae due to movement of the fronds by waves appears to deter movement of seastars towards mussels attached under the macroalgae, thus creating a refuge

From late July through to December the colonies of Alcyonium digitatum remain contracted, during which time they do not feed and assume a shrunken appearance with an obvious reddish or brownish colour. As a result of this period of inactivity the surface of the colonies becomes covered with a layer of epibiota; diatoms and prostrate thalloid and filamentous algae initially, from which arises a forest of erect algae and hydroids. The amphipod Jassa falcata also builds its mucous and detritus tubes amongst the other epibiota, adding to and consolidating the covering (Hartnoll, 1975).

frequently present, often in large numbers, on the leaves of seagrass (e.g. Zostera marina).

Inachus phalangium: the crabs seem content to reside around the base of small anemones, when the anemone is much larger the crabs can be found living right in the centre of the anemone.

Anemonia viridis interacts in many symbiotic relationships. The majid spider crab, Inachus phalangium and A. viridis are commensal. The anemone provides protection for the spider crab. The crab is immune to the possibly dangerous sting. Another commensal relationship includes Bucchichi's goby, Gobius bucchichi which lives in the tentacles of the snakelocks anemone without being harmed. The presence of the zooxanthellae has been shown to improve long term survival for A. viridis. As the zooxanthellae provides food, the anemone provides nitrogen. Without this mutualistic interaction neither species would flourish.

Mussel Beds

The branchial sac of Ascidia mentula is commonly inhabited by the pea crab Pinnotheres pinnotheres or the copepod Notopterophorus papilio. The small bivalve Modiolarca tumida can also live commensally in the test.

Balanus crenatus is an important initial colonizing species, perhaps obscuring material such as antifouling paint that would be toxic to other species.

The high densities at which the species occurs may help to reduce grazing by sea urchins. Stands of Ciona intestinalis provide refugia for several organisms, and some species grow on this species, albeit to a limited extent (e.g., Diplosoma sp.) (John Bishop pers. comm.).

Often associated with hydroids and bryozoans

Seagrasses as well as floating macroalgae, provide support for bryozoan colonies. In turn, bryozoans provide habitat for many species of juvenile fishes and their invertebrate prey such as polychaete worms, amphipods and copepods. (Winston 1995).Bryozoans are also found in association with other species that act as support structures: mangrove roots, oyster beds, mussels, etc.

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

Echinus esculentus is an important grazer of epiflora and epifauna in the subtidal. It may have a keystone role in kelp communities, where grazing by sea urchins may control the lower limit of Laminaria hyperborea beds, increase species diversity in the understorey epiflora/fauna, and habitat diversity through the formation of 'urchin barrens' (see Birkett et al., 1998b and EIR.LhypR Key Information review for discussion).

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.

Moore (1973c) noted that the hairy form of Electra pilosa on kelp holdfasts in north east England acted as a substratum for the settlement of mussel spat.

Often used by dogfish as an anchoring for their "mermaids purse" egg cases.

Eunicella verrucosa provides an important habitat for two associated species: the sea slug Tritonia nilsohdneri and the sea fan anemone Amphianthus dohrnii. The 'poached egg shell' Simnia patula also occurs on Eunicella but Alcyonium spp. are a more favoured habitat.

Aggregations of Flustra foliacea provide a habitat for various species of sponges, hydroids, caprellid amphipods and the suspension feeding crab Pisidia longicornis (as Porcellana), as well as a number of epizoic species (see MCR.Flu and Stebbing, 1971b).

Large lobsters moult very occasionally, if at all and can become covered in encrusting organisms, barnacles etc.

Halidrys siliquosa has been reported to support a number of epiphytic species, depending on location, including microflora (e.g. bacteria, blue green algae, diatoms and juvenile larger algae), Ulothrix and Ceramium sp., hydroids (e.g. Laomedea flexuosa andObelia spp.), bryozoans (e.g. Scrupocellaria spp.), and ascidians (e.g. Apilidium spp. and Botrylloides leachi ). However, Halidrys siliquosa was considered to be relatively clear of epiphytes due to its ability to shed the outer layer of epidermal cell walls, together with adherent epiphytes (Moss, 1982; Lobban & Harrison, 1997).

Kelps provide a unique habitat and substratum for many organisms and kelp forests are species rich habitats (Birkett et al., 1998b). Laminaria hyperborea provide three separate habitats, the blade (or frond), the stipe and the holdfast. The blades support Helcion pellucidum, the bryozoan Membranipora membranacea and the hydroid Obelia geniculata (Erwin et al., 1990; Birkett et al., 1998b) as well as endophytes and epiphytes, e.g. Myrionema corunnae (only found on Laminaria blades), and Pogotrichum filiforme and Chilionema sp., which are mainly restricted to kelp blades (Birkett et al., 1998b).The stipes support a diverse flora and fauna, especially foliose algae, depending on the age of the stipe, density of the kelp plants, and depth (Norton et al., 1977; Birkett et al. 1998b,). Hiscock & Mitchell (1980) list 15 species of algae associated with kelp stipes including, Palmaria palmata, Membranoptera alata, and Phycodrys rubens which are found mainly or solely on kelp stipes in the sublittoral.Kelp holdfasts support a diverse fauna that represents a sample of the surrounding mobile fauna and crevice dwelling organisms, e.g. cnidaria, polychaetes, nematodes, gastropods, bivalves, cirripedes, amphipods, isopods, copepods (mainly harpacticoids), and small crabs (Hoare & Hiscock, 1974; Jones, 1971; Moore, 1973a & b; Sheppard et al., 1980). Moore (1973a) lists 389 species from holdfasts collected from the north east coast of Britain.

Epizoid Speices associated with l. hyperborea: Halichondria reticulata (only on stipes), Obelia geniculata (on Thalli), Sertularia operculata (On stipes generally), Aglaophenia pluma (On stipes generally), Mebranipora membranacea (on Thalli), Electra Pilosa (on stipes and epiphytes), Celleporlla hyalina, Tridemnium and didemnium spp. (on stipes).

The tube which Lanice conchilega builds provides structure to the sediment. Lanice conchilega consolidates the sediment, obstructing the activities of predatory burrowers and enabling other sedentary animals to establish themselves. Lanice conchilega can be found as a solitary individual or in populations of several thousand per m2

Even in relatively low numbers Lanice alters the species composition of a habitat by creating favourable conditions for other species to live. Gammarus locusta has been associated with Lanice in intertidal studies. Lanice is potential refuge for species and an attachment site for juveniles.

Mussels only offer passive resistance to most predators but they can take an active stance against dog whelfs, sometimes tying them down until they starve to death.

Metridium senile may dominate large areas of rock and artificial substrata such as jetty piles and wrecks. It is capable of aggressively displacing other species through the use of 'catchtentacles' that sting adjacent fauna causing necrotic patches to develop.

The field observations showed a firmer attachment of mussels on stations with Asterias, and field and laboratory experiments revealed that mussels attach more strongly when starfish are present. Either the field or the laboratory experiment demonstrated a correlation between starfish predation rates and the mussel attachment strength, suggesting that the increased mussel attachment was caused by the presence of Asterias by a tactile or chemical stimulation of the mussels, and not as a result of higher predation rates on loosely attached mussels.Suchanek (1992), Seed (1996) and Holt et al. (1998).

Ecosystem importanceLarval production represents a significant contribution to the zooplankton, forming an important food source for herring larvae and carnivorous zooplankton (Seed & Suchanek, 1992). Dense beds of bivalve suspension feeders increase turnover of nutrients and organic carbon in estuarine (and presumably coastal) environments by effectively transferring pelagic phytoplanktonic primary production to secondary production (pelagicbenthic coupling) (Dame, 1996).

Nehls & Thiel (1993) suggested that removal or exploitation of persistent mussel beds may reduce or remove food reserves crucial to birds such as eider and oystercatchers. Holt et al. (1998) noted that gross declines in mussel stocks due to overexploitation or recruitment failure may adversely affect the surrounding ecosystem. For example, in the Dutch Wadden Sea unprecedentedly low mussel numbers in 1990 resulted in death or migration of eider, and oystercatchers seeking alternative prey and increased pressure on cockles, Macoma balthica and Mya arenaria (Holt et al., 1998).

Where farmed shellfish rafts cover 10% or more in areas of poor water exchange (e.g. sea lochs) suspension feeding by mussels can significantly reduce levels of phytoplankton with indirect effects on other filter feeders. Reduced phytoplankton levels in one Scottish loch also adversely affected the farmed mussel production resulting in the closure of the farm. However, most Scottish mussel farms were thought to be too small to have serious effects on plankton levels (McKay & Fowler, 1997; Holt et al., 1998).

Although overexploitation of local fisheries was identified as a potential problem (Holt et al., 1998) the longevity of the major natural fisheries was probably a result of careful management by Sea Fisheries Committees.

While a dogwhelk is preoccupied with its drilling, the mussel and its neighbours may attach so many of their byssus threads to the dog whelk that it

In Torbay, Nemertesia antennina, a similar species, has been recorded as hosting more than 150 epizoic species, most of which are not present on other local substrata.

Associated fauna, with a dominance of large suspension feeders such as alcyonium digitatum, anemones, metridium senile, urticina felina and nemertesia antennina fixed on rock outcrops or boulders protruding above the brittlestar covered substratum.

Hydractinia echinata (hydroid), Nereis fucata (Annelid), Calliactis parasitica (Cnidaria).

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.

Algae or encrusting animals such as broyzoans or serpulid worms are often found growing on it.

Often found epizoically on crustacean carapaces, the shells of molluscs, tests of ascidians and on algae.

Home to pycnogonids, caprellids, mussel spat and naudibranchs.

Fouling by this species can compete with and exclude other species.

Bryozoans can be found growing on the tubes

Additional Notes

Has a variety of body forms.

A. Diaphanum is a brooding species, incubating multiple embyros in single autozooids

Usually brownish in colour with wavy surfaces, the fingers can be branched and are often found in bunches.

Maximum size 30cm

Migné et al. (1996) conducted a biometric study on Alcyonium digitatum with the aim of relating both the carbon and the nitrogen content of this species to a simple and rapid measurement. It appeared useful to express the biomass of this species in terms of carbon or nitrogen and then to consider dynamic processes such as respiration or excretion as fluxes of carbon and nitrogen. Converting biomass into a more fundamental measurement of living matter such as carbon or nitrogen content should enable understanding and quantification of the species role in carbon and nitrogen cycles. For instance, the bottom of the Dover Strait (eastern English Channel) is dominated by suspension feeders (Davoult, 1990) that probably play a leading role in the exchange of organic matter at the bottom boundary layer (Migné et al., 1996). Three species, including Alcyonium digitatum accounted for 97% of the total biomass of the suspension feeding community studied in the English Channel (Migné & Davoult, 1995 b).

In the Autumn, most colonies stop feeding and withdraw their tentacles for several months, while they prepare to spawn. When feeding is resumed the outer skin is shed, along with any encrusting growths that settles with the tentacles were out of action.

200 tentacles, cant retract. Large populations of symbiotic algae. In this close association, the algae gain protection and a supply of carbon dioxide and nutrient salts. The host anemone benefits from organic compounds synthesised by the algae using light.

Longlived species which produces small eggs

Its grey tunic has a rough surface which tends to trap a layer of silt and other detritus. Althought classified as solitary, it is usually found in clumps.

Individuals are often grouped together, so what may look like two siphons of one squirt are actually the intake siphons of two individuals whose outflow siphons are hidden.

In a few cases, dislocation of the animals was observed, either by removal from the substratum by browsing sea urchins (Echinus esculentus) or when older individuals were used as a substratum and later died.

Past attempts by shellfish collectors to kill dredged up starfish by simply chopping them in half and throwing them back into the sea were doomed because each half starfish simply grew another half and carried on as before.

The plates of dead Balanus crenatus are probably an important part of the unique shell gravel banks in the Menai Strait, North Wales.

3.5cm at its largest.

Slow growing (0.51mm per year).

Not classified as a true sea anemone. Rather than attching itself to rock, or a similar firm surface, it lives in a soft feltlike tube constructed by specially adapted discharge capsules. Can not retract tentacles but retracts entire body into tube.

Fast growing and will rapidly take over freshly available habitats such as marina pilings and chains. They can form extremely dense aggragations of up to 5000 squirts per square metre. Individuals are not fused in any way so known as solitary as oppposed to colonial.

Pest to shellfish cultivation. Highly efficient filter feeder, each pumping 3 litres of water an hour and removing particles as small as a thousandth of a mm.

Live in bunches that may contain a few or dozens of individuals. Colonial species, although individuals are free along most of their length and are joined to each other only at their base.

Bores its way into soft rock such as limestone. In many instances most fot he sponge is hidden within a network of passages and chambers that is has excavated in the substrate. Boring process is chemical and employs an acid produced as a byproduct of respiration. Also bores into the shells of molluscs, can be a serious pest for commercial oyster beds.

Colonies of C. eburnea form fragile white tufts as they grow on the stems of algae.

Desmarestia spp. Contain sulphuric acid in their cells. When collecting, the acid is released into the water, causing other seaweeds kept in the same contained to disintegrate.

Perennial Species

Most common and widely distributed sponges but because it is fairly small is overlooked.

Sponges skeleton is made of portein fibres. No calcium or silicon based spicules.

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

Max Diameter 16cm

Echinus are grown in aquaculture due to demand for urchin roe in the Asian marketDescribed as a key functional species, due to its grazing traits

Colonies can only grow for part of the year so annual growth lines can sometimes be seen.

Fans usually only branch in one plane which is at right angles to the prevailing current, thus giving each animal the maximum opportunity for feeding. The fans grow slowly at approximately 1cm/ year and may be over 100 years old. Easily knocked off and will die. Protected by the wildlife and country side act. BAP Species.

Green coloration is due to symbiotic algae, and is dependant on the amount of light available.

The name cottonspinner comes from the animal's habit of producing long white threads from its rear end if molested. The threads are part of its internal organs and are extremely sticky, serving to confuse of entangle the attacker.

Usually only emerge at night but can be seem roaming the seabed during the day, such encounters usually happen when on elobster has been disturbed from its hiding place.

Such meshencrusted fronds in shallow water are a good place to find a variety of sea slugs and their eggs.

Deposit feeders when Lanice is in low densities. But when densities of the species increases the worm can switch feeding method to suspension feeding.

Where Lanice occurs in high densities the water flow around the tubes is deflected around the aggregations. Can switch between suspension and deposit feeding.

The worm itself is hardly ever seen, but it extends its sticky tentacles along the "tree's" branches to catch suspended food. The tentacles also reach out over the seabed to pick up deposited food particles and any sand grains required for maintenance. The branches, orientated across the current, serve to slow down water flowing past so more suspended material collects around them.

Colonies may vary in size. Membranipora membranacea grows quickly (several mm/day) in response to predation from sea slugs and in order to maintain its position on the kelp frond. Prolific growth is favoured by fast flowing water which provides food and oxygen. Egg production occurs once a year in the spring; the larva (cyphonautes) is planktotrophic and is the most abundant and largest coastal larva in Britain between June and August. The larva settles in late summerearly autumn.

Mussels only offer passive resistance to most predators but they can take an active stance against dog whelfs, sometimes tying them down until they starve to death.

Sensitive indicator of pollution, caused by tbt.

If currents get too strong they simply link arms to form a huge mat that is less likely to get swept away.

O fragilis plays a major role in pelagobenthic transfer of particles from the water colum to benthic habitats due to their suspension feeding activity.

Species reacted to mechanical distrubance.

In the subtidal Area, Ophiothrix shows gregarious behaviour, and forms dense beds up to several thousand individuals per m2. brun 1969 suggested that such aggregations results from the actvie assoication of animals with their conspecifics rather than simply from the individual responses of the brittlestars to the characteristics of their physical environment. Living in dense aggregations may reduce displacement by stong currents. it is also favourable in maximising the capture of fine particle from the water column, mazimises success of fertilisation.

Large aggregations could have an impact on the transfer of suspended matter between water column and sea bed.

Where dense O.fragilis aggregations are found on bedrock surfaces, they may monopolise the substratum, virtually to the exclusion of other epifauna.

Protula tubularia is often solitary and reproduces throughout June, August and September.

Very sensitive and will retracts into tube at slightest disturbance.

Invasive Species Regarded as a threat to human activities in sheltered shallow water

It can outcompete local species because it is fast growing, can reproduce within the first year of life and being monoecious can fertilize itself.

Maximum size 15cmFood source for flat fish species.Stings prey with nematocysts

Tubes are calcerious

Confidence

R1 Medium

R23 Medium

R1 MediumR124 Low

R126 Medium

R123 LowR24 Medium R1 MediumR34 High

R54 High

R116 High

R301 MediumR307 MediumR88 MediumR26 High

R1 Medium

R2 Medium

Reference Code

R301 MediumR307 MediumR118 Low

R55 Medium Expert Opinion Low

R301 MediumR308 Medium

R56 Medium

R1 Medium

R301 Medium

R3 MediumR57 Medium Expert Opinion LowR60 Medium

R59 Medium

R1 MediumR58 MediumR33 HighR62 Medium

R61 Medium

R301 MediumExpert Opinion LowR305 MediumR306 High

Expert Opinion Low

R122 Medium

R63 Medium

R301 MediumR306 HighR66 Low


R301 Medium

R4 Medium

R1 Medium

R1 Medium

R25 Medium

R1 Medium

R5 Medium

R51 MediumR1 MediumR35 HighR64 MediumR301 MediumR88 MediumR65 MediumR1 MediumR301 Medium

R6 MediumR1 MediumR308 Medium

R127 High

R1 Medium


R310 Medium R1 Medium

R7 Medium

R1 Medium

R301 Medium

R8 MediumR122 Low

R1 Medium

R9 Medium

R1 Medium

R301 Medium

R1 Medium

R10 Medium

R1 Medium

R301 Medium

R1 Medium

R11 MediumR1 MediumR301 MediumR307 Medium

R306 MediumR62 MediumR1 MediumR88 MediumR311 Medium

Expert Opinion Low

R67 Low

R1 Medium


Expert Opinion Medium

R69 HighR1 MediumR74 MediumR312 High

Expert Opinion LowR71 MediumR70 HighR1 MediumR73 High R72 MediumR70 High

Expert Opinion LowR1 MediumR313 HighR307 Medium

Expert Opinion MediumR109 MediumR329 MediumR330 MediumR301 Medium

R1 Medium

R12 Medium

R35 High

R36 High

R37 HighR38 HighR39 High

R1 MediumR109 MediumR121 High

R13

R75 MediumR314 High

Expert Opinion LowR308 Medium

R97 MediumExpert opinion Low

R1 Medium

R301 Medium


R14 MediumR118 LowR76 MediumR301 MediumR97 MediumR109 MediumR1 Medium

R315 Medium R78 HighR77 Medium



Expert Opinion LowR82 MediumR80 Medium

R1 Medium

R15 MediumR1 MediumR97 MediumR331 MediumR107 Medium R83 Medium

R301 Medium

Expert Opinion Medium R329 MediumR62 MediumR84 Low


R301 MediumR85 MediumR301 Medium R86 Medium

R87 MediumR88 HighR332 Medium

R62 Medium

R1 Medium

R16 Medium



R28 Medium R40 High

R41 High

R301 HighR42 HighR1 MediumR301 MediumR1 MediumR91 MediumR333 MediumR92 MediumR90 HighR88 HighR128 MediumR88 High

R93 MediumR94 HighR121 MediumR1 MediumR301 Medium

R103 Low


R125 Medium




R1 Medium

R29 Medium

R1 Medium

R325 Medium

R30 Medium


R303 Medium

R88 HighR97 MediumR323 Medium R96 Low


R1 MediumR120 High

R301 Medium

R119 Low

R18 Medium

R1 MediumR301 HighR43 HighR44 HighR45 Medium

R46 HighR1 Medium


Expert Opinion LowR336 MediumR337 MediumR338 Medium R301 MediumR88 High

R322 LowR1 MediumR98 Medium



R20 MediumR101 HighR77 MediumR33 MediumR320 MediumR105 Medium R104 MediumR1 Medium

