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. Northern Sea Fan Communities –
supplementary document.
Northern Sea Fan Communities
Component of MPA search feature: Northern sea fan and sponge communities
Biotopes:
CR.HCR.XFa.SwiLgAs
Mixed turf of hydroids and large ascidians with Swiftia pallida and
Caryophyllia smithii on weakly tide-swept circalittoral rock
CR.MCR.EcCr.CarSwi
Caryophyllia smithii and Swiftia pallida on circalittoral rock.
Sub-biotope: CR.MCR.EcCr.CarSwi.LgAs
Caryophyllia smithii, Swiftia pallida and large solitary ascidians on
exposed or moderately exposed circalittoral rock.
Territorial/Offshore waters: Territorial
The boulders and bedrock found in northern sea fan communities create a complex
environment resulting in a number of ecological niches, i.e. boulders, crevices,
overhangs, under boulder areas, bedrock, and silt. The presence of algae and sessile
fauna provides additional attachment sites and living space for other organisms. For
example, Swiftia pallida is the only known habitat (host) for the nationally rare sea fan
anemone Amphianthus dohrnii in Scotland (Hiscock et al., 2001). Boulders are
characteristically colonised by the cup coral Caryophyllia smithii and the sea fan S.
pallida along with other soft corals. On the rocky surfaces a variety of epibionts can be
found such as bryozoans, encrusting red algae and tube building polychaetes, whilst
overhangs and crevices shelter sea cucumbers, squat lobsters and wrasse (Connor et
al., 2004.).
Functional Links
Functional links and associations with Priority Marine Features

Pink sea fingers:
The hard rock substratum within northern sea fan
communities provides a source of attachment for the octocoral Alcyonium
hibernicum (pink sea fingers), which can therefore be found in the same
location as Caryophyllia smithii and Swiftia pallida. The functional significance
of the association is not known.

White cluster anemone; European spiny lobster: Predatory species associated
with Nothern sea fan communities, such as the white cluster anemone
(Parazoanthus anguicomus) and European spiny lobster (Palinurus elephas),
will remove a number of organisms through predator-prey interactions and may
also create additional living space by this process.

Northern feather star: The northern feather star (Leptometra celtica) can be
found in the sub-biotope CR.MCR.EcCr.CarSwi.LgAs; however, it is unknown
how this species contributes towards the functioning of the ecosystem other
than filter-feeding on plankton.

Kelp and seaweed communities in tide-swept sheltered conditions: Sheltered
kelp forests can often be found above the biotope CR.HCR.XFa.SwiLgAs and
the sub-biotope CR.MCR.EcCrCarSwi.LgAs (Connor et al., 2004; Seasearch
Firth of Lorn sea fan surveys, 2007-2009). The kelp forest may incorporate the
biotope Kelp and seaweed communities in tide-swept sheltered conditions
(IR.MIR.KT). The exact manner of the functional link with northern sea fan
communities has not been specifically studied; however, kelp plants provide an
important source of particulate organic matter (POM) through the erosion or
decay of the outer edges of the kelp blades and frond (Birkett et al., 1998). This
POM is an important source of food for many filter feeders, allowing an increase
in biological diversity to develop within the surrounding waters. As well as
increasing levels of POM within the area, kelp plants can reduce current flow,
attenuate wave energy, reduce light availability (Birkett et al., 1998), thereby
facilitating conditions that favour sessile animals rather than plants.

Burrowed mud: In deeper water, below the biotope CR.HCR.XFa.SwiLgAs and
sub-biotope CR.MCR.EcCrCarSwi.LgAs, burrowed mud habitats have been
recorded (Connor et al., 2004; Seasearch Firth of Lorn sea fan surveys, 20072009). These contain organisms such as the sea pens Pennatula phosphorea
and Funiculina quadrangularis, the mud burrowing amphipod Maera loveni, and
the echiuran worm Maxmuelleria lankesteri.

Deep sponge communities: Sponges are often found within the
CR.HCR.XFa.SwiLgAs biotope, and where this biotope reaches a depth of 20m
or more may crossover, to some extent, with the biotope complex
CR.HCR.DpSp (Deep Sponge Communities). It is unknown if these
communities have a functional link despite having a recorded association.
