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Please quote as: Lancaster, J. (Ed.), McCallum, S., Lowe A.C., Taylor, E., Chapman A. & Pomfret, J. (2014).
Development of detailed ecological guidance to support the application of the Scottish MPA selection guidelines in
Scotland’s seas. Scottish Natural Heritage Commissioned Report No.491. Shallow Tide-Swept Coarse Sands with
Burrowing Bivalves – supplementary document.
Shallow Tide-Swept Coarse Sands with Burrowing
Bivalves
Biotope:
SS.SCS.ICS.MoeVen
Moerella1 spp. with venerid bivalves in infralittoral gravelly sand
Offshore/Territorial waters: Territorial
The ‘shallow tide-swept coarse sands with burrowing bivalves’ search feature is
characterised by infaunal burrowing bivalves and polychaetes. The burrows and tubes
of the infaunal species stabilise the sediment and provide structural complexity, which
in turn provides habitats for other species. Burrowing infauna also greatly increase
bioturbation which has been shown to have a beneficial effect on the maintenance of
biodiversity (Widdicombe et al., 2004). Burrowing fauna such as polychaetes
significantly influence nutrient fluxes of nitrogen and phosphorus at the sediment-water
interface, as their burrowing activity promotes oxygenation of the substrata
(Widdicombe & Austen, 1999).
Long-lived venerid bivalves, a major constituent of this feature, provide a food source
for many species of fish, invertebrates and birds (Malthus et al., 2006; Durkin, 2008;
Howson et al., 2009), and represent an example of a bentho-pelagic link. This feature
is very sparsely distributed around the UK and Ireland and therefore Scottish records
are likely to be of national importance (Connor et al., 2004).
Functional Links
Functional links and associations with other Priority Marine Features

Eelgrass beds: Shallow tide-swept coarse sands with burrowing bivalves may
provide a habitat for the formation of Zostera marina/angustifolia beds on lower
shore or infralittoral clean or muddy-sand (SS.SMp.SSgr.Zmar).

Maerl beds: Scottish maerl beds can occur in association with shallow tideswept coarse sands with burrowing bivalves and, where this occurs, may have
an epifauna typified by crustaceans and echinoderms (Malthus et al., 2006).
Functional links with the wider Scottish marine ecosystem
Long-lived venerid bivalves provide a food source for many species of fish (such as
gurnards, dragonets and whiting), crustaceans and echinoderms, representing an
example of a bentho-pelagic link (Durkin, 2008). They are important in supporting
commercial fisheries e.g. for bivalve and flatfish species, the juveniles of which have
been observed to use this habitat (Malthus et al., 2006). Long-lived venerid bivalves
can be an important food source for birds (Howson et al., 2009).
1
This name of this group has now been changed to Tellina, although the biotope name remains Moerella
1
The infaunal species within this habitat stabilise sediments and provide structural
complexity, in otherwise habitat-poor areas, through the creation of burrows and tubes
which increase biodiversity (Widdicombe et al., 2004). Additionally, infralittoral rock with
Laminaria hyperborea and Saccharina latissima have been found alongside this habitat
(Malthus et al., 2006).
Biological Diversity
Habitat/Biotope description for Scottish waters
The following habitat description is taken directly from JNCC’s Marine Habitat
Classification Hierarchy (Connor et al., 2004). Below the box this has been
supplemented with additional information specific to Scotland from the scientific
literature.
Moerella spp. with
(SS.SCS.ICS.MoeVen).
venerid
bivalves
in
infralittoral
gravelly
sand
“Infralittoral medium to coarse sand and gravelly sand which is subject to moderately
strong water movement from tidal streams may be characterised by Moerella spp. with
the polychaete Glycera lapidum (agg.) and venerid bivalves. Typical species include
Moerella pygmaea or M. donacina2 with other robust bivalves such as Dosinia lupinus,
Timoclea ovata, Goodallia triangularis and Chamelea gallina. Other infauna include
nephtyd and spionid polychaetes and amphipod crustacea. Another important
component of this biotope in some areas is the bivalve Spisula solida (see Khne &
Rachnor 1996) which may be common or abundant. In conjunction with .FfabMag3 this
biotope may form part of the 'Shallow Venus Community', the 'Boreal Off-shore Sand
Association' and the 'Goniadella-Spisula association' of previous workers (see
Petersen 1918; Jones 1951; Thorson 1957; Salzwedel, Rachor & Gerdes 1985).