R319 Medium

R304 MediumR97 MediumR69 HighR62 MediumR1 MediumR107 MediumR106 Medium

Expert Opinion LowR339 MediumR340 Medium




R112 Medium

R47 HighR48 HighR318 Medium

R49 HighR50 HighR51 High R88 High

Expert Opinion R1 Medium


R53 Medium

R51 Medium

Expert Opinion


R113 Medium R62 Medium

R32 Medium

R32 MediumR97 Medium R308 Medium


R1 Medium

R22 Medium

Species Gro

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Ahnfeltia plicataAlaria esculentaAlcyonidium diaphanumAlcyonium digitatumAnemonia viridisAntedon bifidaAntedon petasusAscidia mentulaAscidiella aspersaAsterias reubensAxinella dissimilisBalanus crenatusBugula plumosaCancer pagurusCaryophyllia smithiiCerianthus lloydiiChondrus crispusCiona intestinalisCladophora rupestrisClavelina lepadiformisCliona celataCorallina officinalisCorella parallelogrammaCrisia eburneaDelesseria sanguineaDesmarestia aculeataDexamine spinosaDyopedos porrectusDysidea fragilisEchinus esculentusElectra pilosaEpizoanthus couchiiEunicella verrucosaFlustra foliceaGibbula cinerariaHalichondria panicea

Haliclona viscosaHalidrys siliquosaHarmothoeHolothuria forskaliHomarus gammarusJanolus cristatusKirchenpaueria pinnataLaminaria hyperboreaLaminaria ochroleucaLanice conchilegaLuidia ciliarisMargarites helicinusMembranipora membranaceaMetridium senileMytilus edulisMyxilla incrustansNecora puberNemertesia ramosaNeocrania anomalaNucella lapillusObelia geniculataOphiothrix fragilisPagurus bernhardusPalmaria palmataPandalus montaguiParasmittina trispinosaPhellia gausapataPholas dactylusPolyclinum aurantiumPolydoraPolyides rotundusProtula tubulariaSabella pavoninaSargassum muticumSertularia argenteaSpirobranchus triquiterSuberites carnosusSwiftia pallidaUlva lactucaUrticina felina

Literature ReviewExpert OpinionData Gap

Sub

stra

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Tida

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Phy

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Tem

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Model Level

R97 5. Output processes Secondary Production

R100 1. Regional to global drivers Temperature

R100 1. Regional to global drivers Water Currents

R100 1. Regional to global drivers Depth

R103 1. Regional to global drivers DepthR103 1. Regional to global drivers Temperature


R104 5. Output processes Biodeposition

R104 6. Local ecosystem functions Food Resource



R110 1. Regional to global drivers Climate

Source Code

Model Parameter Assessed

R113 1. Regional to global drivers Seasonal variability


R114 6. Local ecosystem functions Ecosystem functions

R114 6. Local ecosystem functions Habitat provision



R117 2. Water column processes Dissolved Oxygen

R119 2. Water column processes Suspended Sediments


R123 3. Local processes at the seabed Benthic topography

R126 5. Output processes Bioengineering


R130 1. Regional to global drivers


R132 2. Water Column Processes Suspended Sediments

R132 3. Local processes at the seabed Seabed Energy



R155 2. Water column processes Suspended Sediments

R155 2. Water column processesDepth

R156

2. Water column processes

Dissolved oxygen

R157


Primary production

R1572. Water column processes

Primary production

R158

1. Regional to global drivers

Climate

Temperature, hydrodynamics, currents

R161


Depth

R161


Light attenuation

R161


Depth

R162


Light attenuation

R163

5. Output processes

Secondary Production

R163

6. Local ecosystem functions

Biogeochemical cycling

R164


Wave exposure

R170 3. Local processes at the seabed Habitat Complexity


R171 1. Regional to global drivers Wave Exposure

R171 6. Local ecosystem functions Mobile Invertebrate fauna

R171 5. Output processes KelpR171 1. Regional to global drivers Wave exposure

R171 1. Regional to global drivers Wave exposureR172 1. Regional to global drivers Wave exposure

R173 2. Water column processes TurbidityR174 2. Water column processes Water Movement

R175 6. Local ecosystem functions Kelp


R176 5. Output processes Kelp

R177 5. Output processes Macroalgae

R178 5. Output processes Ecosystem functions










R179 2. Water column processes Climate/Wave energyR179 1. Regional to global drivers Seasonal variability

R180 1. Regional to global drivers Wave exposure

R181 3. Local processes at the seabed Secondary Production

R182 3. Local processes at the seabed Sediment deposition



R185 3. Local processes at the seabed Habitat complexity




R188 6. Local ecosystem functions Ecosystem functionsR189 3. Local processes at the seabed Tidal flow


R190 3. Local processes at the seabed Habitst StructureR191 3. Local processes at the seabed Water Currents






R195 3. Local processes at the seabed Speciies diversity

R195 3. Local processes at the seabed Species competition


R195 3. Local processes at the seabed Light attenuation

R160


GeologyR165 1. Regional to global drivers Depth

R1641. Regional to global drivers

DepthR165 1. Regional to global drivers Water Currents


Wave exposure


Wave exposure


Water CurrentsR88 6. Local ecosystem functions Habitat provision

Linking to Fauna/Species specific focus (if applicable)

Venerid bivalves Asterias rubens, bivalves

Fauna

Fauna

Fauna

Biological ProductionBiological Production

Fauna

Epifauna Brittlestars

Higher trophic levels Brittlestars

Epifauna Hydroids

Fauna Suspension feeders

Fauna

Fauna Pomatoceros, filter feeders, deposit feeders.

Fauna Bivalves

Fauna Bivalves

Epifauna Bivalves

Fauna

Fauna Hermit crabs

Epifauna Hermit crabs

Fauna Bivalves

Fauna Bivalves

Fauna

Fauna

Nutrient cycling

Fauna

Primary production

Primary production

Microbial activity Microorganisms and algae

Epifauna Actiniaria

Epifauna Actiniaria

Light attenuation

Light attenuation

Temperature

Food sources

Water Chemistry and temperature

Primary Production

Light attenuation

Primary production

Temperature

Food sources

Fauna

Water Chemistry and temperature

Suspended Sediments

Species diversity

Species diversity Urchins


Carbon transfer Mobile Invertebrate fauna

Production of POM Laminaria hyperboreaFlora Algae

FaunaSpecies Diversity Kelp

Kelp KelpKelp Kelp

Modification of hydrodynamic streams Kelp

Sediment transport Kelp

Recruitment Kelp

Macrofauna/Habitat complexity

Bioerosion Sponges

Habitat Stabilisation Sponges

Water column chemistry Sponges

Carbon transfer Sponges

Biogeochemical cycling Sponges

Bacterial activity Sponges

Secondary Production Sponges

Habitat provision Sponges

Habitat provision Sponges

Hydrodynamic conditions Sponges

AlgaeAlgae

Species diversity Algae

Species diversity Urchins



Recruitment Gastropods

Gastropods Gastropods

Fauna Asterias rubens

Bioengineering Mytilus

Water column chemistry/biodeposition Mytilus

Secondary Production GastropodsFauna Brittlestars

Nutrient cycling Brittlestars

Species diversityFeeding ability Feather stars

Biodeposition Mytilus

Habitat provision Mytilus

Water Chemistry and temperature Mytilus

Biodeposition Mytilus

Nursery Habitat Kelp beds

Light attenuation Kelp

Fauna

Export of organic matter Kelp

Light attenuation Kelp

Suspended SedimentsWave exposure

Water Chemistry and temperatureSuspended Sediment

Dissolved Oxygen

Fauna

FaunaFauna Hydroids

Limitations in evidence

Output High

Input High

Input Medium

Input High

Input MediumInput Medium

Output High

Output Positive High

Output High

Output Medium

Input High

Input Medium

Direction of interaction

Positive/Negative (if output process)

Magnitude of Interaction

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

Info based on Ophiothrix fragilis

Info based on Ophiothrix fragilis

Input Medium

Feedback Low


Output Positive Medium Based on Mytilus edulis

Input Low

Output Low

Input Medium

Input Low

Feedback Negative Low

Input Medium Study from Australia.

Input Negative Low

Input Medium

Based on interactions in an estuarine environment

Study based on Paguristes eremita in Adriatic Sea

Study based on Paguristes eremita in Adriatic Sea

Study based on Mytilus edulis farms in SW Ireland

Input Medium

Input High

Input High

Input Medium

Output Positive Low

Output Positive Medium

Input

High

InputHigh

Input Medium

Input

High

InputHigh

Input

High

Input

High

Input

High

Input

High

Input

High

Output Positive

High

Output Positive

High

Input

High

Input Medium

Output NegativeHigh

Input Medium

Output PositiveHigh

Output PositiveHigh

Input Medium

Input MediumInput High

Input MediumInput Low Study based in Ireland

Output Negative Medium Study based in USA

Output Negative Medium Study based in USA

Output Negative Low Study based in USA

Output Positive Low Study based in Spain

Output Negative Medium



Output PositiveHigh




Output PositiveHigh

Output PositiveHigh

Output Negative Low

InputHigh

Input High

Input Medium

Input High

Input Medium

Input Medium


Input Medium Study based in Australia

Input Medium



Output Negative MediumInput Positive Low


Input Positive HighInput Positive Medium


Output Positive

High




Input Medium

Input Medium


Input Medium

Input LargeInput Large

Input MediumInput Medium

Input Medium

Input Medium

Input MediumOutput Positive Low

Information Summary Source Scale/Location

Lab based

National - UK Wide

Tidal stress has a positive effect on benthic production National - UK Wide

Depth has a negative influence on benthic production National - UK Wide

Biological production negatively related to water depth GeneralBiological production positively related to temperature General

General

Regional - English Channel

Regional - English Channel

General

Local - Bay of Brest, France

General

A Rubens is a voracious predator of bivalve molluscs. Abra, Spisula and Nucula were considered in this study.

Temperature has a positive effect on benthic abundance, diversity and biomass

Highest biological production recorded for filter feeders, bivalves, omnivores, predators and arthropods.

Brittlestars play a major role in pelago-benthic transfer of particles from the water column to benthic habitats due to their suspension feeding activityBrittlestars preyed upon by crabs, dragonets Callionymus lyra and plaice Pleuronectes platessaHydrozoans capture large amounts of zooplankton and seston, and play an important role in transferring energy from pelagic to benthic ecosystemsStrong currents favour suspension feeders and mixing helps phytoplankton growth.

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. Plankton bloom might change. Has info on some specific species.

Local - Estuary in Portugal

General

General

General

General

Regional - Adriatic Sea

Regional - Adriatic Sea

General

General

General

General

Local - SW Ireland

Seasonal variation in dominant functional groups corresponding to changes in salinity, oxygen, chlorophyll and nutrient concentrations. The taxa that showed clear seasonal changes were C. multisetosum, Streblospio shrubsolii, H. diversicolor, C. carinata and H. ulvae. In winter, filter feeders became increasingly more important. although burrowing species were the most abundant, seasonal differences were observed for some parts of the estuary. In summer, sessile species (e.g. Pomatoceros lamarcki and Modiolus Modiolus) were only found at the lower estuary, while in winter they were absent.

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.Climate change will likely change species distribution. Increased rainfall may influence salinity. Hermit crabs offer habitat provision to symbionts and epibiota, affecting abundance and distribution of other invertebrates. .

Low dissolved oxygen in the water column triggers a response in hermit crabs. High anoxia may cause fatalities. Turbidity can have a positive influencing effect on filter feeding bivalve populations. Filter feeding bivalves can have a top-down control on phytoplankton abundance.

Microtopography clearly influenced the structure of the benthic macrofaunal assemblages. Numbers of taxa of the benthic macrofaunal assemblages and almost all individual taxa analysed showed significant greater abundances in troughs than in crests. The results strongly support the model that benthic macrofauna are affected by locally varying hydrodynamic environments produced by ripple-beds.

Tubes of invertebrates may reduce the mobility of other benthic burrowing fauna, reducing ecosystem function, abundance and diversity.

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

Regional - North Sea

General

General

General

General

General

General

General

General

General

General

General

Drivers affecting macrobenthic and epibenthic species distribution in the North Sea. As depth increases, light attenuation decreases, thus primary production decreases. As turbidity increases (often with a depth decrease) light attenuation will decrease, thus primary production may decrease.

Microbial activity is low in areas of higher energy as there is limited organic detritus available for bacterial degradation coupled with the particles’ comparatively low surface area to volume ratio that provides a surface for microbial populations.

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

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

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.Significant relationship between depth and light attenuation

Decreasing dissolved oxygen levels were being influenced by increasing temperatures and biomass input (which in turn was being driven by the spring blooms and stratification).

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

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.

General

General

General

General

General

General

General

Epiphytic algae increases with wave exposure.

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 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.

All environments depend on the surface primary production. Phytoplankton die and sink to the sea bed where they are eaten by other species, or decay releasing organic material.

Secondary production is the amount of biomass of an animal created as a direct result of trophic level consumption E.g. biomass of secondary consumer, tertiary consumers etc.

Minerals, particularly ammonia, are excreted by secondary producers and are formed during bacterial decomposition or organic matter. These minerals are either consumed directly or are oxidised and dissolve into the water column.

Wave action on sediments is more variable than currents due to intensity of atmospheric weather. Waves resuspend sediment particles and transport them. In calm weather water depth of 10-20m will be affected whilst during storms sediment down to 200m could be altered.

Vertical rock walls are usually inaccessible to urchins and have a different association of organisms. On these walls are a variety of encrusting, mostly colonial invertebrates.Sea urchins are major grazers of kelps and algaes and can act to control their distribution.Secondary production of mobile invertebrate fauna in kelp forests increases with wave exposure levels.

Mobile invertebrate fauna has a key function in transferring kelp carbon to higher levels in the food web.Kelps produce significant POM, some of which is exported outside of the ecosystem.

Faunal abundance increases with wave exposure.Faunal diversity increases with wave exposure.

Water motion has little impact on kelp growth rates.

Local - Spain

General

General

General

General

General

General

General

General

General

General

Local - AustraliaAlgaes subject to high levels of seasonal variability. Local - Australia

Local - Australia

Local - Australia

Silting and turbidity has a negative effect on kelps, affecting the holdfast fauna.

Kelp has a negative effect on water flow rate near the seabed and may account for increased deposition of sediments contained within the water column near the bed. Kelp can have a negative effect on sediment particle transport from overlying water column to the bottom.

Kelp can have a negative effect on the recruitment of benthic organsims through altered hydrodynamics beneath the canopy, increased sedimentation and the blocking of light.

Habitat structure from epilithic algae has a positive effect on benthic faunal abundance, although other factors may be more important. Eplithic biomass is linked to faunal biomass.Sponges are one of the primary bioeroders on reef structures, esp. boring sponges.

Sponge colonisation may lead to the stabilisation and consolidation of habitat in the rocky subtidal zone, binding boulders to rock.

Sponges remove nutrients and from the water column and have the capacity to significantly impact pelagic ecosystemsSponges are imporant factors for carbon flow through the ecosystem being such large filter feeders.

Sponges and their associated microorganisms are important fixers of nitrogen in the marine enviornment. The presence of sponges has been shown to increase bacterial biomass in an ecosystem

Sponges are consumed by a range of organisms including fish, molluscs, crustaceans and echinoderms. Sponges act as microhabitats and can provide habitats for other organisms.Sponges may act as a settlement surface for other organisms. Sponges may be responsible for altering localised current flow dynamicsBiomass of algae correlated with changes in light availability, wave energy and sea temperature.

Wave exposure is a controling factor on strucure, diversity and assmeblages of macroalgaeUrchins can be a controling factor on macroalgae and turf algae.

Local - Medeterranean

Local - Baltic

Local - UK

Local - Australia

Local - Wadden Sea

Regional - Baltic

Regional - Baltic

Local - UKBrittlestar feeding intensity is influenced by tidal flow. Local - UK

Local - UK

Local - South AfricaCurrent speed influences featherstar feeding. Lab based

Regional - Baltic

General

General

Sediment deposition significantly affects the local structure and diversity of macroalgal assemblages. Erect algae is more likely to be affected. Growth of algae was shown to be enhanced in periods of reduced sedimentation.

Depth is a crucial factor in determining macroalgal assemblage diversity. Abiotic factors (sedimentation, wave exposure, wave and water currents, slope of seabed) are important in algae distribution.

Grazing by Gibbula can have a negative impact on recruitment of other organisms in rocky suntidal habitats.The complexity of habitats positively affects the density and richness of gastropods

Temperature is a controling factor in starfish feeding rates, as is light intensity. Asterias reubens is subject to strong seasonal variation.

Mytilus sp. is a habitat-modifying species that can influence biodiversity by facilitation and inhibition of species. Species diversity of macro and meio fauna greatly increased in the presence of mussel beds.

Contents of total carbon and nitrogen are noted to be significantly higher within mussel bed patches than outside, mainly due to the mussels’ filtration and biodeposition of small nutrientrich particles.Gastropod grazers including Patella and Gibbula are controlers of marine algae.

Brittlestars important for the flux of nutrients in marine ecosystems.

Strong link between structual complexity of habitat and meiofaunal/macrofaunal abundance and diversity.

Mytilus are important for sedimentation and biodeposition through the production of faces and pseudofaeces.

Species richness and biomass of associated macrofauna and flora was enhanced even by the presence of single Mytilus individuals and increased rapidly with increasing mussel numbers. This is mainly due to physical structure and creation of habitat.

Abundance and biomass of flora and fauna associated with Mytilus were higher if live mussels were present because mussel biodeposition and nutrient regeneration supplied limiting resources and increased carrying capacity.

Local - Wadden Sea

General

General

General

General

General

GeneralWave Exposure decreases with depth General

GeneralSuspended sediments increase with water currents General

General

Wave exposure is a controlling factor in benthic assemblages. General

Water currents are a controlling factor in benthic assemblages. GeneralHydroids provide a habitat to other marine life. General

The presence of Mytilus can lead to increased deposition of fine sedimentary and organic material on the sea bed because of the filtering and particle selection activities of the mussels and the reduction of current velocities by the structure of the bed. With these habitat modifying properties mussels can be regarded as physical ecosystem engineers.

Kelp biotopes, with their enormous numbers of species, high biomass and high rates of productivity are important nursery areas for a diverse range of species. It is likely that juvenile forms of all the animals that are present as adults in the kelp bed make use of the habitat as a nursery area.

The zonation patterns of the understorey algae are related to the available light rather than to the physical depth.

It is probable that kelp plants and other algae are in direct competition with the colonial tunicates, sponges etc. for space on the substratum.

The high productivity of kelp forests (mg C m-2 yr- 1) in comparison to other marine biotopes suggests that the surrounding coastal areas are dependent on the kelp biotopes as a major source of food energy. 90% of kelp production is estimated to enter the detrital food webs as POM or DOM, being exported from the immediate area of the kelp bed.

The blades of mature kelps in the kelp forest form a canopy layer which, under certain conditions, may cut off as much as 90% of the incident irradiance.

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).

Water temperature decreases with depth and chemistry may become altered.

Dissolved oxygen increases with water column mixing caused through wave exposure and water currents

Author Year Comments/Further references

Allen 1983High

Bolam 2010High

Bolam 2010High

Bolam 2010

HighCusson 2005 MediumCusson 2005 Medium

Cusson 2005Medium

Dauvin 2013Medium

Dauvin 2013 Warner 1971Medium

Gili 1997Medium

Hilly 1991High

Hiscock 2004

High

Confidence Score

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

Linder 2012

Medium

Meadows 2012 Prins et al., 1995).