Functional links with wider Scottish marine ecosystem
The abundance of epibiont filter feeding organisms in these environments (e.g.
anthozoans, ascidians, hydrozoans and poriferans) play a significant role in regulation
of the plankton at a local scale (Officer et al., 1982). The plankton consumed by filter
feeders takes a variety of forms including detritus, bacteria, phytoplankton and
zooplankton (including eggs and larval stages of nekton and benthos); thus they are
important consumers of the lowest trophic levels.
2
Biological Diversity
Habitat/Biotope description for Scottish waters
There are two biotopes and one sub-biotope described for northern sea fan
communities that occur in Scottish waters Their habitat descriptions are taken directly
from JNCC’s Marine Habitat Classification Hierarchy (Connor et al., 2004) (see boxed
text). Below each box this has been supplemented with additional information specific
to Scotland from scientific literature.
Caryophyllia
smithii
(CR.MCR.EcCr.CarSwi)
and
Swiftia
pallida
on
circalittoral
rock
“This biotope is typically found on the upper and vertical faces of very exposed through
to wave-sheltered circalittoral bedrock and boulders, which are typically subject to
weak tidal streams. It is characterised by dense aggregations of the cup coral C.
smithii and the sea fan S. pallida on the silty substratum. Under the silt, bryozoan
crusts such as Parasmittina trispinosa and encrusting red algae may be seen. This
biotope may have a grazed appearance, perhaps attributable to the frequently
occurring E. esculentus. There may be a sparse hydroid turf present, with species
such as Nemertesia antennina, Nemertesia ramosa and Halecium halecinum present.
The soft corals A. glomeratum and A. digitatum may be present on the tops of boulders
along with the crinoids Antedon petasus and Antedon bifida. Other echinoderms
occasionally observed include the starfish Marthasterias glacialis, Asterias rubens and
Luidia ciliaris. Sponges feature only occasionally in this biotope, including species such
as the boring sponge Cliona celata. The bryozoan Porella compressa may also be
recorded. Ascidians occasionally present include Ascidia mentula, Clavelina
lepadiformis and Ciona intestinalis. Under-boulder fauna typically consists of the
crustacean M. rugosa. The polychaete P. triqueter may be seen encrusting the rocky
surface” (Connor et al., 2004).
There is one sub-biotope of CR.MCR.EcCr.CarSwi which is described below:
Caryophyllia smithii, Swiftia pallida and large solitary ascidians on exposed or
moderately exposed circalittoral rock (CR.MCR.EcCr.CarSwi.LgAs)
“This variant typically occurs on exposed to moderately wave-exposed, circalittoral
bedrock and boulders rock subject to mainly weak tidal streams and has a thin layer of
silt present. It is found predominantly from 10-30m water depth. From afar, this
biotope is mostly distinguished by the frequently occurring sea fan S. pallida,
encrusting red algae and the abundant cup coral C. smithii. This sub-biotope has quite
an impoverished appearance, compared with CR.HCR.XFa.SwiLgAs1 which has a
strong sponge component. Other species are typically present in low abundance.
Echinoderms such as E. esculentus, A. bifida, A. petasus, Leptometra celtica, M.
glacialis, L. ciliaris and A. rubens may be recorded. Large hydroids such as N.
antennina and N. ramosa may occasionally be seen in isolated clumps on the tops of
boulders and rocky outcrops. The anthozoan Parazoanthus anguicomus may be
recorded. Bryozoans such as Parasmittina trispinosa and Porella compressa are
occasionally observed. The polychaete P. triqueter may be observed encrusting the
1
Mixed turf of hydroids and large ascidians with Swiftia pallida and Caryophyllia smithii on
weakly tide-swept circalittoral rock.