Epifaunal communities may be reduced in this biotope when compared to .FfabMag;
both types may have surface sand waves which may be indicative of the presence of
venerid bivalves (Warwick & Davies 1977). This hypothesis, however, requires testing.
Remote grab sampling is likely to under-estimate venerid bivalves and other deepburrowing and more dispersed species such as Paphia, Ensis and Spatangus. In
southern areas of the UK and the North Sea, in slightly siltier sand and shelly sand,
SCS.MoeVen4 may give way to the other Spisula biotope SSA.SsubNhom5. Together
these two biotopes replace the old biotope IGS.Sell6.” (Connor et al., 2004)
2
Tellina pygmaea and T. donacina
3
Fabulina fabula and Magelona mirabilis with venerid bivalves and amphipods in infralittoral compacted
fine muddy sand (SS.SSa.IMuSa.FfabMag)
4
Moerella spp. with venerid bivalves in infralittoral gravelly sand
5
SS.SSa.IMuSa.SsubNhom - Spisula subtruncata and Nephtys hombergii in shallow muddy sand
6
SS.IGS.FaG.Sell - Spisula elliptica and venerid bivalves in infralittoral clean sand or shell gravel
2
Studies from the Outer Hebrides provide further information on the diversity of this
habitat in Scotland. In the Sound of Barra this habitat has been recorded containing
both T. pygmaea and T. donacina, however T. donacina was recorded in very few
locations, and T. pygmaea was the dominant tellin, found during both infaunal and
epifaunal surveys (Harris et al., 2007). The polychaete Nephtys cirrosa and amphipod
Bathyporeia were also recorded in abundance in this habitat (Harris et al., 2007). In the
Sound of Harris only T. pygmaea was found, although a wide range of associated
habitats and species were described occurring alongside this habitat (Malthus et al.,
2006). Habitats such as those dominated by Laminaria hyperborea and Saccharina
latissima were found adjacent to this habitat, as were eelgrass beds and maerl beds
(Phymatolithon calcareum) (Malthus et al., 2006). Additional infaunal organisms
present included Arenicola marina, Amphiura sp., Lanice conchilega and the burrowing
anemone Cerianthus lloydii. Epifauna recorded in the Sound of Harris included Echinus
sp., Asterias sp., Carcinus sp., hydroids and ascidians (e.g. Ascidia mentula) and a
diatom film was also found on top of the sediment.
Species diversity
Records from the Outer Hebrides (Sounds of Barra and Harris) indicate the typical
levels of biodiversity present in this habitat in Scotland. The number of species per
grab (0.1m2) varied, with generally more species being found in the Sound of Harris (853 species; Malthus et al., 2006) compared to the Sound of Barra (10-29 species;
Harris et al., 2007). Species richness indices values per grab (Shannon Index) also
varied from 1.866-3.325 in the Sound of Harris (Malthus et al., 2006) to 1.10-2.93 in the
Sound of Barra (Harris et al., 2007).
Key and characterising Species
These have been taken from JNCC’s biotope description (Connor et al., 2004).
Biotope type
SS.SCS.ICS.MoeVen
Key species for identification
The tellins T. pygmaea and T.
donacina.
3
Additional characterising species
Glycera lapidum, S. solida, A. latreillii,
Anemonia viridis, Nemertea, Nematoda,
Pisione remota, Glycera lapidum,
Streptosyllis websteri, Nephtys cirrosa,
Aonides paucibranchiata, Spio filicornis,
Spiophanes bombyx, Lanice conchilega,
Bathyporeia pelagica, Apseudes latreillii,
Pagurus bernhardus, Liocarcinus depurator,
Carcinus maenas, G. triangularis, Spisula
solida, Chamelea gallina, T. ovata; Dosinia
lupinus; Echinocyamus pusillus;
Callionymus lyra, Pomatoschistus minutes,
Chondrus crispus, Polyides rotundus,
Gracilaria gracilis, Saccharina latissima and
Ulva spp.
Coherence
Typicalness
This habitat is found on coarse, gravelly infralittoral sand (0-20m) on exposed open
coasts and in estuaries, subject to moderately strong tidal water movement (Connor et
al., 2004). It is characterised by Tellina spp. with the polychaete Glycera lapidum and
venerid bivalves. Other infaunal species include other polychaete, tanaids (shrimp-like
crustaceans) and sand hoppers (Connor et al., 2004). Many of the faunal components
of this habitat are moderately robust as the environment is relatively dynamic and sand
ripples may be seen on the surface of the sediment (Connor et al., 2004). This habitat
has been found in association with eelgrass beds and maerl beds in Scotland, and
where this occurs, may have an epifauna characterised by crustaceans and
echinoderms (Malthus et al., 2006).