High

Meadows 2012

High

Meadows 2012High

Phillipart 2012High

Pretterebner 2012Medium

Pretterebner 2012Medium

Saraiva 2011High

Saraiva 2011 Zemlys 2003High

Barros 2004

Medium

Brenchley 1982Medium

Chamberlain 2001Medium

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

Reiss 2009High

Jones 2000Medium

Jones 2000 Carter 1988Medium

Jones 2000

Medium

Vader 1984High

Daly 2008

High

Devlin 2008 High

Devlin 2009 High

Greenwood 2010 High

Hiscock 2006 High

Hiscock 2006 High

Eppley 1972 Medium

Munn 2004 Medium

Munn 2004 Medium

Munn 2005 Low

Levinton 2001 Low

Lalli 2006 Medium

Lalli 2006 Medium

Brown 2002 Medium

Nybakken 2001 Medium

Nybakken 2001 Medium

Nauderhaug & Christie 2011 High

Nauderhaug & Christie 2011 High

Nauderhaug & Christie 2011 HighNauderhaug & Christie 2011 High

Nauderhaug & Christie 2011 HighNauderhaug et al. 2012 High

Moore 1972 HighKregting et al. 2013 High

Echman et al. 1989 Medium

Echman et al. 1989 Medium

Duggins et al. 1990 Medium

Cacabelos et al. 2010 Medium

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Bell 2008 High

Wernberg & Goldberg 2008 LowWernberg & Goldberg 2008 Low

Wernberg & Connell 2008 Low

Livore & Connell 2012 Low

Airoldi & Cinelli 1997 Low

Eriksson & Bergstrom 2005 Low

Turner & Todd 1991 High

Beck 2000 Low

Aguera et al. 2012 Medium

Norling & Kautsky 2008 Medium


Boaventura et al. 2002 HighDavoult 1995 High

Davoult 1995 High

Gibbons 1988 LowLeonard 1989 Low

Kautsky & Evans 1987 Medium



Drent & Dekker 2013 Low

Birkett et al 1998 High





Masselink 2003 MediumBrown 2002b Medium

Brown 2002 MediumBrown 2002b Medium

Brown 2002 Medium

Little 1996 Medium

Little 1996 MediumPorter 2012 Medium

Code Full Reference

R1 MarLIN Biotic

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R7

Georgina Budd 2008. Alcyonium digitatum. Dead man's fingers. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 03/11/2014]. Available from: <http://www.marlin.ac.uk/specieshabitats.php?speciesID=2442>

Jacqueline Hill 2008. Antedon bifida. Rosy feather-star. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 03/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2542>

Georgina Budd 2008. Asterias rubens. Common starfish. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 03/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2657>

Nicola White 2004. Balanus crenatus. An acorn barnacle. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 03/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2718>

Ken Neal and Emily Wilson 2008. Cancer pagurus. Edible crab. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 03/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2872>

Will Rayment and Paolo Pizzola 2008. Chondrus crispus. Carrageen. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2971>

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Angus Jackson 2008. Ciona intestinalis. A sea squirt. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2991>

Georgina Budd 2007. Cladophora rupestris. A green seaweed. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=2999>

Karen Riley 2008. Clavelina lepadiformis. Light bulb sea squirt. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3009>

Dr Harvey Tyler-Walters 2008. Corallina officinalis. Coral weed. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3039>

Dr Harvey Tyler-Walters 2008. Echinus esculentus. Edible sea urchin. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3237>

Dr Harvey Tyler-Walters 2005. Electra pilosa. A sea mat. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3246>

Dr Harvey Tyler-Walters and Susie Ballerstedt 2007. Flustra foliacea. Hornwrack. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3342>

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Dr Harvey Tyler-Walters and Paolo Pizzolla 2008. Halidrys siliquosa. Sea oak. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=3447>

Dr Harvey Tyler-Walters 2007. Laminaria hyperborea. Tangle or cuvie. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=3614>

Dr Harvey Tyler-Walters 2008. Mytilus edulis. Common mussel. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 04/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3848>

Angus Jackson 2008. Ophiothrix fragilis. Common brittlestar. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3976>

Ana Ruiz 2008. Pandalus montagui. Pink shrimp. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesinformation.php?speciesID=4034>

Jacqueline Hill 2006. Pholas dactylus. Common piddock. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=4102>

Karen Riley and Susie Ballerstedt 2005. Pomatoceros triqueter. A tubeworm. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=4179>

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Angus Jackson and Dr Keith Hiscock 2008. Urticina felina. Dahlia anemone. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=4556>

Will Rayment 2004. Ahnfeltia plicata. A red seaweed. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2417>

Dr Harvey Tyler-Walters 2008. Alaria esculenta. Dabberlocks. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2431>

Angus Jackson 2008. Axinella dissimilis. Yellow staghorn sponge. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=2695>

Dr Harvey Tyler-Walters 2006. Delesseria sanguinea. Sea beech. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesfullreview.php?speciesID=3122>

Dr Keith Hiscock 2007. Eunicella verrucosa. Pink sea fan. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3317>

Olwen Ager 2008. Lanice conchilega. Sand mason. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciessensitivity.php?speciesID=3633>

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Angus Jackson 2004. Nemertesia ramosa. A hydroid. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3864>

Angus Jackson 2000. Neocrania anomala. A brachiopod. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=3879>

Jacqueline Hill 2008. Palmaria palmata. Dulse. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=4028>

Catherine Wilding and Emily Wilson 2009. Swiftia pallida. Northern sea fan. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 05/11/2014]. Available from: <http://www.marlin.ac.uk/speciesbenchmarks.php?speciesID=4407>

R. H. Millar (1970). British Ascidians. Tunicata: Ascidiacea. Keys and Notes for the Identification of the Species. Published for the Linnean Society of London by Academic Press, London and New York, 1970. Hayward, P.J. 1985. Ctenostome Bryozoans. Synopses of the British Fauna. No. 33.Southward, A.J. 2008. Barnacles. Synopses of the British Fauna. No. 57

Lock, K., Gibbs, R., Burton, M., Newman, P. 2008. Skomer Marine Nature Reserve Distribution And Abundance Of Echinus esculentus And Selected Starfish Species 2007. CCW Regional Report CCW/WW/08/2

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Schlappy, M-L., Saskov, A., Dahlgren, T.G. In Press. Impact hypothesis for offshore winf farms: Explanatory models for species distribution at extremely exposed rocky areas. Continental Shelf Research.

Jimmy, R.A, Kelly, M.S, Beaumont, A.R. 2003. The effect of diet type and quantity on the developmnet of common sea urchin larvae Echinus esculentus. Aquaculture. 220. 261-275.

Hiscock, K., Marshall, C., Sewell, J. & Hawkins, S.J. 2006. The structure and functioning of marine ecosystems: an environmental protection and management perspective.English Nature Research Reports, No 699.

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Jncc.gov.uk, (2014). LR.FLR.Rkp.FK.Sar - Sargassum muticum in eulittoral rockpools - Marine Habitat Classification. [online] Available at: http://www.jncc.gov.uk/marine/biotopes/biotope.aspx?biotope=JNCCMNCR00000764 [Accessed 27 Nov. 2014].

R344

R345

R346

Jncc.gov.uk, (2014). IR.LIR.K.Sar - Sargassum muticum on shallow slightly tide-swept infralittoral mixed substrata - Marine Habitat Classification. [online] Available at: http://www.jncc.gov.uk/marine/biotopes/biotope.aspx?biotope=JNCCMNCR00002080 [Accessed 27 Nov. 2014].

Little, C., Kitching, J. A. 1996. The Biology of Rocky Shores. Oxford University Press: Oxford.

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

Abstract

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.

The increasing need for renewable and clean energy production is likely to result in a diversi fication of locations for the implementation of offshore wind farms which have been so far predominantly sited on soft substrata. In contrast, offshore wind turbines placed on rocky reefs in highly exposed areas are much less common and the impacts on localflora and fauna can only be 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 rchinpopulations. 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 information about the potential impacts of the construction of an offshore wind farm on rocky substratum and allows us to generate impact hypotheses that go beyond what could behypothesized through a mere general knowledge of the area.

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.

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.

This paper evaluates the effect of the tube-building, habitat structuring polychaete Lanice conchilega on the macrobenthic community and sediment characteristics of its habitat. To investigate which factors make species occur in a well-known bio-engineered habitat, macrofaunal and sedimentological data, gathered over a period of 10 years in a shallow, fine sediment bottom of the Belgian Part of the North Sea, were submitted to analyses. Both sediment composition and community structure of the associated macrofaunal matrix were affected by the presence of L. conchilega. The effect of the protruding tubes on hydrodynamics clearly resulted in the retention of fine sediment particles, while the increased coarse fraction was assumed to reflect a dynamic population build-up. This study confirmed that tube aggregations of L. conchilega expand the realized niche of several species without forming their own association. A species rank list was created according to each species’ association with L. conchilega. This species rank list is extensively discussed based on all ecological knowledge available. Species are favoured by the habitat modifying ability of the polychaete tubes, which create and regulate refuge for species, alter the interactions between local species and change the physical factors of the environment. This descriptive and correlative data study examines the ecological importance of the bioengineer L. conchilega on species level.

A laboratory flume study was conducted to determine the effect of flow velocity on clearance rates of the Polychaeta Terebellidea Lanice conchilega (Pallas, 1966). Using sets of 75 individuals collected along the east coast of the English Channel, we measured clearance rates at five flow velocities of ≈4, 9, 15, 22 and 27 cm s−1 with a culture of the diatom Chaetoceros calcitrans as food source. During each control (without polychaetes) or clearance (with Lanice conchilega) experiment, in vivo fluorescence was continuously monitored. Further HPLC analysis confirmed the sediment resuspension for the highest flow velocities tested, indicating the need for 'corrections' from control experiments. The global pattern resulted in a dome-shaped curve, with a corrected clearance rate per individual increasing with flow velocity from an average value of 0.091±0.041 L h−1 gDW−1 at 4 cm s−1 up to a maximal value of 0.171±0.046 L h−1 ind−1 at 15 cm s−1 and decreasing for higher flow velocity (0.063±0.029 L h−1 ind−1 at 27 cm s−1). When normalised to average Dry Weight (DW), the clearance rates varied in a wider range, with maximal clearance rates of up to 0.749 L h−1 gDW−1 and a marked dome-shaped structure for small individuals (11.9±5.4 mg Ash Free DryWeight), whereas larger individuals (up to 30 mg AFDW) showed maximal clearance rates of around 0.400 L h−1 gDW−1. Our results indicate the need to take flow velocity into account in estimating the ecological impact of filter-feeding polychaetes on available food, as large budget underestimates may arise from clearance rates measured in non-controlled flow velocity.

As part of a long-term study to examine the ecological effects of beam-trawling, we 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.

Epibiont communities of Pagurus bernhardus and its inhabiting shell collected in the same month of two consecutive years near the 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 environmental parameters on these differences is considered

This paper is an account of Serpulidae (Annelida, Polychaeta) collected from the Arctic and Northern Atlantic Oceans by Soviet expeditions. This material is part of the collection of the Department of Hydrobiology at Moscow State University and the Zoological Museum of the Russian Academy of Sciences in St. Petersburg. We redescribe eleven species: Chitinopoma serrula, Ditrupa arietina, Filograna implexa, Hyalopomatus claparedii, Hydroides norvegicus, Placostegus tridentatus, Pomatoceros triqueter, Protula globifera, P. tubularia, Protis arctica, and Serpula vermicularis. A new species from this collection, Metavermilia arctica, was described in a previous publication. Original illustrations and distribution maps accompany all descriptions. We provide two separate taxonomic keys to specimens and tubes. The previous records of arctic serpulids and the present state of the taxonomy of the family are discussed. Based on the operculum-bearing specimens of Protis arctica found, and on previous similar records, the taxonomic significance of the operculum in serpulids is further re-evaluated. As a result, Salmacina and Apomatus are considered synonyms of Filograna and Protula, respectively. We believe that many of the species and genera considered in this paper need revision.

Dyopedos monacanthus (Metzger) and L). porrectus Bate secrete numerous threads of mucus from special setae and a solid spinning thread from each of the pereopods 3-4. The threads are used for the construction of a mast-like structure on the bottom out of sediment, seston, and faecal pellets, and for h~dling collected seston. The mast serves Dyo~e~o~ to reach a higher level with more rapid current. Dyopedos feeds mainly on seston, which it sieves from the current with its richly setose antennae. The mast is a territory. Enlarged second gnathopods and enhanced aggressiveness in the males are manifestations of territorial behaviour and hierarchism. The two species tolerate the close presence of each other when on their awn masts but do not tolerate each other on the same mast. Methods of mast-building, locomotion, and feeding, and features of functional mo~holo~ and behaviour are described. Dyopedos monacanthus accepts weaker currents than D. porrectus. According to data from fish stomach contents, the two species were nearly equally common in the area studied and had a high degree of distributional overlap. From April to October, both species recruited, attained their highest abundance, and were most intensely preyed upon by fish. Dyopedos was found in 1704 of the fishes examined and made up 9% of the total number of food individuals in them. Observations suggest that Dyopedos defends itself against a predatory fish by quickly descending from its high feeding position and clutching the mast tightly lengthwise. This would make it impossible for the fish to ingest it without also ingesting a section of the mast, which is practically devoid of nutritive substance. By thus reducing the nutritional value of the potential morsel, Dyopedos would make itself less attractive as prey. Dulichia tuberculata Boeck probably also builds masts but, in contrast to Dyopedos, can also sit directly on the bottom when feeding. Like Dyopedos, it collects seston from the current with its antennae, the structure ofwhich suggests that aesthetascal mucus threads, rather than setae, are used for seston-collecting by this species.

In the framework of a wider research programme aimed at identifying the functioning and the temporal dynamics of seagrass ecosystems along the Italian coasts of the Adriatic Sea, crustacean amphipods were studied in Zostera marina, Cymodocea nodosa and Posidonia oceanica beds. Data on Z. marina, which is very rare in the Mediterranean Sea and is scarcely studiedas it concerns amphipods, are of particular interest. Samples were collected by hand net and air-lift sampler in Z. marina (- 0.5 m) and C. nodosa (- 1.2 m) at Grado (northern Adriatic Sea), and in P. oceanica (- 6.5 m) at Otranto (southern Adriatic Sea). First data from ten samples collected in February and May 1997 are given. Fifty-five species (Z. marina: 24; C. nodosa: 29; P. oceanica: 31) in 41 genera and 25 families belonging to 7324 individuals were identified. The cluster and nMDS analyses showed the presence of different assemblages in the three seagrasses. The lowest average dissimilarity was present between Z. marina and C. nodosa (43.13: air-lift sampler; 51.66: hand net) and the higher between Z. marina and P. oceanica (81.36: air-lift sampler), and between C. nodosa and P. oceanica (77.41: hand net). The species which are dominant and/ or discriminate each assemblage are Dexamine spinosa, Perioculodes aequimanus and Gammarus insensibilis in Z. marina, Metaphoxus fultoni, Ampithoe helleri and Ericthonius difformis in C. nodosa, and the small sized Apolochus neapolitanus, Peltocoxa marioni and Stenothoe monoculoides in P. oceanica. The features of the sampling sites and the plant structure play an important role in defining assemblages.

Recruitment ofepifaunal invertebrates was followed on 16 x 16 cm Perspex panels immersed at a low imertidal site at St Andrews, on the east coast of Scotland. Short-term experimental enclosures assessed the importance of grazing by the herbivorous prosobranch Gibbula cineraria (L.), and the nontrophic activities of the predatory species Nucella iapiii~ts (L.) and Asterias rubens L., on developing epifaunal assemblages. At the end of each experimental period, recruits of all seven taxonomic groups examined were less abundant on G. cineraria grazed treatments, than on those from which grazers were excluded. Furthermore, none of the epifauna were apparently able to achieve an escape-in-size during a 2-month period of exclusion of grazers. N. lapillus and A. rubens did not appear to have similar deleterious effects on the developing assemblages (with the possible exceptions of barnacles, hydroids and ascidians during periods of peak larval settlement). Similar numbers of recruits were recorded on the treatment panels and the net controls. Enclosure of N. lapillus appeared to enhance the recruitment of some species (e.g., anomiids). These experiments provide no information regarding the nature of the effects ofthese consumers on the developing assemblages. However, there is circumstantial evidence that the effects of G. cineraria may be attributable to the rasping-off of recently established recruits, as the snails browse across the substrata.

ABSTRACT: Many bryozoans, exemplified by species of Membranipora, are obligate epiphytes. We used laboratory and field experiments to ascertain whether colonies of Mernbranipora isabelleana benefit from contact with the fronds and exudates of Lessonia trabeculata, their natural substratum in central Chile. Type of substratum, presence or absence of microalgal food and of L. trabeculata fronds were the experimental factors, while colonial growth and percentage of non-degenerated zooids were the measured response variables. Colonies on L. trabeculata fronds did not grow faster than thoseattached to glass slides, either in the laboratory or in the field Colonies grown for l moon fronds, however, retained a significantly higher proportion of non-degenerated zooids than colonies grown on slldes (88 and 73 % respectively in the laboratory; 100 and 84 % respectively in the field). When grown on slides, without microalgal food, colonies retained 79% of non-degenerated zooids in the presence of fronds but only 28% in their absence. These results suggest that colonies absorbed algal exudates when fronds were present, thus preventing the early degeneration of zooids. Even when fronds were separated from colonies by borosilicate membranes, allowing only the diffusion of molecules smaller than 1200 daltons, the above effect was still pronounced. In the field, colonies on fronds had a higher survival rate than those on slides. Association with an algal host therefore enhances at least 2 important life-history parameters in M isabelleana: the percentage of non-degenerated zooids and colonial survivorship. Our results suggest that the trophic association with seaweeds is likely to be particularly important to epiphytic bryozoans under conditions of reduced particulate-food concentration.

The occurrence of encrusting colonies of the hydroid Obelia geniculata on farmed Saccharina japonica was examined between December 2007 and July 2008 at a Saccharina farm at Wando on the southwestern coast of Korea. The growth stages of S. japonica can be divided into two phases: an active growth phase from February to the end of May and a decay phase from June to July. There was a significant increase in the level of incrustation by colonies on fronds (measured as the percentage of fronds with encrusting colonies) between February and July (p<0.05). The encrusting colonies occurred first on the upper part of the frond in February and progressed to the basal part in July. The abundance of encrusting colonies in relation to the growth phase on farms over time was limited by the harvest of the seaweed crop at the end of the cultivation period in July. The stipes and holdfasts of fronds showed no signs of infestation at any time during the cultivation period. The extent of the infestation appeared to be related to a combination of factors. These could be reduced physiological activity and subsequent tissue aging that occurred simultaneously in the sporophytic life phase of Saccharina frond, and a rapid increase in reproduction and growth of O. geniculata coinciding with rising seawater temperature.

Estimates of secondary production (P/B ratio and total production) by macrobenthic communities across the UK continental shelf are 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 of the variation in production estimates was not explained by the present study are presented.

No consistent -functional grouping of organisms as stabilizers vs destabilizers, respectively decreasing or enhancing credibility, is possible . Benthic organisms can affect erodibility in particular-and sediment transport in general-via alternation (1) of fluid momentum impinging on the bed, (2) of particle exposure to the flow, (3) of adhesion between particles, and (4) of particle momentum. The net effects of a species or individual on erosion and deposition thresholds or on transport rates are not in general predictable from extant data . Furthermore, they depend upon the context of flow conditions, bed configuration, and community composition into which the organism is set. Separation of organism effects into these four categories does, however, allow their explicit incorporation into DuBoys-type and stochastic sediment dynamic models already in use and thus permits the specification of parameters whose measurement will enhance predictability of sediment transport modes and rates in natural, organism-influenced, marine settings. If the variable of prime concern is the total amount of sediment transported, rather than the frequency of transport events or the spatial pattern of erosion and deposition, and if most transport occurs in rare but intense bouts (e .g ., winter storms on boreal continental shelves), then it may be possible to ignore organism effects without major sacrifices in accuracy or precision. Under high transport rates, suspended load effects override organism-produced bottom roughness, abrasion removes adhesives from transporting grains, and transport rates (normalized per unit width of the channel or bed) exceed feeding and pelletization rates. Moreover, at high rates most material transports as suspended load, effectively out of reach of the benthos. The transport rates at which organism effects are overridden, however, remain to be determined. For lower transport rates, foraging theory promises to provide insights into organism effects.

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 organicmatter 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 advectiondominated 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.

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.

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.

Relationships between benthic macrofauna and natural abiotic factors were studied along the coastal fringe of South Brittany, situated north of the Gulf of Biscay on the French Atlantic continental shelf. Within the framework of the REBENT network, sediment characteristics, depth, and macrofaunal abundance were determined for 95 stations spread over five subtidal sectors, using a combination of seabed acoustic remote sensing systems and grab sampling. The physico-chemical properties of the water column and the hydrodynamic conditions were generated by validated three-dimensional environmental models which take into account variations over shorter temporal scales. Multivariate analyses ranked 16 natural abiotic variables according to the significance of their influence on the macrofauna. Together these variables explained 51% of spatial variation in the macrofauna, with morpho-sedimentological and hydrological factors contributing 22% and 26%, respectively. The outputs from validated three-dimensional environmental models appear to be useful interpretational tools for benthic ecology studies, especially in estuarine and coastal ecosystems with high environmental variability due to regular freshwater inputs. Ten major species assemblages were identified using biological and physical characteristics. The results provide important baseline knowledge for future ecosystem and resource management.