3
sides of rocks and boulders while occasional A. digitatum may also be seen. A small
suite of large ascidians may be present, including A. mentula, C. lepadiformis, C.
intestinalis, Diazona violacea and Ascidia virginea. Sponges are typically absent from
this biotope, although Cliona celata may be recorded occasionally. The top shell
Gibbula cineraria is usually present. Under boulders and overhangs, the squat lobster
M. rugosa can usually be seen hiding.” (Connor et al., 2004).
A Seasearch survey in 2005 recorded this biotope about 500m south of Culanach Reef
in the Firth of Lorn and noted the presence of the spiny lobster (Palinurus elephas) in
the area. The cup coral Caryophyllia inornata reaches its northern limit in the Firth of
Lorn and is found in the vicinity of Swiftia pallida within this region (Howson, 1990).
Mixed turf of hydroids and large ascidians with Swiftia pallida and Caryophyllia
smithii on weakly tide-swept circalittoral rock (CR.HCR.XFa.SwiLgAs)
“This biotope is distinguished by frequently occurring S. pallida, abundant C. smithii
and a diverse range of ascidians including Clavelina lepadiformis, Ascidia mentula,
Polycarpa pomaria, Diazona violacea and Corella parallelogramma. A sparse, yet
diverse hydroid turf is often apparent, with species such as Aglaophenia tubulifera,
Nemertesia antennina, Polyplumaria frutescens, Halecium halecinum, Abietinaria
abietina, Nemertesia ramosa and Halopteris catharina often recorded. Spaces
amongst the turf are usually colonised by the polychaete Pomatoceros triqueter and
encrusting red algae. Crinoids such as Antedon petasus, Antedon bifida and
Leptometra celtica may be seen filter feeding on the tops of outcrops and boulders,
along with the soft coral Alcyonium digitatum. Other echinoderms such as Echinus
esculentus, Crossaster papposus and Asterias rubens may also be recorded. There
may also be a bryozoan component to the sparse faunal turf. Species such as
Securiflustra securifrons and Eucratea loricata as well as the crustose Parasmittina
trispinosa are all usually present. There may be a few isolated growths of sponge,
such as Iophonopsis nigricans, Axinella infundibuliformis and Haliclona urceolus.
Other species that may be present include the brachiopod Terebratulina retusa and the
top shell Calliostoma zizyphinum. The crustacean Munida rugosa may be visible in
crevices.” (Connor et al., 2004).
A survey of the Sound of Harris (Malthus et al., 2006) found this biotope at a number of
sites in the eastern end of the Sound and included the following organisms within the
community: the hydroids Tubularia sp, Nemertesia sp, Abietinaria sp; the anemone
Protanthea simplex; the bryozoan Alcyonidium diaphanum; the tunicates Ascidia
mentula and Diazona violacea; the crustacean Munida rugosa and several Axinellid
sponges. Further information from Seasearch surveys of the Firth of Lorn (2007-2009)
also identified additional fauna, including Ballan wrasse (Labrus bergylta), cluster
anemones (Parazoanthus spp.), plumose anemones (Metridium senile), red sea fingers
(Alcyonium glomeratum), jewel anemones (Corynactis spp.), lobsters (Homarus
gammarus), cotton spinners (Holothuria forskali), and the anemone Actinothoe
sphyrodeta. Work by Davies (1999) between the Garvellachs and the south-east coast
of Mull, in the Firth of Lorn, found S. pallida on sediment covered stony areas in
association with hydroid turfs, sponges and dense aggregations of the northern feather
star Leptometra celtica in deep (>100m) sediment troughs. At other areas within the
Firth of Lorn, such as Ardnoe Point to the south of Loch Crinan, C. smithii and S.
pallida have been found associated with the colonial ascidian Diazona violacea, the
sponges Axinella infundibuliformis and Mycale lingua and the nationally rare sea fan
anemone Amphianthus dohrnii (Wilding et al., 2005).
4
Species diversity
No information currently available.
Key and characterising species
These have been taken from JNCC’s biotope descriptions (Connor et al., 2004).