Ecological variations across Scottish waters
The species composition of this habitat is thought to be largely determined by
substratum characteristics and hydrodynamics (Durkin, 2008). Around the islands of
Yell, Fetlar and Unst (Shetland) associated species vary according to location and
include Moerella pygmaea (now Tellina) in the majority of samples, as well as maerl P.
calcareum at a few locations, together with the bivalve Spisula elliptica, the razor shell
Ensis arcuatus, and the polychaetes L. conchilega, Glycera spp., Spio spp.,
Spiophanes spp. and Aonides paucibranchiata (Howson,1988; Hirst et al., 2013). The
habitat was found often situated adjacent to records of the related but deeper
SS.SCS.CCS.MedLumVen7 biotope.
In Lamlash Bay (Arran) this habitat can be found in, muddy, gravelly sand with some
shell fragments and supported clumps of red algal turf (Aglaothamnion sp. and other
species) and bootlace weed C. filum. Mobile species included Frequent edible crabs C.
pagurus and hermit crabs Pagurus bernhardus, the gobies P. pictus and P. minutus,
and juvenile common starfish A. rubens. Some samples contained relatively high
densities of other venerid bivalves including the striped venus Chamelea gallina and
smooth Artemis Dosinia lupinus, and low densities of the polychaetes Mediomastus
fragilis and Lumbrineris gracilis (Munro et al., 2012). These samples share
characteristics with the related SS.SCS.CCS.MedLumVen biotope and areas of sea
bed within Lamlash Bay were considered to comprise a mosaic of
SS.SCS.ICS.MoeVen and SS.SCS.CCS.MedLumVen biotopes in 2010 (Munro et al.,
2012)
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Mediomastus fragilis, Lumbrineris spp. and venerid bivalves in circalittoral coarse sand or gravel
4
Viability
Very little information exists for the bivalves present in this community, with no
information available regarding population densities or Minimum Viable Populations.
There are also only a few locations where this search feature is known within Scotland.
In most known instances (e.g. Malthus et al., 2006) the area covered by this feature is
much smaller than would be needed to protect the entire life cycle of the bivalves.
Therefore, regardless of the size of the feature, it would be recommended that
examples of its entire extent are protected. Where the feature occurs in discrete
patches, amongst other habitats such as maerl beds, seagrass beds, kelp or reefs, it is
recommended that all of the patches of this search feature within specific examples of
these habitat mosaics are protected.
Fragmentation
No detailed information was available on the level of fragmentation typical for this
habitat. However, it is known to occur in small discrete patches amongst other habitats.
Indicators of Least Damaged/More Natural
Up to date information on the sensitivity of this search feature to pressures associated
with human activities are included in the Feature Activity Sensitivity Tool (FeAST;
Marine Scotland, 2013). Below, information on indicators of damage and naturalness
are taken from MarLIN sensitivity data (Durkin, 2008).
Table 1: Indicators of damage and naturalness
Indicators of Naturalness
Indicators of Damage
Potential Sources of Damage
Continued presence of this
biotope and its characteristic
species, including S. solida and
other populations of infauna and
sessile epifauna present.
Loss of habitat feature and/or
associated species diversity
particularly loss or reduction of S.
solida.
Physical disturbance, substrate
removal, changes to
hydrography.
No reduction in species diversity
and/or abundance.
Notable reductions in species
abundance and/or diversity.
Pollution/Eutrophication
Risk Assessment
The details of the assessment of risk for each MPA search feature is addressed in a
separate report (Chaniotis et al., 2014).
Recovery Potential
Physical damage/substrate removal hydrographic change
MarLIN lists the .MoeVen biotope’s ability to recover from substratum loss and physical
damage as high, however, this will depend on whether or not sufficient numbers of
individuals remain to supply recruits to the area (Durkin, 2008). A loss of key
characterising species will take up to 5 years to recover, due to the time required for
the majority of key bivalves to reach maturity (Durkin, 2008). However, the rarity of
some characterising species in British waters means that there may be a lack of recruit
sources to seed a particular area, causing recovery to be hindered or not possible.