Benthic organisms appear to be accurate proxies for assessing coastal ecosystem structures and changes due to climatic and anthropogenic stresses. Functional studies of benthic systems are relatively recent, mainly because of the difficulties in obtaining 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 connected through hydrodynamics, a model of the entire eastern basin of the English Channel would appear to be acceptable. However, the main sediment types must be taken into account when establishing relationships between the functional units.

Benthic suspension feeders are abundant in littoral and shallow sub-littoral ecosystems, where they feed on the plankton and on organic matter suspended in the water column. Recent studies indicate that active suspension feeders with powerful water filtration mechanisms (e.g., bivalve molluscs) may exert an important influence on the abundance and production of phytoplankton, and probably zoo-plankton as well. Passive suspension feeders, such as hydrozoans, have received less attention, and their effect on shallow planktonic communities is poorly understood. This paper presents evidence that hydrozoans, which make only a minor contribution to benthic community biomass, capture large amounts of zooplankton and seston, and that they may play an important role in transferring energy from pelagic to benthic ecosystems over a wide range of latitudes.

The expectation that long-term, broad-scale changes in the relative abundance of species, homogenisation of habitats and decreases in diversity will affect ecosystem function has led to an increasing number of studies on functional diversity and composition. Such studies frequently consider the effect of environmental gradients and anthropogenic impacts, but rarely the effect of biogenic habitat variation. In marine soft sediment systems, habitat variability is likely to be of particular importance because of the strong link between habitat and species diversity. In this study we examine the link between functional trait diversity (as richness and evenness) and composition, and habitat variation in two locations with different regional species pools. We found similar functional traits occurring in the two locations, but differences between habitats within the locations. High evenness within traits was apparent (across both locations and habitats) reflecting the potential for the maintenance of function with the loss of individual species. Between-habitat differences in functional traits were driven by differences in organism densities rather than the presence/ absence of individual traits, emphasising the importance of density shifts in driving function. Furthermore, our demonstration of habitat variation as a driver of functional composition and diversity suggests that habitat heterogeneity should be explicitly included within studies trying to predict the effect of species loss on ecosystem function.

In the Bay of Brest, France, suspension feeders are broadly dominant in the trophic structure of macrobenthic assemblages in terms of species number, number of individuals and biomass. A succession of species exists on an east-west gradient, which is an ecological continuum from a < 10 m depth polyhaline muddy environment to a > 20 m depth marine environment with coarse and clean sediments. The abundance of biogenic substrates (shells, calcareous algae, etc.) on the sediment surface favours a high development of sessile epifauna. Some species, such as Crepidula fornicata, Ascidiella aspersa, Ficulina ficus, Ophiothrix fragilis and Ophiocomina nigra, can locally proliferate, reaching 100 g ODW (organic dry wt) m-2. Fertilization of the bay by streams loaded with nutrients induces a very high primary production (280 g C m-' yr-l) characterized by high annual and seasonal variability of phytoplankton blooms. However, so far, eutrophication has not been observed. While water renewal by tidal currents is one important factor in the control of eutrophication, suspension feeder activity is undoubtedly another effective control factor. Benthic suspension feeders can filter 7.18 X 10' m3 daily, which represents ca 30% of the total volume of the bay. The phytoplankton production is easily accessible to the benthos because of the relatively long water residence time and the high tidal current mixing. Spatial distribution of populations was found to be related to the tidal current distribution.

The coastal waters surrounding Britain and Ireland became warmer during the 20th century and, according to the UK Climate Impact Programme 2002 scenarios of change and other sources, average annual seawater temperatures may rise a further 28C or more by the 2050s. This warming is part of a global rise in sea- and air-surface temperatures that will cause changes in the distribution 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 good candidates to track change. Using the key, many species are shown as likely to increase their range northwards significantly. In contrast, fewer will decline in abundance and extent in the north. If, as anticipated, global warming continues, then species with distributions already accurately mapped, or being mapped at present, will provide baseline data to test forecasts.

The influence of bed density (from 0 to 1000 ind.m−2) and flow speed (2, 5, and 15 cms−1, i.e. “low”, “medium” and “high”, respectively) on clearance rates (CR) of an infaunal suspension-feeding bivalve (Austrovenus stutchburyi) and boundary layer dynamics were investigated in annular flumes. Near-bed flow speeds decreased and bed shear stresses increased with bivalve density due to the bed roughness generated by the shallow burrowers. At low densities (b500 ind.m−2) there was no detectable effect of feeding currents on bed shear stress, but at high densities (1000 ind.m−2) andat the lowflowspeed bed shear stress was almost 80% higher when Austrovenus were feeding (0.0016 N m−2), compared to when they were not feeding (0.0009 N m−2). These results suggest that bed roughness generated by Austrovenus and their feeding currents are capable of influencing food supply to the bed. Individual CRwere significantly (pb0.01) greater at the high flow speed (0.88 L h−1 ind.−1), compared to the medium and low flow speeds (0.34 to 0.54 L h−1 ind.−1). With increasing bed density, individual CR was significantly (pb0.001) reduced; at low densities CR ranged from 0.6 to 1.4 L h−1 ind.−1, but at high densities they were 0.1 to 0.8 L h−1 ind.−1. Bed CR (L h−1 m−2) did not scale proportionally with density, initially increasing up to 500 ind.m−2 but remaining relatively constant thereafter. This was in part due to the negative effect of density on individual CR but also because the proportion of bivalves feeding decreased with increasing density. Our results demonstrate that recording the number of bivalves feeding is necessary to interpret the effect of density on bed CR. Moreover, if the effects of both bivalve density and flow speed are not considered, then scaling up individual CR obtained using animals at low densities will over estimate population filtration capacity. As high (but equivalent to in situ) densities reduced both the number of animals feeding and individual CR other factors may offset this negative interaction, such as protection from wave disturbance or enhanced resuspension of benthic microalgae.

The present paper reviews the current knowledge on diagenetic carbon transformations at the oxic/anoxic interface in coastal marine sediments. Oxygen microelectrodes have revealed that most coastal sediments are covered only by a thin oxic surface layer. The penetration depth of oxygen into sediments is controlled by the balance between downward transport and consumption processes. Consumption of oxygen is directly or indirectly caused by respiration of benthic organisms. Aerobic organisms have the enzymatic capacity for complete oxidation of organic carbon. Anaerobic decay occurs stepwise, involving several types of bacteria. Large organic molecules are first fermented into small moieties. These are then oxidized completely by anaerobic respirers using a sequence of electron acceptors: Mn4+, NO3-, Fe3+, SO4 2- and CO2. The quantitative role of each electron acceptor depends on the sediment type and water depth. Since most of the sediment oxygen uptake is due to reoxidation of reduced metabolites, aerobic respiration is of limited importance. It has been suggested that sediments contain three major organic fractions: (1) fresh material that is oxidized regardless of oxygen conditions; (2) oxygen sensitive material that is only degraded in the presence of oxygen; and (3) totally refractory organic matter. Processes occurring at the oxic/anoxic boundaries are controlled by a number of factors. The most important are: (1) temperature, (2) organic supply, (3) light, (4) water currents, and (5) bioturbation. The role of bioturbation is important because the infauna creates a three-dimensional mosaic of oxic/anoxic interfaces in sediments. The volume of oxic burrow walls may be several times the volume of oxic surface sediment. The infauna increases the capacity, but not the overall organic matter decay in sediments, thus decreasing the pool of reactive organic matter. The increase in decay capacity is partly caused by injection of oxygen into the sediment, and thereby enhancing the decay of old, oxygen sensitive organic matter several fold. Finally, some future research directions to improve our understanding of diagenetic processes at the oxic/anoxic interface are suggested.

Within transitional/estuarine environments ‘ecosystem functioning’ has been mostly investigated with “traditional” taxonomic analysis, based on the taxonomic composition of benthic invertebrate communities. However, ‘ecosystem functioning’ depends also greatly on the functional characteristics (biological traits) of organisms. It was a priori suggested that the biological traits of the subtidal benthic invertebrate communities within an estuarine environment would respond to the high variability of environmental pressures (natural and human induced) within this type of ecosystem. For this study, traditional taxonomic analysis (species richness, species density and Shannon–Wiener diversity) as well as biological trait analysis were used together for the first time to investigate the response of the subtidal benthic invertebrate communities to the environmental pressures within the Mondego estuary (Portugal). Biological trait analysis, in addition to traditional taxonomic analysis provided a more comprehensive understanding of the functioning within this type of ecosystem. Some of the most important outcomes are: (i) the trait “salinity preference” was the most important trait that distributed the species along the estuary, (ii) the central part of the estuary appeared to be under higher environmental stress levels than the other areas, as suggested by a dominance of some “opportunistic” traits (e.g. small short-lived species), (iii) the ratio between functional diversity (FD) and Shannon–Wiener diversity (H_) indicated lower functional redundancy at the upper reaches of the estuary. Our results, suggest that the ratio (FD/H_) might be a helpful tool to visualize this functional attribute and could potentially be applied to different communities from distinct environments. Using the traditional taxonomic analysis alone, this last functional aspect would not be detectable. Therefore, the inclusion of biological traits analysis is recommendable for estuarine ecological studies.

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.

Fauna have been found to regulate important biogeochemical properties and ecosystem functions in benthic environments. In this study, we focused on how functional biodiversity and species-specific traits of benthic macrofauna affect key ecosystem functions related to organic matter mineralization and cycling of nutrients in surface sediments. Dominant benthic invertebrates from the Baltic Sea and the Skagerrak were classified into functional groups in accordance with their behaviour, feeding and sediment reworking activities. Macrofauna species were added in different combinations to defaunated Baltic sediments in 2 parallel microcosm systems fuelled with brackish and marine water. In total, there were 12 treatments that differed in terms of functional diversity of benthic fauna. The experiments demonstrated that faunal activities directly affected benthic oxygen and nutrient fluxes, sediment reactivity and pore-water distribution under both Baltic and Skagerrak conditions. Benthic fluxes, sediment reactivity and pore-water distribution were similar in Baltic and Skagerrak treatments, in which the same functional biodiversity and species-specific traits of benthic macrofauna were observed. Although no significant effects of functional biodiversity could be detected under Baltic or Skagerrak conditions, treatments with bioturbating fauna from the Skagerrak enhanced oxygen consumption and nutrient fluxes compared to treatments with Baltic fauna and Skagerrak fauna with functional groups similar (parallel) to the Baltic fauna. Moreover, species specific traits related to the Skagerrak fauna (e.g. the thalassinid shrimp Calocaris macandreae) exceeded the effects of all other faunal treatments. This suggests that species-specific traits of macrofauna may override species richness and functional biodiversity of macrofauna when regulating important ecosystem properties and functions in benthic environments.

The Northern Hemisphere has been warmer since 1980 than at any other time during the last 2000 years. The observed increase in 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 essential to formulate and inform better adaptive strategies to address the inevitable consequences of climate change.

Hermit crabs play an important role in the Northern Adriatic Sea due to their abundance, wide range of symbionts, and function in structuring the benthic community. Small-scale (0.25 m2) hypoxia and anoxia were experimentally generated on a sublittoral soft 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 relative tolerance of crabs and certain symbionts (e.g. the sea anemone Calliactis parasitica) – as potential survivors and recolonizers of affected areas – may influence and promote community recovery after oxygen crises.

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.

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.

The relationship between biological community structure and particle size composition is investigated in coastal deposits off the southeast of England. Sediments in the survey area fall into well-defined groups when analysed by multivariate techniques, indicating similarities and differences which could not be identified by mere inspection of the data. Biological resources also fall into relatively distinct groups, or communities, when analysed for species composition and population density, although similarity within the groups is lower than that obtained for the sediments. There is, however, little evidence of a close correspondence between the distribution of different sediment types and benthic communities in the survey area: comparison of the similarity matrices yields weighted Spearman rank correlation of less than 0.37. This suggests that factors other than sediment composition play a significant part in controlling biological community structure on the seabed. Still, there is evidence that some species such as the tube-dwelling worm Sabellaria spinulosa are associated mainly with sands and gravels whilst fine mobile silts and sands are characterized by ‘‘opportunistic’’ species such as the tube-worm Lagis koreni. The results suggest that although modification of sediment composition from mixed sands and gravels to silts would be expected to result in colonization by ‘‘opportunistic’’ species capable of survival in mobile deposits, restoration of sediment composition after cessation of dredging for marine aggregates is not, within broad limits, a prerequisite for establishment of biological communities which are comparable with those that occurred in the deposits prior to dredging.

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.

Dense beds of the suspension-feeding brittle-star Ophiothrix fragilis are common in European waters. Their potential importance in benthic-pelagic coupling has been highlighted, but little is known about the feeding dynamics of this species. Encounter-rate models provide a potential mechanism for the estimation of feeding rates on suspended material of varying sizes. This work investigates factors essential to the application of such models. Particle-retention efficiency (RE) converts encounter rate into capture, or clearance rate. Laboratory studies demonstrated that RE varied with the interactive effects of flow velocity and particle size. RE was lowest for large particles, particularly at high flow velocity where RE as low as 59% was observed. This indicates that if RE is not accounted for in encounter-rate models, significant overestimates of feeding rates on large particles may occur. Flow around feeding arms and tube feet was characterised by intermediate Reynolds numbers, precluding application of the most simple encounter-rate models. Complex secondary-flow patterns were observed, which carried particles along the downstream side of the feeding arms, but these did not appear to increase the area available for particle capture. Previously reported particle capture by arm spines was not observed. Evidence of active rejection of large particles by tube feet was recorded. Difficulties in the application of encounter-rate models for prediction of seston-removal rates are highlighted by these results. Predicted encounter rate may deviate from actual clearance rate due to the effects of retention dynamics, localised flow patterns and differential particle handling. Other methods of estimation of seston-removal rates are equally problematic however, so that encounter-rate models are likely to remain a useful tool for such estimates.

This study tested the hypothesis that the small-scale topography caused by ripples in sediment would affect benthic macrofaunal assemblages. Ripple marks were measured and macrofauna and sediments were sampled near to and far from subtidal rocky reefs at two sites. Ripples were significantly wider and taller close to than far from reefs. Sediment grain-size was significantly different between crests and troughs. Microtopography clearly influenced the structure of the benthic macrofaunal assemblages in three of the four locations (and 8 of the 12 sites) examined. Numbers of taxa of the benthic macrofaunal assemblages and almost all individual taxa analysed showed significant greater abundances in troughs than in crests. The results strongly support the model that benthic macrofauna are affected by locally varying hydrodynamic environments produced by ripple-beds. These, in turn, were influenced by their proximity to a reef. Models about passive transport of the macrofauna by water-movement and active movements of the fauna are discussed. Furthermore, it appeared that there was a relationship between spatial variability of the macrofaunal assemblages and size of ripples. It is suggested that microtopography should be considered in experimental designs when patterns of distribution of benthic organisms are being evaluated.

Biological dinitrogen (N2) fixation is the primary input of fixed nitrogen (N) into the marine biosphere, making it an essential process 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 marine N and carbon (C) cycles.

The effect of flow and biodiversity on ecosystem functioning was investigated in an estuarine system using in situ benthic chambers. Macrofaunal communities were artificially assembled to manipulate both species richness and functional trait richness. In addition, naturally occurring communities were sampled in order to determine the effect of macrofaunal and sediment disruption. Ecosystem functioning was assessed by measurement of nutrient release (NH4–N) from the sediment, a process essential for primary production. Natural and assembled communities were found to differ significantly, demonstrating the effect of experimental manipulation on the system. Flow was found to have a highly significant effect on ecosystem functioning in both natural and assembled communities in treatments containing macrofauna. No significant difference between static and flow treatments was found in macrofaunal-free controls, indicating that flow generates an effect through promoting changes in bioturbatory activity of the infauna causing greater disruption to the sediment. In assembled communities, functional richness significantly increased ecosystem functioning. Species richness had no influence in assembled communities.

Two kinds of biogenic structures, roots of seagrasses and tubes of invertebrates, were found to reduce the mobility of a variety of burrowing species, including two polychaetes (Abarenicola pacifica Healy & Wells, A. claparedii Healy & Wells), two bivalves (Macoma nasuta Conrad, Clinocardium nuttallii Conrad), two crustaceans (Upogebia pugettensis (Dana), Callianassa californiensis Dana), a holothurian (Leptosynapta clarkii Heding), and an echinoid (Dendraster excentricus (Eschschohz)). Comparisons of burrowing in similar sediments showed that: (a) all species had about a two-fold increase in mean time to burial in dense tube-mats of polychaetes and crustaceans; (b) hard-bodied taxa were particularly restricted by roots of Zostera marina L. and were often incapable of penetrating them; (c) roots and tubes together had the greatest impact, with mobility 3 to 37 times slower; and (d) burrowing ability increased when sediments were “pre-burrowed” for I day. Comparisons of burrowing rates in different sediments showed effects of physical origin (sediment compaction or sorting). Overall these physical effects were second to those of purely biogenic origin, sediment binding and cohesion by roots and tubes. Nonetheless, physical effects exceeded those of extremely high densities of tube-builders (3160 individuals . 0.01 m-?). In all substrata the movement of larger individuals, and the larger of morphologically similar pairs of species (listed second above), were more restricted by roots and tubes than were smaller ones. The results suggest a general mechanism by which dense assemblages of tube-builders and seagrasses may restrict older and larger burrowing organisms. Because physical properties of the sediment and the abundance of other burrowers are also factors affecting mobility, it is not now possible to predict what situations will render this mechanism of interaction important to natural communities.

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.

Suspension-feeding bivalves play an important role in coastal ecosystems by affecting near-bed hydrodynamics and, subsequently, rates of biodeposition. We designed a high-resolution field study to investigate rates of sedimentation and biodeposition around individuals within beds of the large pinnid bivalve, Atrina zelandica, and to link these rates with sediment biogeochemical characteristics and macrofaunal community structure. The study was conducted at three sites arrayed along a gradient of increasing suspended seston concentration, enabling us to assess changes in the strength of the Atrina–macrofauna interaction with background sedimentation. Sedimentation rates and inputs of organic carbon and nitrogen were higher close to individual Atrina (#10 cm) compared to further away (.30 cm). Seafloor sediments nearer Atrina were enriched in carbon and nitrogen and had more diverse and abundant macrofaunal assemblages. The strength of this interaction decreased with increasing background sedimentation. Although other mechanisms, both biotic and abiotic, may explain some of these patterns, we have identified variations in macrofaunal community structure that at least partly can be linked to site-specific sedimentation, at the small scale of single Atrina, nested within the larger landscape scale of Atrina beds, emphasizing Atrina’s role in habitat modification, both structurally and functionally.

Temperature-induced variations in bioturbation could affect sediment mixing processes in the marine benthic environment. In this study, sediment reworking by Neanthes virens (Sars), a widely distributed polychaete in muddy sand communities of northern temperate latitudes, was studied under different temperature conditions representing winter (1°C), spring and fall (6°C), summer (13°C), and tide pool (18°C) temperatures in the lower St. Lawrence Estuary, Québec, Canada. Sediment reworking was quantified using inert fluorescent particles (luminophores) deposited at the sediment surface. Based on the 1-D luminophore distributions obtained after 5 and 30 d, the use of the specific ‘gallery-biodiffusor’ model allowed us to quantify both biodiffusion (Db) and biotransport (Vb) due to the organisms. Our results showed temperature effects on sediment transport. The lowest biotransport and biodiffusion coefficients were measured at 1 and 6°C and did not change with time. The highest biodiffusion occurred at 13°C for both sampling periods. At 18°C, biodiffusion was intermediate while biotransport was maximal. Differences between the 13°C biodiffusive transport and the other temperatures increased with time. Low transport values at 1 and 6°C suggest that a quiescent stage exists for this species at these temperatures, with sediment mixing occurring mostly during burrow construction. On the other hand, sediment mixing resulted from both the burrow construction and maintenance phases at higher temperatures (13 and 18°C).

Understanding the structure and interrelationships of North Sea benthic invertebrate and fish communities and their underlying environmental drivers is an important prerequisite for conservation and spatial ecosystem management on scales relevant to ecological processes. Datasets of North Sea infauna, epifauna, and demersal fish (1999–2002) were compiled and analysed to (i) identify and compare spatial patterns in community structure, and (ii) relate these to environmental variables. The multivariate analyses revealed significantly similar large-scale patterns in all three components with major distinctions between a southern community (Oyster Ground and German Bight), an eastern Channel and southern coastal community, and at least one northern community (.50 m deep). In contrast, species diversity patterns differed between the components with a diversity gradient for infauna and epifauna decreasing from north to south, and diversity hotspots of demersal fish, e.g. near the major inflows of Atlantic water. The largescale hydrodynamic variables were the main drivers for the structuring of communities, whereas sediment characteristics appeared to be less influential, even for the infauna communities. The delineation of ecologically meaningful ecosystem management units in the North Sea might be based on the structure of the main faunal ecosystem components.

How do small, benthic meiofaunal organisms become cosmopolitan over large geographic ranges? Abiotic forces including oceanic currents are believed to be of key importance in aiding marine meiofaunal dispersal. We investigated the effect of distance from substrate and site exposure on meiofaunal colonization and transport in the water column. First, we tested how distance from substrate and sediment grain size affected colonization of azoic sediments by meiofauna in a sheltered inlet. Nematodes, crustacean nauplii and small amphipods colonized distant sediment cages (3 m above bottom) as quickly and abundantly as cages closer to the ocean floor. Some of the 30 recorded genera of nematodes predominated in one height treatment whereas abundance of others was not related to distance from the substrate. Polychaetes and harpacticoid copepods colonized near-benthic cages more rapidly and abundantly than those farther away suggesting active dispersal. Nematodes, harpacticoids and polychaetes were more abundant in fine than in coarse sediments, while nauplii and amphipods did not differ in abundance between sediment types. In part two of this study, we surveyed occurrence of meiofauna in the water column at several sheltered and exposed sites using plankton nets towed at fixed distances from 0.5 to 6.5 m above the ocean floor. Because oceanic currents increase sediment suspension and transport, we expected to see more meiofauna in samples collected from exposed than from sheltered sites. However, with the exception of polychaetes, which were more abundant in the water column of sheltered sites, there was no difference in meiofaunal abundances between the two exposure classes. Meiofauna, including the 14 identified nematode genera, were collected in greatest numbers nearer to the ocean floor and dwindled further up in the water column. The presence of meiofauna high in the water column of even the most sheltered sites combined with the quick and abundant colonization of distant, sheltered sediment cages suggests that even very weak currents are sufficient to suspend and transport these animals or that many meiofaunal taxa are capable of active dispersal into the water column.

The common tube-building polychaete Lanice conchilega is known as a habitat structuring species and can form dense aggregations. The eVects of L. Conchilega on the surrounding benthic community have received little attention, especially in subtidal areas. Therefore, the presence of L. conchilega in diVerent habitats in the North Sea and its eVect on the abundance, species richness, diversity and community structure in these habitats are evaluated in the present paper, based on data from the ICES North Sea Benthos Survey of 2000. Lanice conchilega has a wide geographical distribution and a low habitat specialization, but optimally occurs in shallow Wne sands. In the present study, the presence of L. conchilega resulted in a density increase and a signiWcant (positive) correlation of the benthos density with the density of L. conchilega. Furthermore, the species richness (number of species) increased with increasing density of L. conchilega. This trend was, however, not consistent: the number of species reached more or less an asymptotic value or even decreased after reaching a critical density of L. conchilega (>500–1,000 ind/m²), as observed in shallow Wne sands. The same overall pattern was detected concerning the expected number of species. The N1-diversity index showed similar or slightly higher values in L. conchilega patches compared to patches without L. conchilega. From the results of the community analysis, it can be concluded that the species, which were responsible for the increase of the diversity, belonged to the overall species-pool of that habitat. The eVects on density and diversity diVered between the four discerned habitats (shallow muddy sand, shallow Wne sand, shallow medium sand and deep Wne sand), and were most pronounced in shallow Wne sands. These patterns can be attributed to the habitat structuring capacity of L. conchilega. The mechanisms responsible for the increase of the habitat quality in patches of L. conchilega can be summarized as (1) changes in the hydrodynamics, (2) increases of the habitat stability and oxygen supply, and (3) a creation of habitat heterogeneity in a uniform environment. In this way, L. Conchilega alters the habitat characteristics and aVects other organisms, and can therefore even be considered as an ecosystem engineer. In other words, L. conchilega patches are responsible for an increased habitat quality in an otherwise uniform habitat, which results in a higher survival of the surrounding benthic species.

This study reports diatom cell abundance, concentrations of water-soluble and water-insoluble carbohydrates, and concentrations of chlorophyll—a equivalents as indicators of microbial exudates and biomass in the uppermost sediment layer of two coastal sites (19 m water depth: fine sand; 25 m water depth: mud) in Mecklenburg Bight, south-western Baltic Sea. Correlation analysis with experimentally determined erosion thresholds measured in five different periods of a year showed that the sediment water content and the content of water-soluble carbohydrates may indicate erosion thresholds for both sediment types. Erosion thresholds were significantly negatively correlated with sediment water content at the mud site. At both sites, colloidal carbohydrates, as indicators for diatom EPS, showed significantly negative correlations with sediment water content and no correlation with diatom biomass. Therefore, following current interpretation, we argue that micro-biostabilsation effects of benthic microalgae play a minor role in the investigation area. Light is likely to be insufficient for phototrophic growth on the seafloor of the investigated sites most of the year, as indicated by in situ measurements and calculated values of photosynthetically active radiation. We consider lateral transport of benthic diatoms from shallower to deeper areas of Mecklenburg Bight and subsequent to be the mechanism influencing diatom abundance. Exhaustive biofilms are absent at these depths and mass erosion is not likely to be affected by benthic diatoms.

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.

Echinoderms play a key role in structuring many marine ecosystems and are notorious for large population density variations in so-called ‘‘outbreak’’ or ‘‘dieoff’’ events. In a review of this phenomenon, we assess the causal factors and ecological and evolutionary consequences. We identified 28 species (6 Asteroidea, 8 Echinoidea, 10 Holothuroidea, 4 Ophiuroidea) that exhibit large (more than two population doublings or halvings) population density changes. Three generalized patterns were identified and named for exemplary species: (1) rapid decreases followed by no or slow recovery (Diadema–Paracentrotus Model), (2), rapid increase and apparent stability at a new population density (Amperima–Amphiura Model), and (3) population density fluctuations Acanthaster–Asterias Model). Echinoderms identified were distributed from the shallow intertidal to the deep sea, and from tropical to temperate regions. In most cases, significant impacts on the respective ecosystems were observed. The most striking similarity among all species identified was possession of the ancestral-type planktotrophic larva. This larval type was significantly overrepresented in species identified within the Asteroidea, Echinoidea, Holothuroidea, and for the combined data set. We suggest three main factors that render a life history with planktotrophic larvae a high-risk–high-gain strategy: (1) a strong nonlinear dependency of larval production on adult densities (Allee effects), (2) a low potential for compensatory feedback mechanisms, and (3) an uncoupling of larval and adult ecology. The alternative (derived) lecithotrophic larva occurs in 68% of recent echinoderm species, suggesting an evolutionary trend toward this larval type. Lecithotrophic development represents a more buffered life history because compensatory feedback between adult densities and larval output is likely to be more efficient. For lecithotrophic developers, direct nutritive coupling from adult to larva to the early benthic juvenile provides a buffer against starvation. Lecithotrophic larvae are independent of the vagaries of planktonic food supply, and their short planktonic duration may promote local recruitment. Anthropogenic influences contributed to the population density variations in most cases, including increased primary productivity through eutrophication or global change, disease, overfishing, and species introductions. We suggest that anthropogenic disturbance, through its influence on the frequency and/or amplitude of echinoderm population density changes, may go beyond present ecosystem impacts and alter future evolutionary trends.

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.

In the Firth of Forth, Amphipholis squamata has continuous brooding, and pulsed breeding and recruitment, with highest numbers of brooded embryos and newly released juveniles occurring in the summer months. The estimated size of maturity was 1·5 mm disc diameter although the majority of reproductive output came from individuals of between 2·00–2·99 mm disc diameter. Brittlestar size and season had significant effects on embryo production. Embryos of different developmental stages were present in the bursae of the same individual, and newly spawned embryos and those of a potentially releasable size were present each month. The majority of juveniles apparently emerge at a disc diameter of between 0·8–1·2 mm. Brittlestars were found with full stomachs throughout the year but a peak of feeding was recorded during summer when growth and seawater temperatures were highest. Stomach analyses indicated that almost all animals (99·4%) ingest fine particulate matter but a relatively wide range of plant and animal food items was also identified. Approximately 3% of this Scottish population of A. squamata was infested by the ectoparasitic copepod Cancerilla tubulata.

The influence of macrofaunal reworking activities on the redistribution of particle associated hydrocarbon compounds (HC) was experimentally investigated. Two distinct hydrocarbon mixtures adsorbed on montmorillonite particles (< 4 μm diameter) were added to the surface and deeper (2.5 cm) sediment layers. For comparison, luminophores (100–160 μm diameter) were added in the two deposit layers. At the start of the experiment, four macrobenthic species (the bivalve Abra nitida, the polychaete Scalibregma inflatum, and the echinoderms Amphiura filiformis and Echinocardium cordatum) were added to the sediment surface. The macrofauna added rapidly transferred HC from the surface sediment down to 5 cm depth by both continuous (biodiffusion) and ∼non-continuous (biotransport) transport. Hydrocarbon compounds initially added to the deeper sediment layer were only subject to biodiffusion-like transport. Apparent biodiffusion coefficients (Db) quantified by using a 1-D model were between 0.5 and 8.4×10−3 cm 2 d−1, and biotransport coefficients (r) ranged from 2.0 to 27.6×10−3 d−1. Thus, the four species studied did not have the same effect on particle redistribution and, consequently, on HC repartition in the sediments. E. cordatum was the most efficient reworker. The present study demonstrated the importance of particle size selectivity by benthic fauna, and verified that macrofaunal reworking activities may redeposit sediment from deeper sediment layers on the sediment surface. Both processes have obvious implications for rates and pathways during organic matter mineralisation in marine sediments.

The potential threat to the North Sea ecosystem due to pollution by organic and chemical wastes, has directed attention to the benthic assemblages in the sea. Because these assemblages comprise a wide range of ecological groups and species, many of which have a life span surpassing one year, they integrate effects of pollution over long periods of time. Knowledge on the spatial distribution of different types of assemblages allows assessment of the extension of possible effects over large areas. Analysis of the macrobenthos collected in the Dutch, German and Danish sectors of the North Sea during the synoptical ICES NORTH SEA BENTHOS SURVEY, showed that the samples from this area can be grouped into four clusters on the basis of macrofaunal similarity. Because the geographical distribution of the clusters correlated well with sediment characteristics and depth, for convenience the clusters can be described as shallow coarse- (medium-)sand, shallow fine-sand, shallow muddy-sand and deep muddy-sand. These four clusters differ not only with respect to species composition, diversity and biomass, but also with respect to their consumption and production of organic material. The differences between the structure and functioning of the clusters are interpreted in the light of local differences in the production and deposition of organic material and physical stress on the bottom. Because of the paucity of long-term series of observations, the important question as to whether the present situation shows signs of eutrophication cannot be addressed for the entire investigated area. In the German Bight, where nutrient levels have increased during the past decades and heavy algal blooms have been reported, the macrobenthic communities have not shown a major change. Even after mass mortality among benthic animals due to oxygen deficiency following the breakdown of algal blooms, the benthic community rapidly regained its former structure. This is explained by the dominance of physical factors in shaping the community of this shallow sand habitat. In the more protected muddy-sand habitats, on the other hand, similar events may lead to more permanent changes due to the importance of biological interactions. The available data from studies on the effects of single-point organic inputs on macrobenthic species can be used to predict the sort of changes that would take place in the case of large-scale eutrophication and can help to determine whether this is taking place.

Most experimental studies that deal with the effects of biotic interactions in marine soft sediments are designed to examine the direct effects of one species (or group of species) on some subset of the community. However, species within a community influence one another through a complex web comprising both direct and indirect effects. Ignoring this complexity in the development of hypotheses and experimental designs will likely lead to unexpected or conflicting findings. Few studies of marine soft-sediment communities have emphasized indirect effects and even fewer have included experimental designs capable of testing for them. To illustrate the potential importance of indirect effects in marine soft sediments, I provide several examples from the literature and describe the type of experimental design required to demonstrate a simple indirect effect involving three species. Perspective on indirect effects in a particular community changes as more information is collected about individual species and their effects. The influence of one species or guild may be confounded by that of another and even effects expected to be strong may appear weak or nonexistent as a result of unidentified indirect effects. Recognizing indirect effects in soft-sediment communities may help to explain apparent paradoxical results in past experiments and provide an improved template for the design of future experiments.

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 structure 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.

Predicting the consequences of species loss is critically important, given present threats to biological diversity such as habitat destruction, overharvesting and climate change1. Several empirical studies have reported decreased ecosystem performance (for example, primary productivity) coincident with decreased biodiversity2, 3, 4, although the relative influence of biotic effects and confounding abiotic factors has been vigorously debated5, 6, 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 properties10, 11, making the consequences of species loss more difficult to predict12. 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 documented13, 14, and our research linking these bioturbators to important benthic–pelagic fluxes highlights potential ramifications for productivity in coastal oceans.

On the west coast of Ireland extremely dense aggregations of some echinoderm species were observed. The values determined for the population densities of some of these echinoderms are among the highest that have been known. The investigations focus on the lifestyle of HolothuriePseudocucumis mixta Östergren with special reference to habitat preference, the type of Einbohrens into the ground, the food way of defecation and the meaning life. In situ studies showed that an obviously controlled by the day-night cycle Freßrhythmus present.

The mixing and displacement of sediment by benthic macrofauna (bioturbation) has major biogeochemical implications, and can control rates of organic matter degradation and carbon burial. Large, abundant, mobile macrofauna often dominate sediment bioturbation, and heart urchins of the genus Echinocardium are regarded as key sediment bioturbators in marine systems throughout the world. To better understand the bioturbation potential and functional role of Echinocardium, we developed a mathematical model and parameterized it with field data from six locations in northern New Zealand in order to estimate bioturbation rates in these places. Although urchin sizes and densities were measured in consecutive years at all six locations, we obtained a third model parameter, urchin movement rate, from one time and place only (Site OB5). Because confidence in model output was greatest at OB5, and since OB5 had the highest sediment reworking rate of all sites, our model yielded a good upper bound estimate for the bioturbation potential of Echinocardium in the areas examined. The volume of sediment displaced by Echinocardium populations reached 20,000 cm3 m−2 d−1 at OB5, suggesting that surface sediment is reworked about every 3 days at sites where Echinocardium is abundant. Experimental work with a fluorescent tracer at OB5 suggested limited downward particle movement as a result of Echinocardium bioturbation, though vertical profiles of chlorophyll a and organic matter content indicated well mixed sediment. The loss of Echinocardium because of broad-scale anthropogenic disturbance to the seabed could have major consequences on marine ecosystem functioning.

Near the island of Sylt in the Wadden Sea (German Bight, North Sea), seastars Asterias rubens (L.) co-occur with their preferred prey, mussels Mytilus edulis (L.), which form extensive beds from the intertidal down to the subtidal zone. Mussel density within these beds is significantly lower in the subtidal than the intertidal zone. Laboratory and field experiments were conducted to check if this was due to seastar predation. Feeding experiments did not indicate size refuge of M. edulis from predation by A. rubens, but showed that seastars preferred clean subtidal mussels above barnacle-overgrown intertidal ones. This preference coincided with higher abundances of both large View the MathML source and smaller seastars in the shallow subtidal, but their abundance was too low to account for the decreased subtidal mussel density in the area studied. However, seastars may indirectly reduce mussel recruitment in the subtidal zone. This is caused by juvenile seastar predation upon the barnacles that grow on mussels, because such epigrowth strongly enhances recruitment in mussels. Such an indirect effect on mussel recruitment may affect mussel density more than adult seastar predation. An exception may be mass invasions of A. rubens on subtidal mussel beds. One such event happened during this study, clearing a large patch of mussels.

We investigated the choice preferences exhibited by the spider crab Inachus phalangium when using algae for feeding and masking, together with the pattern of decoration present on the body surface and the morphology of the setae, which are used to attach masking material. We showed that feeding and masking preferences are decoupled, and that I. phalangium exploited the algal nutritional and chemical properties differently. When feeding, the crab revealed a differentialchoice among the offered algae, preferring the protein-rich cyanobacteria. When decorating, both algal components that match the background and the Phaeophyta Dictyota dichotoma, which acts as a chemical deterrent against predators, were actively chosen. At the same time, decoration in I. phalangium was used as a short-term food store. We also provided evidence that masking patterns varied in different body regions, depending on the functional morphology of the hooked setae. This work discusses the origin and evolution of the decorative habit, showing for the first time that decoration is a multifaceted mechanism, tightly linked with morpho-functional and behavioural adaptations, and that masking material can also be used as a food store.

Laboratory and in situ studies have shown that the prosobranch Balcis alba is a temporary ectoparasite on the boreal holothuroid Neopentadactyla mixta. This association is described in relation to aspects of the species' functional anatomy and ecology.

(1) The importance of large epibenthic predators in structuring marine sublittoral assemblages is unclear. Several previous investigations have yielded conflicting results, possibly because of limitations in experimental design. The effects of predation by the crab Liocarcinus depurator on prey assemblages are examined in a field experiment in a subtidal habitat on the west coast of Scotland. (2) Cages covering 1 m2 maintained crab densities from zero to eight crabs m-2 (0-32 times natural densities) for a period of 24 days. The total number of prey individuals, the number of species in samples and abundance of selected species were compared between treatments using ANOVA. Overall differences in assemblages were explored using multidimensional scaling. Effects on population size structure for four common species were also examined. (3) Contrary to expectation, there was little evidence that predation by Liocarcinus had any significant effect on the abundance of benthic species. The number of species in samples and the total number of individuals in samples were not significantly different between treatments and there were no effects on individual species which could be interpreted in terms of predation. The multivariate analysis, however, suggested some effects of caging, possibly due to the exclusion of other predators in addition to crabs in the case of infauna. (4) Potentially confounding processes such as prey mobility, crabs not feeding normally and insufficient power in the experiment design appear not to be significant features of our results. Therefore, we conclude that Liocarcinus has trivial effects on the benthic assemblages in the study area.

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.

The increasing need for renewable and clean energy production is likely to result in a diversi fication of locations for the implementation of offshore wind farms which have been so far predominantly sited on soft substrata. In contrast, offshore wind turbines placed on rocky reefs in highly exposed areas are much less common and the impacts on localflora and fauna can only be 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 variablesmay 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 urchinpopulations. 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 information about the potential impacts of the construction of an offshore wind farm on rocky substratum and allows us to generate impact hypotheses that go beyond what could behypothesized through a mere general knowledge of the area.

Measurements of sub-surface light attenuation (Kd), Secchi depth and suspended particulate material (SPM) were made at 382 locations in transitional, coastal and offshore waters around the United Kingdom (hereafter UK) between August 2004 and December 2005. Data were analysed statistically in relation to a marine water typology characterised by differences in tidal range, 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 rateof 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 in both estuarine and coastal waters. In terms of susceptibility to eutrophication, large macrotidal, well mixed estuarine waters, such as the Thames embayment and the Humber estuary were identified at least risk from eutrophic conditions due to light-limiting conditions of the water type.