5
Changes to hydrography
Permanent changes in the hydrography caused by development or coastal defence
may cause changes in the benthic conditions rendering the habitat unsuitable for
recolonisation. MarLIN lists the .MoeVen biotope as being intermediate in its tolerance
to increases in water flow rate and its sensitivity is moderate since its ability to recover
is high if the activity is only temporary (Durkin, 2008).
Contamination by pollutants
MarLIN lists this .MoeVen biotope as being highly intolerant to contamination with
pollutants and its sensitivity as moderate since its ability to recover from pollution is
high after the cessation of the impacting activity (Durkin, 2008).
Eutrophication
MarLIN lists the .MoeVen biotope as being intermediate in its tolerance to
eutrophication and its sensitivity as moderate since its ability to recover from
eutrophication is high after the cessation of the impacting activity (Durkin, 2008).
Geographical Variation
This search feature is uncommon in the British Isles, and in Scotland has only been
recorded in the West and North territorial MPA regions in territorial waters. The majority
of these records are from Shetland and Orkney with some instances on the west coast
of Scotland (e.g. Loch Slapin, Loch Kishorn and Arran) and Outer Hebrides (e.g.
Sounds of Barra and Harris) (Connor et al., 2004 ).
Geographical context
Beyond Scotland this habitat is very scarcely distributed within the UK and Ireland, with
a few records reported from south Devon and Cornwall, the Scilly Isles and north-west
Wales (Connor et al., 2004). Scottish records of this habitat are therefore likely to be of
national importance.
6
References
Connor, D. W., James A. H., Golding, N., 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
Chaniotis, P.D., Cunningham, S., Gillham, K. & Epstein, G. 2014. Assessing risk to
Scottish MPA search features at the MPA regional scale. Final report produced by the
Joint Nature Conservation Committee, Scottish Natural Heritage and Marine Scotland
for the Scottish Marine Protected Areas Project.
Durkin, O.C. 2008. Moerella spp. with venerid bivalves in infralittoral gravelly sand.
Marine Life Information Network: Biology and Sensitivity Key Information Subprogramme [on-line]. Plymouth: Marine Biological Association of the United Kingdom.
[cited 12/04/2010]. Available from: http://www.marlin.ac.uk/habitatecology.php?
habitatid=388&code=1997
Harris, D.B., Malthus, T.J., Karpouzli, E., Moore, C.G., Lyndon, A.R., Mair, J.M.,
Service, M. & Mitchell, A. 2007. Biotope mapping of the Sound of Barra, Scotland.
Scottish Natural Heritage, Commissioned Report No.258 (ROAME No. F06PA04).
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Network Principle of Viability: Feature specific recommendations for species and
habitats of conservation importance. Natural England Commissioned Reports, Number
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Hirst, N.E., Kamphausen, L.M., Cook, R.L., Porter, J.S. & Sanderson, W.G. 2013. The
distribution and status of proposed protected features within the Fetlar and Haroldswick
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Howson, C.M. 1988. Marine Nature Conservation Review: Survey of Shetland, Foula
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of Harris, Scotland. Scottish Natural Heritage, Commissioned Report No. 212 (ROAME
No. F01AC401/2).
Marine Scotland. 2013. Features, Activities, Sensitivities Tool (FeAST) online resource.
Available from:
http://www.scotland.gov.uk/Topics/marine/marine-environment/FEAST-Intro
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habitats and species in Lamlash Bay, Arran: methodological developments and 2010
baseline data. Scottish Natural Heritage, Commissioned Report.
Widdicombe, S. & Austen, M.C., 1999. Mesocosm investigation into the effects of
bioturbation on the diversity and structure of a subtidal macrobenthic community.
Marine Ecology Progress Series, 189, 181-193.
Widdicombe, S. & Austen, M.C. 1998 . Experimental evidence for the role of Brissopsis
lyrifera (Forbes, 1841) as a critical species in the maintenance of benthic diversity and
the modification of sediment chemistry Journal of Experimental Marine Biology and
Ecology, 228, 241–255
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Widdicombe, S., Austen, M.C., Kendall, M.A., Olsgard, F., Schaanning M.T, Dashfield,
S.L, & Needham. H.R. 2004. Importance of bioturbators for biodiversity maintenance:
indirect effects of fishing disturbance. Marine Ecology Progress Series, 275, 1-10.
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