This paper presents new results from high temporal resolution observations over two years (2007 and 2008) from instrumented moorings deployed in the central North Sea, at the Oyster Grounds and on the northern slope ofDogger Bank (North Dogger). The water column was stratified in the summer at both sites, leading to limited exchange of the water in the bottom mixed layer. Data from these moorings revealed the variable nature of summer oxygen depletion at the Oyster Grounds. The combination of in situ and ship-based measurements allowed the physical and biological conditions leading to decreasing dissolved oxygen concentrations in bottom water to be examined. In 2007 and 2008, the concentration of dissolved oxygen in the bottom water at both sites was observed to decrease throughout the summer period after the onset of stratification. Depleted dissolved oxygen concentration (6.5 mg l−1, 71% saturation) was measured at the North Dogger, a site which is not significantly influenced by anthropogenic nutrient inputs. Lower oxygen saturation (5.2 mg l −1. 60% saturation) was measured for short durations at the Oyster Grounds. The seasonal increase in bottom water temperature accounted for 55% of the decrease in dissolved oxygen concentration at the Oyster Grounds compared to 10% at North Dogger. Dissolved oxygen concentration in bottom water at the Oyster Grounds was shown to be strongly influenced by short term events including storms and pulses of particulate organic matter input. In contrast, dissolved oxygen concentration in bottom water at the North Dogger reflected longer seasonal processes such as a gradual temperature increase over the summer and a more steady supply of particulate organic matter to the bottom mixed layer. The differences between the study sites shows the need for an improved understanding of the mechanisms driving these processes if the use of oxygen in marine management and ensuring ecosystem health is to be meaningful and successful in the future. These high frequency observations provide greater understanding of the nature of the depletion in bottom oxygen concentration in the North Sea.

Suspension-feeding eastern oysters, Crassostrea virginica,were once abundant in Chesapeake Bay and may then have exerted top-down control on phytoplankton and also reduced turbidities, thereby increasing light available to benthic plants. Alternatively, oysters may have simply recycled inorganic nutrients rapidly back to the water column, with no long-lasting reduction in phytoplankton biomass resulting from oyster feeding activity. To help distinguish between these scenarios, we explored changes in nitrogen fluxes and denitrification in laboratory incubations of sediment cores held under oxic and anoxic conditions in response to loading by pelletized phytoplankton cells, an experimental analog for oyster feces and pseudofeces. When organics were regenerated under aerobic conditions, typical of those associated with oyster habitat, coupled nitrification–denitrification was promoted, resulting in denitrification of ;20% of the total added nitrogen. In contrast, under anoxic conditions, typical of current summertime conditions in main-stem Chesapeake Bay where phytoplankton is microbially degraded beneath the pycnocline, nitrogen was released solely as ammonium from the added organics. We postulate that denitrification of particulate nitrogen remaining in oyster feces and pseudofeces may enhance nitrogen removal from estuaries. In aerobic incubations with sufficient light (70mmol m22s21), a benthic microalgal/cyanobacterial community grew that not only absorbed the inorganic nitrogen released from the added organics but also fixed N2. This result suggests hat an ecosystem dominated by benthic primary production may develop in shallow waters when reduced turbidity associated with bivalve feeding increases light penetration to a level that can sustain benthic microalgal production.

The secondary production of mobile invertebrate fauna in the Laminaria hyperborea (Gunn.) Foslie kelp forest increases with wave exposure level. This faunal group has a key function in transferring kelpcarbon to higher levels in the food web. By using a size-frequency method the calculated production was68 ( 18) g D.W. m 2 yr 1 ( S.E.) at low, 250 ( 57) at medium and 308 ( 64) at high exposure levels. The calculations included 30 macrofauna species, which accounted for 96% of the specimens registered, withGastropods, amphipods and bivalves being the most abundant taxa. The calculated secondary productionis high, but comparable to that previously reported from other macrophyte systems and was 3%, 8% and 8% of the total primary production at low, medium and high exposure levels, respectively. Our results indicate that large quantities of Laminaria kelp are exported from the system, although the production of sessile animals was not taken into account. The most important factor in determining faunal densities and secondary production was probably habitat size but at low exposure levels the percentage of eggcarrying crustacean females and juveniles were lower than at medium and high exposure levels, thereby indicating lower fitness for animals at low exposure stations.

Kelp (Laminaria hyperborea (Gunn.) Foslie) forests support a diverse community of invertebrates. The aim of the present study was to test if the abundance, diversity (measured as average species number) and community structure of macrofauna associated with epiphytic algae on kelp stipes depend on physical stress from wave surge and habitat diversity, or if differences simply result from differences in habitat size. Results show that faunal diversity increased with wave exposure, mainly due to increasing wave stress and habitat diversity. Abundance was highest at medium wave exposed stations, and lowest at low exposure stations. However, the habitat structural diversity is high in kelp forests regardless of wave exposure, and a typical kelp forest fauna is associated with functional groups of epiphytic algae across different levels of wave exposure.

As part of a multidisciplinary investigation of pollution in northeast Britain a study of the sublittoral kelp fauna is described. Sampling strategy is discussed and a programme adopted involving the investigation of an instantaneous fauna1 pattern over a wide area and its environmental correlation. The kelp holdfast as a sample unit is briefly evaluated. The total fauna] content of the survey samples is described in gross terms. A review of the physical parameters which impinge upon the holdfast fauna is made and on this basis significant variables are resolved into turbidity, pollution, water movement, and holdfast morphology. Detailed treatment of these factors follows and their distribution over the northeast coast transect is described. Seasonal studies on suspended solids enable an approximate area north of the River Coquet in Northumberland to be considered as continually clear, whilst the area to the south is considered turbid. The region of turbid water is considered to be the consequence of erosion of superficial coastal deposits under the action of marine and atmospheric forces. The role of discharges in contributing to suspended solids is considered. Nutrient data are given; the amounts were spatially similar over the whole coastline. An area of ‘pollution’ is provisionally designated on the basis of published figures. The problems of assessment of wave exposure and present inadequate knowledge of local inshore current systems preclude other than crude separation of one site on considerations of water movement. Holdfast parameters are generally interdependent: however the degree of holdfast branching is shown to be independent of age and to influence significantly (along with water movement external to the holdfast) the degree of holdfast silting.

The kelp Laminaria hyperborea is a dominant component of the subtidal nearshore ecosystem and is subjected to a heterogeneous wave and current climate.Water motion is known to influence physiological processes in macroalgae such as photosynthesis and nutrient uptake attributed to mass-transfer limitation. The study attempts to establish the effect of water motion on the growth rates of blades and elongation rates of the stipes of L. hyperborea at adjacent wave-exposed and wave-sheltered locations over a 12 month period from field observations. The observations were supported by detailed physical and chemical measurements (light, temperature, seawater nutrient concentrations and hydrodynamics) and of tissue carbon and nitrogen concentrations together with δ13 carbon. Despite a 30% difference in the root mean square of the velocity (Velrms) between the two survey locations, there was no evidence to suggest that water motion had any direct influence on the growth rates of either the blades or elongation of stipes of L. hyperborea. No significant differences were observed between either environmental or plant physiological variables between the sheltered and exposed locations. Using an integral velocity parameter (Velrms) the present study also highlighted the importance of the tidally induced current component ofwater flowin the subtidal zone.

Because of their likely ecological importance, the effects of understory kelps on fluid and particulate transport near the bottom were assessed in waters of the San Juan Archipelago, Washington, U.S.A. Relative to more exposed rocky substrata at identical depths, bottoms beneath kelp canopies were exposed to weak fluid transport and were characterized by greater rates of deposition of particulates. A tracer particle experiment demonstrated that kelps inhibited transport of suspended particles from the overlying water column to the bottom. Thus, the higher rates ofparticulate deposition beneath kelp canopies probably were caused by longer particle residence times and higher probabilities of particulate redeposition beneath canopies and not by higher rates ofparticle import. These hydrodynamic effects may play important roles in the ecology of animals that inhabit understory kelp environments. We propose several potential effects of flow modification by kelps on larval recruitment and dispersal and growth of suspension-feeders.

Previous studies demonstrated that understory kelps altered rates of mass and particulate transport beneath kelp canopies. This study examined the implications ofthese physical effects, plus shading by canopies, on the recruitment of four taxa of benthic invertebrates. Using field manipulations of understory kelp density and substratum orientation, we examined the effect of kelps on invertebrate recruitment and assessed the mechanisms involved. These mechanisms included reduced flow, increased sedimentation, reduced light intensity and reduced microalgal cover. All effects were significant to recruitment, but their relative importance varied among taxa. Two species examined, the serpulid polychaete Pseudochitinopoma occidentalis (Bush) and the cheilostome bryozoan Membranipora membranacea (Linnaeus) have long-lived planktonic larvae. Kelp canopies signiticantly affected recruitment of these two species, but in different ways. Pseudochitinopoma recruitment was reduced by sedimentation and light but was unaffected by ambient flow directly. Recruitment of Membranipora exhibited patterns consistent with that of passive transport of larvae with higher recruitment in high flow, no-kelp environments. Its recruitment was reduced by higher sedimentation beneath canopies, but showed no response to changes in light intensity. Spirorbid polychaetes have short planktonic phases of unknown durations. Kelps significantly affected their recruitment but difference in recruitment rates between kelp-covered and exposed substrata varied in direction with season. Spirorbid recruitment was reduced by light and high levels of sedimentation, but their recruitment patterns were unaffected by ambient flow directly. The cyclostome bryozoan Tubulipora sp. has a very short planktonic period (seconds to minutes) and factors afFecting its recruitment were consequently ditlicult to decipher. It apparently responded negatively to higher light intensity and increased sedimentation. The effects of kelp canopies on recruitment depend on the complex interaction of these variables.

Patterns of distribution and abundance of epifauna often differ markedly among macroalgal species. The hypotheses tested were that (1) assemblages of mobile epifauna associated with Laminaria ochroleuca and Sargassum muticum differed because they have different structure, and (2) assemblages of mobile epifauna associated with S. muticum differed between heights on the shore because tidal height affects physical and biological conditions.We also investigated the effect of epiphytic biomass on the composition of epifaunal assemblages. Hypotheses were tested with measuring and manipulative experiments using natural and artificial algae, and by measuring uni- and multivariate assemblage descriptors. The results indicated that epifaunal assemblages associated with natural L. ochroleuca and S. muticum differed, but only differences in epifaunal densities were likely to be related to the structure of algae since all other variables did not clearly differ between the two algae. Although structure might play an important role, other factors need to be taken into account and further experimental tests are necessary. Epifaunal assemblages associated with S. muticum did vary depending on the height on the shore, but inconsistently over time in the case of natural algae. In addition, epifaunal densities of natural algae were positively related to biomass of epiphytes in both species. Time of sampling, epiphytic load and height on the shore were the most important factors in structuring epifaunal assemblages rather than complexity of the host algae.

Despite the wide range of functional roles performed by marine sponges they are still poorly represented in many research, monitoring and conservation programmes. The aim of this review is to examine recent developments in our understanding of sponge functional roles in tropical, temperate and polar ecosystems. Functions have been categorised into three areas: (a) impacts on substrate (including bioerosion, reef creation, and substrate stabilisation, consolidation and regeneration); (b) bentho-pelagic coupling (including carbon cycling, silicon cycling, oxygen depletion and nitrogen cycling); and (c) associations with other organisms (facilitating primary production, secondary production, provision of microhabitat, enhanced predation protection, survival success, range expansions and camouflage though association with sponges, sponges as a settlement substrate, disrupting near-boundary and reef level flow regimes, sponges as agents of biological disturbance, sponges as releasers of chemicals and sponges as tools for other organisms). The importance of sponges on substrate, sponge bentho-pelagic coupling, and sponge interactions and associations is described. Although the scientific evidence strongly supports the significance and widespread nature of these functional roles sponges still remain underappreciated in marine systems.

Understanding temporal variation at the scale of weeks to months is critical to understanding broad temporal patterns in diversity in the same way as understanding diversity across landscapes relies on understanding variation at the scale of meters. However, whereas small-scale spatial variation in temperate reef algal assemblages has been extensively studied, fine-scale temporal changes have not been well addressed. By sampling the macroalgae of a subtidal reef near Perth (Australia), dominated by the small kelp Ecklonia radiata, every w40 days over a 2-year period, we were able to test whether temporal changes in species richness, assemblage structure and species turn-over were related to seasonal changes in surface temperature, solar radiation and wave height. A total of 93 macroalgal taxa were identified, and species richness per sampling time ranged from 25 to 64 taxa 1.25 m 2. Biomass of E. radiata was positively correlated with changes in sea surface temperature and light, and negatively correlated with wave height. Species richness, assemblage structure and turn-over of other macroalgae were more associated with seasonal changes in kelp biomass than environmental variables per se. We conclude that seasonal changes in environmental conditions drive changes in the kelp canopy, which in turn drive changes in species richness and assemblage structure. This suggests that habitat-formers such as kelps can exert a strong temporal influence on associated communities, analogous to well-described spatial influences. Thus, as kelp canopy biomass expands and retracts over time-scales of weeks to months, so does available space for colonization and growth, resulting in a high species turn-over. Species richness is therefore increased and maintained through time, in the same way as canopy-gap mosaics increase and maintain species richness across spatial landscapes.

Canopy-forming algae occur across of range of energy environments (i.e., wave sheltered to exposed coasts) where disturbances are frequent (i.e., gap formation) and benthic patterns largely reflect variation in post-disturbance processes. Disturbances vary in extent (area affected) and intensity (degree of damage), and this may affect recolonisation at local scales. On an open oceanic coast, we tested whether habitat structure (patches of canopy algae) differed between heavy and relatively lighter wave exposure (sheltered vs. exposed sides of islands), and whether wave exposure affected the response of prominent habitat-formers to varying disturbance regimes (different sizes of partial and complete canopy removal). Observations of naturally occurring patterns showed sheltered coasts to be characterised by small patches of fucoids, whereas exposed coasts were characterised by large patches of kelp. Canopy-gaps were larger at exposed than sheltered coasts, and mixed canopies constituted N24% of the subtidal rocky habitat independently of wave exposure. Experimental disturbances showed the local density of kelps to affect recovery through greater recruitment to partial clearings (80% canopy removal). Fucalean algae, on the other hand, mainly recruited into complete clearings (100% removal), but when their recruits were abundant, they also recruited into partial clearings. The covers of filamentous, turfforming algae increased in all clearings, and more so at exposed than sheltered sites. Extent of disturbance had no detectable effect on recolonisation by canopy-forming algae across the scales examined (i.e., 1.5 m, 3 m diameter loss of canopy). Recolonisation varied among islands kilometres apart, and correlations (rN0.85) between cover of canopies and cover of their recruits in clearings at the scale of sites, suggested that differences in propagule supply could account for variation in patterns of recolonisation at scales of kilometres. There was no evidence to suggest that the effect of disturbance depended on wave exposure within the range of exposures tested in this study (i.e. open coasts). We recognise that wave exposure can be fundamental to habitat structure of subtidal rocky coasts, but we suggest that its influence may be mediated by the biological setting (e.g., canopy composition).

The effect of sea urchins on algae has often been studied when sea urchins forage over areas to create or maintain urchin barrens. The effect of sea urchins that are not associated with barrens, however, is not so clear. This study tested the hypothesis that sedentary and largely site attached sea urchins (Heliocidaris erythrogramma) have detectable effects on the covers of canopy-forming and understory algae, whether that be due to their mere presence or grazing. Algal cover was estimated beneath and immediately surrounding urchins in a location where they do not create barrens but attain densities at the upper end of their local range. The area immediately beneath the urchins was mostly devoid of visible algae. Experimental removal of urchins over 3 years revealed that canopy-forming algae were able to develop in more extensive covers to match those in areas naturally lacking urchins. In the presence of urchins, the immediate area (i.e. first 5–6 cm from urchins) was primarily comprised of turf-forming algae, thereafter the cover of canopyforming algae sharply increased until at ~8 cm their covermatched areas without urchins. These data support the model that composition and cover of algae can be altered in the presence of non-barren forming urchins, albeit only within a few centimetres of individual urchins. Where populations of sedentary urchins are dense and widespread (e.g. sheltered coasts for H. erythrogramma) such fine-scale effects may repeat over large areas such that urchins do not need to be associated with barrens before their effects are detectable.

The effects of sedimentation on the development and structure of macroalgal assemblages were investigated from June 1992 to December 1993 on a rocky subtidal shore South of Livorno (Ligurian Sea). Experimental procedures were specifically designed to address: (1) which species recruit in patches of bare rock under different sedimentation rates; (2) whether small-scale variations of sediment deposition influence the local diversity of the assemblage; (3) whether the effects of sedimentation are independent from the time of the year when succession begins. Depositional environment was manipulated in the field by means of transparent Plexiglas panels which on average reduced the amount of sediment sinking over the algal assemblage by 35%. Possible effects on the penetration of light and flow microenvironment were assessed. Small-scale variations of sedimentation rates significantly influenced the local structure and diversity of the algal assemblage. Such effects were particularly evident during the recolonisation of patches of bare rock and were dependent upon the time at which succession was initiated. The erect algae were the ones more affected by depositional environment and showed different responses to the experimental reduction of sediment supplies. All the plots were quantitatively dominated by a filamentous turf mostly consisting of Polysiphonia setacea, whose growth was potentially enhanced by reduced sedimentation rates. At the end of the experiments, the overall diversity and evenness of the assemblage were lower in treatments with lessened sediment supplies. It is suggested that small-scale spatial variability of the depositional environment may affect the within-habitat diversity of algal assemblages either through direct effects on individual species or on their propagules, and through indirect effects mediated by competitive outcomes.

The relationship between macroalgal assemblages and abiotic factors was quantified by gradient analyses in an area where longterm changes in macroalgal depth distributions have previously been documented. Biomass data from 4, 6, 8 and 10 m depth in an area of similar salinity (5) and substrate (rock) in the northern Baltic Proper was constrained by a set of environmental variables defining different aspects of abiotic control of species distributions (sediment cover, effective fetch, clarity index, the curvature and slope of the bottom, and direction of exposure) in multivariate analyses at different scales. Fucus vesiculosus dominated the biomass at 4, 6 and 8 m depth, and Furcellaria lumbricalis at 10 m. The applied models explained 30.7–53.3% of the total variance in community structure, and 49.3–60.9% when analysed separately for each depth. A separate analysis of species depth distributions demonstrated that effective fetch was most strongly related to upper limits of the algal belts, sediment cover to the lower limit and density of the F. vesiculosus belt, and clarity index to the lower limits of F. vesiculosus, perennial red algae, and of the red algal and Sphacelaria spp. belts. The results show a strong correlation between environmental variables and vegetation structure even on a small, local scale in the northern Baltic Proper, indicating a high suitability of the phytobenthic zone for environmental monitoring. The results add to previous studies that show a strong importance of abiotic factors on large-scale variation in phytobentic community composition in the Baltic Sea.

Recruitment ofepifaunal invertebrates was followed on 16 x 16 cm Perspex panels immersed at a low imertidal site at St Andrews, on the east coast of Scotland. Short-term experimental enclosures assessed the importance of grazing by the herbivorous prosobranch Gibbula cineraria (L.), and the nontrophic activities of the predatory species Nucella iapiii~ts (L.) and Asterias rubens L., on developing epifaunal assemblages. At the end of each experimental period, recruits of all seven taxonomic groups examined were less abundant on G. cineraria grazed treatments, than on those from which grazers were excluded. Furthermore, none of the epifauna were apparently able to achieve an escape-in-size during a 2-month period of exclusion of grazers. N. lapillus and A. rubens did not appear to have similar deleterious effects on the developing assemblages (with the possible exceptions of barnacles, hydroids and ascidians during periods of peak larval settlement). Similar numbers of recruits were recorded on the treatment panels and the net controls. Enclosure of N. lapillus appeared to enhance the recruitment of some species (e.g., anomiids). These experiments provide no information regarding the nature of the effects ofthese consumers on the developing assemblages. However, there is circumstantial evidence that the effects of G. cineraria may be attributable to the rasping-off of recently established recruits, as the snails browse across the substrata.

It has been difficult to understand the effects of habitat structure on assemblages because the different elements of habitat structure are often confounded. For example, few studies consider that the effects of structural components of a habitat (rocks, trees, pits, pneumatophores) may be separate from the complexity (e.g. surface area hSAj) they create. From prior observations and experiments, I developed three hypotheses about the effects of habitat structure on gastropods on rocky intertidal shores in Botany Bay, Australia. (1) The complexity of habitats positively affects the density and richness of gastropods. (2) The fractal dimension (D) represents elements of complexity that affect the density and richness of gastropods better than other indices of complexity. (3) The effects of specific structural components on the density and richness of gastropods are independent of their complexity. To test these hypotheses, treatments composed of pits and pneumatophores were used to independently manipulate complexity and structural components in experiments repeated at five different times on two shores. There was support for hypotheses (1) and (3) at most times and places but not for hypothesis (2). Richness, total density, and the densities of two of the three most common gastropods were greater in treatments with greater complexity. D was not definitively better than other indices of complexity, but D and SA were recommended for further consideration. When complexity was held constant, species richness and the density of most gastropods, except Austrocochlea porcata, was greater in treatments with pits than with pneumatophores. A common mechanistic explanation for the effects of habitat complexity on rocky intertidal gastropods relies on a specific characteristic of pits; they pool water and reduce desiccation stress. This assumption may be appropriate for many gastropods, but it was inappropriate for A. porcata. Habitat complexity affected its density, but this was not because of a characteristic specific to pits. The complexity and structural components of habitats have separate effects on assemblages, and it confuses the study of habitat structure to combine them.

In the Wadden Sea common starfish is an important predator of mussel beds which in turn are relevant ecologicaland economic resource. To improve the management of mussel seedbeds, knowledge is required on over winter predation, a factor affecting mussel survival. The aim of this study was to assess the importance of A. rubens feeding activity during winter and how it relates with changes in temperature. Feeding activity of starfish was monitored during a full winter. The potential impact of temperature change on starfish–mussel seed interactions during winter was analysed. The factor shading was included, as changes in light intensity appear to be a primary governing factor for the timing of feeding activity. The results showed that temperature limits feeding rate and feeding activity of starfish during winter. However, starfish feeding rate exhibited very high sensitivity to temperature changes. Light intensity affected both feeding rate and feeding activity. It is concluded that starfish may not be an important factor destabilising seedbeds during a mean winter, but its importance may grow along with the increasing temperature due to climate change.

Mytilus sp. is a habitat-modifying species that can influence biodiversity by facilitation and inhibition of species. We investigated the relationship between size of Mytilus patches, sediment characteristics and species richness of associated macrofauna, meiofauna and macroalgae on sandy sediments at 7 m depth in the Askö area, northern Baltic proper. A total of 24 associated macrofauna species were identified, 11 of which were not present on bare sediment outside the patches. The bare sediment had on average 5 species, which increased to 6 associated species at a mussel patch size of 5 cm2 and 17 species at 314 cm2, which was >85% of the associated species in a nearby large mussel bed. The diversity of macro-infauna was higher in mussel patches compared to sediment. Macroalgae, being dependent on mussel shell as substrate or on attachment by byssus threads, were not found outside patches. Sediment contents of total carbon and nitrogen were significantly higher within patches than outside, mainly due to the mussels’ filtration and biodeposition of small nutrientrich particles. Diversity of meiofauna did not change with patch size, but total abundance of Nematoda was higher within patches. As patch size increased, the effects of Mytilus on the sediment characteristics became stronger, but few negative effects of increased biodeposition were seen on species richness. The results show that mussel patches represent islands of high biodiversity in sediment areas and belong to the most species-rich habitats in the Baltic Sea, due to structural and functional effects.

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae.

Ophiothrixfragilis is a suspension-feeder which forms dense aggregations (up to 2000 individuals m 2) in the Dover Strait. Its optimal suspension-feeding activity is expressed when currents are less than 20 cm s - 1. Thus, the duration of this behaviour over a tidal cycle depends on the tidal velocity, and can be very short during spring tides. During neap tides, suspension-feeding is almost continuous but the flux of nutrients (phytoplankton, organic carbon and nitrogen) remains low because current increases slowly after slack water, and resuspension phenomena do not occur. In contrast, during spring tides, the current speed rapidly increases and induces an important resuspension of particles. Thus, the flux of nutrients is very important during the short duration of suspension-feeding of the O. fragilis population. Minimum gut contents were observed 3 h before slack current and maximum gut contents were observed about 1 h after slack water.

The impact of inorganic sediments on rocky shore meiofauna was investigated in relation to tidal elevation, using artificial “algal” mats of differing complexity. Sediment accumulation was correlated with habitat structure and increased at higher elevations. The water content and retention time of mats was similarly related to sediment content and mat structure. The meiofauna was numerically dominated by small interstitial forms, notably harpacticoid copepods, copepod nauplii and nematodes. Densities recorded in the high shore were as great as those in the low intertidal, although diversity (especially of larger forms) and biomass were markedly lower. These patterns are discussed in relation to habitat structure and sediment content. By retaining water, sediments can create moderately stable environments within algal mats on the high shore, to the advantage of small meiofauna. As a result of the increasingly favourable environment towards the bottom of the shore, meiofaunal diversity and biomass increased. Traditionally, the positive relationship between structural complexity and meiofaunal abundance and diversity has been viewed in terms of the number of habitats and the provision of refugia from predators. It is concluded here that complexity and sediments also influence microenvironmental conditions and that this in turn has an important impact on meiofaunal communities.

The stalkless crinoid Antedon mediterranea (Lamarck) is a passive suspension-feeder, relying on exogenous currents to supply it with food. The animal uses aerosol filtration mechanisms to remove particles from suspension and thus can feed on particles which are smaller than the mesh of its filter. The influence of food availability on feeding activity of Antedon were monitored quantitatively in a laboratory flow chamber. Because the rate at which the animal encounters food particles, is a combined function of particle concentration and of ambient current speed, both these variables were manipulated simultaneously and their combined influence on feeding activity were monitored. Ambient current speeds were varied between 0.4 and 3.7 cm. s ’ and concentrations of food (the unicellular alga Hymenomonas) ranged from 0 controls to 10000 cells. ml _ ‘. Both particle concentration and ambient current speed shape the functional response of Antedon at all values tested; however, they do not show an interactive effect. I.e., the feeding activity of the crinoid is not a function ofthe rate at which particles approach the filter but depends only on the ambient particle concentration and current speed. This observation can be accounted for by hydromechanical effects at the filter and a complex behavioral modulation of the feeding response. In addition to the above effects, a novel form of hysteresis in the feeding behavior was discovered: response to a given current velocity depends on whether that velocity was reached by an increase or a decrease in speed. This response may result from down regulation of receptor mediated events.

Biodeposit production of Mjrtilus eduLis was studied in situ over an annual cycle in the northern Baltic proper. Faeces and pseudofaeces were collected in traps covered by a layer of mussels and placed on the bottom at 5 m depth. The summed deposition due to sedimentation and biodeposition was compared with deposition in mussel-free control traps on the bottom and suspended in the water column. Both quantity and quality of deposited matter increased through biodeposition. Relative contribution of matenal from M. edulis was hlghest in summer, whereas both sedimentation and biodeposition peaked in autumn. Annual biodeposition by M. edulis was 1.76 g dry weight, 0.33 g ashfree dry weight, 0.13 g carbon, l.? X I O - ~ g nitrogen and 2.6 X 104 g phosphorus per g mussel (dry weight including shells). During the productive season from April to September the organic content of biodeposits was higher than that of naturally sedimenting material (30.4 vs 24.8%), despite having passed the mussels' alimentary tracts. Nitrogen content of biodeposited material was also higher than that in control traps (1.68 vs 1.20 % of dry weight), while phosphorus content was the same (0.19 %) in both. The higher contents of carbon and nitrogen in biodeposits may be explained by the fact that the mussels to a large extent remove and initiate the sedimentation of small particles of high organic content whch would otherwise stay in suspension. We argue that M. edulis occupies an important role as a connecting link between pelagic and benthic ecosystems by increasing total annual deposition of C, N and P by 10 % and by circulating and regenerating 12 and 22 %, respectively, of the annual N and P demands for pelaqc pnmary production in the area.

Habitat-modifying species such as Mytilus edulis strongly impact both community structure and ecosystem functioning through positive or negative interactions with other species and by changing physical and biological conditions. A study of natural patches of mussels showed that C and N content of sediment was higher in mussel patches compared to the surrounding sand community. Species richness and biomass of associated macrofauna and -flora was enhanced even by the presence of single mussels and increased rapidly with patch size up to about 150 cm2, while no change in sediment meiofauna was found. In order to separate the effects of structure and function of M. edulis, a manipulative field experiment was performed with constructed patches of cleaned live mussels or intact empty shells. After 3 mo colonisation, the number of associated species in both treatments approached those in natural mussel patches, indicating that species richness was mainly due to physical structure. Abundance and biomass of associated flora and fauna were higher if live mussels were present because mussel biodeposition and nutrient regeneration supplied limiting resources and increased carrying capacity. Species composition was also affected. Complexity and biodiversity increased with time, especially if Fucus vesiculosus plants established themselves. Measurements of community metabolism showed that the associated community found in mussel patches depends on mussel biodeposition for 24 to 31% of its energy demand. Thus, the energy balance of the community is dependent on the function of live mussels.

Sea anemones in genera Adamsia, Calliactis and Paracalliactis (family Hormathiidae) engage in a mutualistic symbiosis with hermit crabs in which the anemone gains substrate and food in exchange for defending the crab. Some of the sea anemones also expand the living space of the crab by producing a carcinoecium, a chitinous structure that overlies the initial gastropod shell in which the hermit crab lives. The symbiosis is initiated either by the crab, or by the anemone. Although behavioral and physiological aspects of this symbiosis have been studied, interpretations cannot be generalized without an evolutionaryframework. After reconstructing relationships among members of Hormathiidae using DNA sequences, we find that the association has evolved at least twice: Adamsia nests within Calliactis in a single clade, and Paracalliactis belongs to a different clade within the family. The carcinoecium and complex behavioral and anatomical features associated with the symbiosis are interpreted as having evolved at least twice within Hormathiidae and seem to be phylogenetically labile.

The sea anemone Metridium senile is a quantitatively important passive suspension feeder in hard-bottom communities on the west coast of Sweden and occurs in aggregations with different size distributions. This study tests the hypothesis that different polyp sizes have different optimal flow regimes maximizing prey capture. Results showed that prey capture by M. senile is a function of both flow regime and polyp size, and different optimal flow regimes exist for different size classes. Large anemones had a maximum feeding efficiency at the slowest flow, medium-sized anemones at moderate flow, and small anemones at moderate- to high-flow regimes. Small anemones showed consistently higher feeding rates (per unit of biomass and area of tentacle crown) at all velocities above 10 cm s-’ and exhibited less flow-induced deformation of the tentacle crown, suggesting that small anemones are better at feeding in moderate- to high-flow habitats. Different vertical projectionsof large and small anemones in the boundary layer could only partly account for differences in feeding success among size classes. Feeding rate was also a function of upstream conspecifics, declining asymptotically to 30% of the maximum rate. The negative effects of neighbors on feeding in aggregations with clonal rather than polyp growth appear to be compensated for by the generally higher feeding efficiency of small polyps.

The role of native consumers in mediating biological invasions is poorly understood. In theory, there are reasons to expect both strong and weak effects of native consumers on non-native species. However, non-native ranges may include multiple regions or even continents, each with its own suite of consumers and invader–consumer interactions may play out differently in different places and times. In this Washington State (USA) study we found that the common herbivorous snail Lacunavincta was 2–9 times more abundant on the nonnative seaweed Sargassum muticum, compared to native kelps. Choice feeding trials with fresh tissue and artificial foods both suggest that S. muticum is a preferred food for Lacuna vincta. Lab experiments indicated that L. vincta did not experience diminished predation by two common predators on Sargassum muticum compared to native kelp hosts. Our results suggest that Sargassum experiences considerable herbivory by Lacuna vincta in our study region, a conclusion that is consistent with previous work and our own field observations. In our system, L. vincta and S. muticum have been coexisting in the same habitats for at least 50 years and available data suggest that it acquired a preference for S. muticum more than 30 years after the initial invasion. Comparison of our results to recent work on Sargassum– herbivore interactions in Europe suggests that the response of native consumer communities to S. muticum varies both within and among regions. Geographic and temporal variation in the response of native consumers are likely to be hallmarks of many large-scale invasions.

Introduced algae form globally an increasing problem. Grazing on invaders could provide communities with resistance to algal invasions. In this study, we experimentally tested the food choice of mesoherbivores for native macroalgae versus the invader Sargassum muticum. We performed food choice experiments with common grazers (Gammarus insensibilis, Hydrobia ulvae and Stenosoma nadejda) and both the brown algal invader Sargassum muticum and its native macroalgal competitors, from both the south (Cystoseira humilis, Stypocaulon scoparium, Cladostephus spongiosus, Dictyota dichotoma, Dictyopteris polypodioides, Sargassum vulgare) and southwest (Cystoseira humilis) coast of Portugal. We tested whether (i) the invader affects the growth rates of native macro algae, and (ii) the grazers have a food preference for S. muticum compared to native macroalgae. The presence of S. muticum did not affect the growth rates of any of the native species. Grazers affected the growth rate of all seaweeds, but S. muticum had the highest growth rates with and without grazers. For the south coast set of species, Stypocaulon scoparium was the most and S. muticum was the least or among the least preferred by grazers. The grazers from the southwest coast did not show a specific preference for C. humilis or S. muticum. Contrary to our expectations the food choice of mesoherbivores may provide S. muticum with some competitive advantage on the Portuguese south coast, but this may not be applicable to the Southwest communities.

The taxonomic identity of the common subtidal ctenostomate bryozoan Alcyonidium diaphanum is resolved using comparative morphological and molecular genetic techniques. The newly recognized lectotype series and topotype specimens are described and illustrated. The incorporation of spicules and foreign particles during colony growth is described. Colony morphology and genetic characters of the three commonly occurring morphotypes are examined, showing that just a single genetic species may be recognized, based on the 12s rRNA and COl mtDNA genes. Mitochondrial haplotypes show no general correlation with either colony morphotype or geographical location.

The distribution and relative abundance of macroepibenthic echinoderms in the eastern English Channel and Irish Sea is described from beam trawl catches. Echinoderms accounted for approximately 29% (by biomass) of fauna captured. A total of 24 species were recorded, including 12 species of starfish. The most frequently encountered species were Asterias rubens and Psammechinus miliaris, which were recorded at 85.5% and 56.0% of stations respectively. Asterias rubens and Ophiothrix fragilis accounted for 63.7% and 25.5% (by biomass) respectively of the echinoderms sampled. Mean echinoderm catches ranged from 0.8-kg h−1 in the north-eastern English Channel to 329-kg h−1 in the south-eastern Irish Sea. The echinoderm fauna was more diverse in the St George's Channel and western Irish Sea (6.7–7.0 species haul−1) than in the north-eastern English Channel (1.9 species haul−1).

Impact studies examining scallop dredging have been limited to loose-sediment habitats typical of most scallop grounds. However, it is likely that rocky-reef habitats also suffer damage from dredging activity since commercially viable grounds are often found in close proximity. This study presents a photographic survey of four experimental tows performed in two rocky-reef biotopes. Survey images show characteristic dredge damage in both study areas, with the greatest damage apparent on the site that is likely to have been unfished previously. Moreover, whilst vulnerable species such as the elephant sponge, Pachymatisma johnstonia, exhibited visible damage in the majority of recorded cases, damage to other epifauna, although in evidence, was restricted. This unexpected result is likely to be caused by the manner in which the scallop dredge fishes hard, uneven substrates. Hence, in instances were the morphology of the rockyreef is particularly uneven, damage to some epifaunal species may be incremental. This is in contrast to loose sediment habitats where the majority of damage occurs on the first tow of fishing gear through a pristine site. The importance of these findings to the planning of protected areas and the incorporation of fishing effects into ecosystem approaches of fisheries management is discussed.

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 opulations 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 ophiuridaggregations (e.g., storm waves, Seine floods and patches) are suggested to explain these patterns.

Ophiothrixfragilis is a suspension-feeder which forms dense aggregations (up to 2000 individuals m 2) in the Dover Strait. Its optimal suspension-feeding activity is expressed when currents are less than 20 cm s - 1. Thus, the duration of this behaviour over a tidal cycle depends on the tidal velocity, and can be very short during spring tides. During neap tides, suspension-feeding is almost continuous but the flux of nutrients (phytoplankton, organic carbon and nitrogen) remains low because current increases slowly after slack water, and resuspension phenomena do not occur. In contrast, during spring tides, the current speed rapidly increases and induces an important resuspension of particles. Thus, the flux of nutrients isvery important during the short duration of suspension-feeding of the O. fragilis population. Minimum gut contents were observed 3 h before slack current and maximum gut contents were observed about 1 h after slack water.

An extensive in situ survey of Kilkieran Bay (west coast of Ireland) established the presence and distribution of seven discrete epibenthic animal associations. Identified on the basis of qualitative examination and named for'one or more visuallydominating elements of the flora or fauna, these are: the Laminaria saccharina association, the Raspailia-Stelligera association, the Lithothamnium association, the Musculus discors association, the Upper Laminaria hyperborea association, the Lower Latninaria hyperborea association and the Axinella dissimilis association. The ecological background to these associations is discussed and a catalogue of their component species is included.

A population of Ascidia mentuh 0. F. Miiller on the Swedish west coast has been studied over an 8-yr period by aid of non-destructive (photographic) recordings of a vertical rock face at a depth of 25 m. Supplemental sampling by diving at a depth of 25 m was carried out over a I-yr period. A demographical analysis revealed elevated mortality during the late summer and during periods of especially low temperature. The mortality was found to be relatively independent of age and was alsorather uniform between different years and generations. The mortality is discussed in relation to reproduction and ambient temperature. Larval settling took place all the year round with maximum activity in August and September. but was subject to high interannual variability. It was concluded that variation in recruitment was the most important factor regulating populations of A. mmtula. The growth was measured from settling to death and showed great variability between single individuals and between different generations. Negative growth was recorded especially when the ambient temperature reached a high level. Histological examinations showed mature gonads over the full year of study, but with a quantitativedifference. Throughout the year. mature oocytes and sperms were present in the gonoducts in 50 to 80”,, of the collected animals and in close to IOO”,, during the peak of activity.

The common seastars Leptasterias polaris and Asterias vulgaris show competitive interactions in shallow subtidal communities in the northern Gulf of St. Lawrence, particularly during summer when aggregations of the two seastars forage on mussel beds at 1–2 m in depth. We examined interactions between the two seastars in a different situation, in a mussel bed at 6 m in depth (a rare situation in this region). In the deeper mussel bed, seastars were three times more abundant than in the shallower beds, and the mussels were larger. The deeper bed disappeared rapidly due to the intense predation. Although decreased prey abundance should have favored interference interactions, we did not detect either partitioning of mussels by size or avoidance of A. vulgaris by L. polaris as previously reported when mussels are in short supply in shallower water. The lack of an avoidance behavior by L. polaris, together with the higher proportion of L. polaris than A. vulgaris that were feeding, suggests that in this situation, the dominance of A. vulgaris (observed in shallower water) is attenuated, or that L. polaris may dominate.

The Northern Hemisphere has been warmer since 1980 than at any other time during the last 2000 years. The observed increase in 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 essential to formulate and inform better adaptive strategies to address the inevitable consequences of climate change.

The feeding behaviour of asteroids is usually analyzed with optimal foraging models, which assume that asteroids maximize the ratio of energy uptake relative to energy used in the feeding process. The behaviour of Asterias rubens feeding at subtidal beds of Mytilus edulis was studied in order to evaluate effects of mussel bed structure on predation rate and size selection of prey. Field observations showed that Asterias feeds on mussels whose size were the same or larger than the mean size for the population. The solid structure of interconnected mussels forming the bed restricts Asterias predation to only those mussels situated at the bed surface. A field experiment demonstrated that Asterias consumption rate of mussels was higher at beds with an intact structure of mussels compared to beds where the bands interconnecting the mussels were broken and mussel orientations changed. A laboratory experiment revealed that attachment of mussels did not affect starfish predation rates and indicated that change in orientation of mussels may reduce predation rates. In field and laboratory experiments, increased attachment of mussels was observed when exposed to Asterias. The study demonstrated that the feeding behaviour of Asterias is restricted by the complex structure of interconnected mussels in beds, which has to be integrated in optimal foraging models.

Populations of Caryophyllia smithii were studied at three sites of differing sedimentation regime in Lough Hyne, Ireland. The height, length, width and density of individuals were measured along with the depth of accumulated sediment on the rock substratum at each site. Calyx size was largest at the site of least sedimentation and smallest at the site of most sedimentation. In contrast, height of individuals was greatest at the site of most sedimentation and smallest at the site of least sedimentation. The height of individuals was correlated with the level of surrounding sediment (Pearson's correlation 0.836 P<0.05). The uppermost limit of C. smithii corresponded with the downward limit of the infralittoral zone at each site; 15, 13, and 12 m at the low, mid and highly sedimented site respectively. Density averaged over depth was greatest (70 ind m−2) at the site of moderate sedimentation and lowest (13 ind m−2) at the site of least sedimentation. Density was also found to increase with depth at the sites of moderate and high sedimentation.

Dogwhelks foraging for mussels in an aquarium followed tortuous search paths when recently fed and moved faster along straighter paths when hungrier. In nature this “area restricted searching” would tend to keep dogwhelks in areas of high prey density and the increased “directionality” of movement of hungry dogwhelks would tend to take them away from areas of low prey density. Dogwhelks encountered different-sized mussels in the aquarium in frequencies significantly different from those presented, both in terms of numbers and of surface area. Water currents may have caused the bias. Dogwhelks inspected encountered mussels by crawling over them for ≈ 1–2 h. A proportion of mussels were rejected after inspection. Relative abundances of different-sized accepted mussels were significantly different from those of encountered mussels, the mean size of mussels in the diet increasing significantly with increasing size of the dogwhelks. Size selection became more pronounced as dogwhelks, previously fed on barnacles, became experienced with mussels. A mussel encountered by an experienced dogwhelk was more likely to be attacked if closer to the preferred size of 20–25 mm shell length than the previously encountered mussel. Accepted mussels were inspected for longer than rejected mussels, perhaps as a result of selection of the drilling site. Dogwhelks previously fed solely on barnacles drilled in random positions on the mussel shell at first, but after consuming six to seven mussels they had developed a significant tendency to drill in the thinnest area of shells up to 25 mm in length and in the area over the digestive gland of larger shells. This learned behaviour reduced the drilling time for smaller shells by ≈ 27% and, whilst not reducing the drilling time for larger shells, it allowed direct access to the most profitable organs of the prey. Shells were penetrated at a rate of ≈ 0.36 mm·day−1. The percentage of mussel flesh extracted by adult dogwhelks decreased from ≈ 90% for 10-mm to ≈ 60% for 40-mm mussels, the discarded flesh being predominantly from the mantle and foot. The profitability of mussels, calculated as the ratio of predicted weight of flesh extracted to predicted handling time, increased monotonically with increasing mussel shell length. The learned tendency to drill in the thinnest part of the shell increased the profitability of mussels <25 mm by ≈ 17%. Dogwhelks engaged in handling mussels were sometimes interrupted or displaced by contesting dogwhelks, with a probability that increased as handling of the prey progressed. Interlopers devalued the prey of occupant dogwhelks by prolonging drilling and ingestion, by stealing flesh, or by displacing the occupant. The learned tendency of the dogwhelk to drill over the digestive gland of larger mussels, allowing quicker access to the most profitable tissue, perhaps maximizes the average energy return from larger mussels when at risk from interlopers. The preference of dogwhelks for mussels smaller than those predicted to be the most profitable could reflect the devaluation of larger mussels by the risk from interlopers or the increased risks of predation, dislodgement or desiccation associated with longer handling times for larger mussels.

Adult ascidians compensate for their inability to move by the shape of the body, orientation of the siphons, structure of the pharyngeal wall, and consistency of the test. Many of these features appear to enhance the organism's capacity to use mechanical energy available in the environment to maximize its interactions with it. Many are convergent features occurringirrespective of phylogeny while others are major determinants of phylogenetic diversity in the Ascidiacea apparently effected by one or the other pressure for protection from predation or mechanical damage, and for feeding.

Summary of interactions Source Type Confidence

Biological Traits Medium







Grey Literature not all peer reviewed











Biological Traits High

High

High

Technical Report High





Peer Reviewed Book

Biological traits of Alcyonidium diaphanum Peer Reviewed Book

Biological traits of Balanus crenatus Peer Reviewed Book

Biologuical traits of Echnius

High

High


Echinus sp. and their proximinity to kelp bed.

Peer Reviewed Journal Article

Aquaculture potential and biological traits of Echinussp.


Overview of marine ecosystem function and drivers



High

High

High



Biological Traits: Species taken from France but a laboratory experiment.


Biological traits of hermit crab Pagurus bernhardus

Peer Reviewed Book

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




Medium




Biological Traits Technical Report Medium



Biological traits - partiularly epifaunal species

Website based on published books


Peer Reviewed Book


Medium

Grey Literature Medium








Arctice location, though still the same species.


Pomatoceros Triqueter - Biological traits

Peer Reviewed Book












Biological Traits Low



Biological Traits Grey Literature High











Biological Traits Grey Literature Medium



















Biological Traits` Grey Literature High

Biological Traits` Medium

Biological Traits` High

Biological Traits` Medium






Website based on research







Biological Traits Global Database Medium







High

Medium


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 of other variables (e.g., depth, stratification, and tidal bed stress) were more regional.



High

Medium

Medium

High

Sediment particle transport (sediment re-working, bioturbation, feces and pseudofeces deposition) and water exchanges (permeability change,bioirrigation) at the water–sediment interface.


Environmental factors affecting scondary 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


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


Abiotic functions which are the most important in influencing benthic biological cmmunity composition. 16 abiotic variables explained 51% of faunal variability. Sediment characteristics, bathymetry, hydrodynamic conditions contributed to 26% of the variation. Physico-chemical properties explained 25% of the faunal variability.


High

Medium

Medium

High

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


hydrozoans, which make only a minor contribution to benthic community biomass, capture large amounts of zooplankton and seston, and that they may play an important role in transferring energy from pelagic to benthic ecosystems


Higher numbers of organisms that transfer sediment from the surface to depth in shelly-sand habitats than muddy habitats. More suspension-feeders were found in complex biogenic habitats; while simple mud habitats were least likely to have large organisms but most likely to have organisms producing mounds on the sediment surface.


Benthic suspension feeders (especially brittle stars) dominate on coarse sediments, with a low mud content. An abundnace of biogenic substrate favours sessile epifauna. Riverine inputs help to increase the abundance of phytoplankton in the water column as a food source, but yields seasonal variability in production. Stong currents are another important factor.


High

Medium

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.


Near-bed flow speeds decreased and bed shear stresses increased with bivalve density due to the bed roughness generated by the shallow burrowers.These results suggest that bed roughness generated by Austrovenus and their feeding currents are capable of influencing food supply to the bed.


High

Medium

High

Energy + Carbon transfer at the sediment surface boundary. Bioturbation increases oxygen penetration into sediments.


Seasonal variation in dominant functional groups corresponding to changes in salinity, oxygen, chlorophyl and nutrient concentrations. The taxa that showed clear seasonal changes were C. multisetosum, Streblospio shrubsolii, H. diversicolor, C. carinata and H. ulvae. In winter, filter feeders became increasingly more important. lthough burrowing species were the most abundant, seasonal differences were observed for some parts of the estuary. In summer, sessile species (e.g. Pomatoceros lamarcki and Modiolus modiolus) were only found at the lower estuary, while in winter they were absent.


Impacts of benthic infauna as modifiers of their habitat. Ecosystem functions include biogeochemical cycling, altered water column properties, altered hydrodynamics, sediment stability and oxygen penetraion.


Medium

High

Medium

Dominant marine fauna (e.g. crustaceans and echinoderms) from the Skagerrak increased O2 consumption and nutrient fluxes, sediment reactivity and sediment pore-water concentrations owing to the individual traits of the specific species that comprised the fauna rather than to enhanced functional diversity.


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


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


High

High

High

High

Impacts of benthic fauna on sediment stability.


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


Sediment type is a controling factor of macrofaunal distribution, although other factors are also likely to be important.


Stabelisig and destabelising effects of fauna on sediments.


Medium

Medium

Medium

Importance of brittlestars in sediment retention and biodeposition.


Benthic topography is a driver behind small-scale faunal variation. Ripples have a controling effect on benthic fauna, largely as a result of variation in sediment composition between peaks and troughs.


Connections between bioturbation and nitrogen cycling.


High

Medium

Medium

Medium

Influence of hydrodynamic flow on benthic species diversity and functioning.


Competition between benthic burrowing organisms and tube building polycheates.


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


Filter feeding bivalves cause local enrichment of carbon and nitrogen in surroundng sediments and are important diodepositers. As a result, surrounding fauna was noted to be more abundant and more diverse.


Medium

High

Medium

Grey Literature Medium

Temperature has an effect on sediment re-working and bioturbation. Lowest bio-transport was measured at temperatures between 1 and 6 degrees, highest biodiffusion occurred at 13 degrees. Sediment re-working slowed down at higher temperatures.


Drivers affecting macrobenthic and epibenthic species distribution in the North Sea.


Benthic communities typically associated with subtidal sediments do not commonly have planktonic larvae, instead release young at an early stage of adult life.


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

High

High

High

Lanice imporive habiat quality for other organisms by (1) changes in the hydrodynamics, (2) increases of the habitat stability and oxygen supply, and (3) a creation of habitat heterogeneity in a uniform environment. In this way, L. Conchilega alters the habitat characteristics and affects other organisms, and can therefore even be considered as an ecosystem engineer. In other words, L. conchilega patches are responsible for an increased habitat quality in an otherwise uniform habitat, which results in a higher survival of the surrounding benthic species.


Microalgae in subtidal sediments can have an influence on erosion potential and can stabelise sediments, in turn reducing wave energy.


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


High

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."


Role of echinoderms in marine ecosystems, feeding preference and broad scale ecosystem services performed.


Biologuical traits of Echnius

High

High

High

Only Viviparous Echnioderm. Habitats include: intertidal rock pools, coralline algae, underside of stones, stands of mussels, sponges and bryzoans, loose in substrate. Seasonal Feeding patterns.


E. cordatum, transfering hydrocarbons from surface sediments to deeper sediments and vice versa. Implications for rates and pathways during organic matter mineralisation.


Sediment type and associated fauna. North Sea.


High

High

High

High

Indirect effects of one species to another within a community.


Direct and indirect affects of Carcinus maenas predation on nematode community structure. Changes in sediment composition and predation.


Good description of the biogeochemical cycling of bioturbators (echinocardium), including figure.


Neopentadactyla mixta bioturbation description and feeding strategies.


High

High

High

High

Feeding strategy of Spatanga, bioturbation and some biogeochemical info.


Asterias rubens a keystone species for mussel predation.


Anemonia viridis associations with spider crab.


Association between Neopentadactyla mixta and Balcis alba


High

High

High


Liocarcinus - No apprent importance in benthic assemblages.


Aquaculture potential and biological traits of Echinussp.


Echinus sp. and their proximinity to kelp bed.


Biological traits and sensitivity of Sabellaria spinulosa


High

High


Medium

Sensitivity information of Sabellaria spinulosa

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


Drecreased levels of dissolved oxygen are being influenced by increased temperature and increased biomass input (spring blooms).


Overview of marine ecosystem function and drivers

Temperature affects on primary production and its limitations.


Medium

Medium

Medium

Medium

Medium

Medium

The oyster Crassostrea virginicaas a suspension feeder, through its natural process may aid denitrification of the estuarine. By excreating particulate nitrogen, the benthic microalgae community would increased, absorbing the inoganic nitrogen as well as fixing N2.


Sediment composition and suspension caused by wave energy

Biological textbook - industry experts. Sources form peer reviwed journals

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


All environments depend on the surface primary production. Phytoplankton die and sink to the sea bed where they are eaten by other species, or decay releasing organic material.




Medium

Medium

High

High

Turbidity negatively affects kelp growth High


Biological textbook

Secondary production of mobile invertebrate fauna in kelp forests increases with wave exposure levels. This faunal group has a key function in transferring kelpcarbon to higher levels in the food web.


Results show that faunal diversity increased with wave exposure, mainly due to increasing wave stress and habitat diversity. Abundance was highest at medium wave exposed stations, and lowest at low exposure stations.



High

Medium

Medium

Water motion has little impact on kelp growth.


Relative to more exposed rocky substrata at identical depths, bottoms beneath kelp canopies were exposed to weak fluid transport and were characterized by greater rates of deposition of particulates. Kelps inhibited transport of suspended particles from the overlying water column to the bottom.


Kelp canopies can affect benthic species recruitment through sedimentation, light blockage and ambient flow changes.


Medium

High

Low

Habitat structure from epilithic algae has a positive effect on benthic faunal abundance, although other factors may be more important. Eplithic biomass is linked to faunal biomass.


Overview of sponge ecosystem functions - impacts on substrate, bentho-pelagic coupling and associations with other organisms.


Biomass of algae correlated with changes in light availability, wave energy and sea temperature. Algaes subject to high levels of seasonal variability.


Low

Low

Low

Wave exposure is a controling factor on strucure, diversity and assmeblages of macroalgae


Urchins can be a controling factor on macroalgae and turf algae.


Sediment deposition significantly affects the local structure and diversity of macroalgal assemblages. Erect algae is more likely to be affected. Growth of algae was shown to be enhanced in periods of reduced sedimentation.


Low

High

Low

Depth is a crucial factor in determining macroalgal assemblage diversity. Abiotic factors (sedimentation, wave exposure, wave and water currents, slope of seabed) are important in algae distribution.


Grazing by Gibbula can have a negative impact on recruitment of other organisms in rocky suntidal habitats.


The complexity of habitats positively affects the density and richnedd of gastropods


Medium

Medium

High

High

Temperature is a controling factor in starfish feeding rates, as is light intensity. Asterias reubens is subject to strong seasonal variation.


Mytilus sp. is a habitat-modifying species that can influence biodiversity by facilitation and inhibition of species. Species diversity of macro and meio fauna greatly increased in the presence of mussel beds. Sediment contents of total carbon and nitrogen were significantly higher within patches than outside, mainly due to the mussels’ filtration and biodeposition of small nutrientrich particles.


Gastropod grazers including Patella and Gibbula are controlers of marine algae.


Brittlestar feeding intensity is influenced by tidal flow. Brittlestars important for the flux of nutrients in marine ecosystems.


Low

Low

Medium

Strong link between structual complexity of habitat and meiofaunal/macrofaunal abundance and diversity.


Current speed influences featherstar feeding.


Mytilus are important for sedimentation and biodeposition through the production of faces and pseudofaeces.


Medium

Grey Literature

Low

Species richness and biomass of associated macrofauna and -flora was enhanced even by the presence of single mussels and increased rapidly with increasing mussel numbers. This is mainly due to physical sructure and creation of habitat. Abundance and biomass of associated flora and fauna were higher if live mussels were present because mussel biodeposition and nutrient regeneration supplied limiting resources and increased carrying capacity.


Another effect of mussel beds is an increased deposition of fine sedimentary and organic material on the sea bed because of the filtering and particle selection activities of the mussels and the reduction of current velocities by the structure of the bed. With these habitat modifying properties mussels can be regarded as physical ecosystem engineers.

Grey Literature High


Biological Traits Global Database High


Kelp biotopes, with their enormous numbers of species, high biomass and high rates of productivity will be important nursery areas for a diverse range of species. It is likely that juvenile forms of all the animals that are present as adults in the kelp bed make use of the habitat as a nursery area. The zonation patterns of the understorey algae are related to the available light rather than to the physical depth. It is probable that kelp plants and other algae are in direct competition with the colonial tunicates, sponges etc. for space on the substratum. The high productivity of kelp forests (mg C m-2 yr- 1) in comparison to other marine biotopes (Mann, 1972b) suggests that the surrounding III. Biology and ecological functioning Vol. VII. Infralittoral reef biotopes with kelp species 64 coastal areas are dependent on the kelp biotopes as a major source of food energy. 90% of kelp production is estimated to enter the detrital food webs as POM or DOM, being exported from the immediate area of the kelp bed. The blades of mature kelps in the kelp forest form a canopy layer which, under certain conditions, may cut off as much as 90% of the incident irradiance.




High











Biological Traits/Interactions Low















Biological textbookBiological textbook


Biological Traits Grey Literature Medium












Medium

Biotope classification High



Technical report by established agency

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

Salinity preferenceTidal Stream Preference

Physiographic preferences

Lifespan

Sensitivity To Change

Temporal Variable Population

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

Standardised Species Trait Categories

Growth FormBoringCrustose hardCrustose softFlaccidMassiveCushionTurfFolioseShrubArborescent / ArbuscularForestAlgal gravelAccretion

Growth Form

Feeding Method

Environmental Position

Habitat

Size

Distribution

Substratum Preference

Key Prey Taxa

Connectivity to other habitats /species

MatFaunal 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

2. Water Column Processes

3. Local Processes/Inputs at the Seabed

5. Output Processes

Table 4. Model Component Definitions

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

Suspended SedimentLight Attenuation

Dissolved Oxygen

Water Chemistry & Temperature3. Local Processes/Inputs at the Seabed

RecruitmentFood Sources

- Plankton

- POM (Particulate Organic Matter)

- Detritus - Algae - Carrion - Small Invertebrates and Fish - Micro-organismsGrazing and PredationSeabed Mobility4. Habitat and Biological Assemblage

4. Habitat and Biological Assemblage

6. Local Ecosystem Functions

7. Regional to Global Ecosystem Functions

Macroalgae

Attached Active Filter Feeders

Attached Passive Filter Feeders

Tube Building Fauna

Scavengers and Predatory Fauna

Non Predatory Mobile Fauna

OUTPUTS5. Output Processes

Primary Production


Bio-deposition

Hydrodynamic Flow

Bioengineering

Supply of Propagules


Nutrient Cycling

Biogeochemical CyclingFood Resources

Population Control

Control of Algal Growth

Sediment Stability

Habitat Provision

7. Regional to Global Ecosystem FunctionsExport of Biodiversity

Export of Organic Matter

Biodiversity Enhancement

Biotope Stability

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

Bivalves, Brachiopods and Other Encrusting Filter Feeders

Table 5. Direction of Interactions

Direction of Interaction

Positive

Negative

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 favourThe salinity the organism favoursFavoured location in relation to tidal currents

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

An organisms connection and influence to other species or habitats

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


Mobility / MovementSwimmerCrawlerBurrowerDrifterTemporary attachmentPermanent attachmentInsufficient informationNot relevantField unsearchedTemporary attachment / CrawlerCrawler / SwimmerSwimmer / BurrowerSwimmer / Crawler / Burrower

The organisms ability and method of locomoting through its environment

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 regarded as a key food source for other organisms The variation in an organisms population in relation to environmental conditions or annual seasonal fluxes

Swimmer / 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

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 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

3. 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 bivalves, polychaetes or small fishMicrobial organisms (e.g. bacteria, diatoms and protozoa)The predation of organisms by anotherMovement of sediment on the seabed

4. Habitat and Biological Assemblage

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 column

Ecosystem 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 dissolved oxygen concentration in the water column above the seabed

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)Non-living material derived from organic sources within the water column

Marine algae including red, green, brown and kelp variants

OUTPUTS5. Output Processes

Amount of biomass created as a direct result of consumption

Changes to water flow/movement as a result of organism activity


The cycling of organic carbon and nitrogen other than nutrientsThe provision of food resources for other organisms

Control of lower trophic level organism population through predation

Control of algal biomass through grazing pressure

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

Organisms attached to the seabed surface which are predominantly active filter feeders, e.g. create water flow to feed

Organisms attached to the seabed surface which are predominantly passive filter feeders, e.g. rely on natural water movement to feed

Boring and attached bivalves and brachiopods as well as other surface encrusting filter feedersTubicolous fauna which construct and live in tubes made from sedimentary material on the surface of the seabedMobile scavenging and predatory crabs, echinoderms, polychaetes and molluscsMobile fauna which are either predominantly grazers or which feed on detritus or POM

The production of energy through the process of photosynthesis (typically by algae and plankton in the marine environment)

The process by which organisms either deposit material onto the seabed, e.g. through the capture of particulate matter in the water column through filter feeding or the production of faeces/pseudo-faeces

Faunal modification of the natural habitat, e.g. tube building, boring organisms, algal canopy etc. The production and transportation of larvae, spores or body fragments capable of regeneration.

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

Cohesion of sediments into a stable form more resistant to disturbanceProvision 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 within the habitat resulting from increased sediment stability and habitat provisionStability of the habitat through the habitat provision and increased sediment stability

Table 5. Direction of Interactions

Definition

Table 6. Magnitude of Interactions

Requirement

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.

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.


Feeding MethodPhotoautotrophActive Suspension FeederPassive Suspension FeederSurface Deposit FeederSub-surface Deposit FeederOmnivoreHerbivoreScavengerSymbiont contributionPlanktotrophChemoautotrophPredatorInterface feeder

Grazer (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


Environmental 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 TubicolousEpizoicNeustonicPleustonicLithotomous

Hyperbenthic

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