Biotype type
Key species for identification
Additional characterising species
CR.MCR.EcCr.CarSwi
C. smithii and S. pallida
C. celata, Halecium halecinum, N.
antennina, N. ramosa, A. digitatum,
A. glomeratum, P. triqueter, M.
rugosa, P. compressa, P. trispinosa,
Antedon bifida, A. petasus, L. ciliaris,
A.
rubens,
M.
glacialis,
E.
esculentus, C. lepadiformis, C.
intestinalis,
A.
mentula,
Corallinaceae
CR.MCR.EcCr.CarSwi.LgAs
C. smithii and S. pallida with
large ascidians such as A.
mentula, C. lepadiformis, C.
intestinalis and Diazona violacea
C. celata, N. antennina, N. ramosa,
A. digitatum, P. triqueter, M. rugosa,
G. cineraria, P. compressa, P.
trispinosa, A. bifida, A. petasus, L.
ciliaris, A. rubens, M. glacialis, E.
esculentus,
A.
virginea,
Corallinaceae
CR.HCR.XFa.SwiLgAs
S. pallida and C. smithii along
with a diverse range of ascidians
including Clavelina lepadiformis,
Ascidia
mentula,
Polycarpa
pomaria, Diazona violacea and
Corella parallelogramma.
A. infundibuliformis, L. nigricans, H.
urceolus, H. halecinum, A. abietina,
H. catharina, N. antennina, N.
ramosa, P. frutescens, A. tubulifera,
A. digitatum, P. triqueter, Antedon
bifida
Coherence
Typicalness
S. pallida is typically found in relatively sheltered deep water (>20m), on flat or sloping
bedrock with little wave action but sufficient current to provide an adequate supply of
food to this sessile filter feeder. These conditions are frequently found off the east
coasts of the southern Western Isles, at the entrances to some mainland sea lochs,
and on deeper submerged reefs in the Minches (Connor et al., 2004).
In Scottish waters the biotope CR.MCR.EcCr.CarSwi and its sub-biotope are generally
found on the upper and vertical faces of very exposed, through to wave-sheltered,
circalittoral bedrock and boulders on silty sediment, which are subject to weak tidal
streams at depths of 10-50m. The biotope CR.HCR.XFa.SwiLgAs differs in that it is
found in areas of stronger currents and generally within the depth range 4-37m.
Despite the depths given above as typical values for these biotopes there is growing
evidence to suggest that they can be found at greater depths in Scottish waters.
Davies (1999) records the old biotope MCR.ErSSwi (Connor et al., 1997; now divided
into CR.HCR.XFa.SwiLgAs and CR.MCR.EcCr.CarSwi) in deep water (>100m)
between Mull and the Garvellachs in the Firth of Lorn. Likewise Malthus et al. (2006)
found the biotope CR.HCR.XFa.SwiLgAs at several sites within the eastern part of the
Sound of Harris at a depth >40m, even recording one site at a depth of 75.1m.
5
Northern sea fan communities are very diverse, and are characteristic of moderately
exposed reefs on the Scottish west coast (Howson et al., 2009). The circalittoral
boulders and bedrock of CR.MCR.EcCr.CarSwi may be home to dense aggregations of
the cup coral C. smithii with the sea fan S. pallida. Alongside these, soft corals,
crinoids, ascidians, hydroids, sponges, and starfish can often be found. The biotope
may appear grazed due to the activity of the urchin E. esculentus. Echinoderms are
especially prevalent in the sub-biotope CR.MCR.EcCrCarSwi.LgAs which has a lower
diversity of some species but greater numbers of large sea squirts and supports the
white cluster anemone P. anguicomus (Connor et al., 2004). Aside from S. pallida, C.
smithii and abundant ascidians the more current swept CR.HCR.XFa.SwiLgAs shelters
a diverse range of species from many taxa; e.g. hydroids, polychaetes, encrusting
algae, echinoderms, bryozoans and sponges. In the Firth of Lorn, northern sea fan
communities occur in various isolated locations. In areas of vertical rock they are
bordered by kelp communities in the shallower waters above and deep inshore
burrowed mud with sea pens on the muddy plains below (Seasearch Firth of Lorn sea
fan surveys, 2007-2009).
Ecological variations across Scottish waters
Northern sea fan communities are found from the lower-infralittoral to depths beyond
the photic zone, and as a result clear changes in the associated algal diversity and
distribution can occur. Furthermore, the penetration of light through the water column is
dependent on water clarity, thus differences in the turbidity of sea water will result in
variations of algal presence/coverage between locations, independent of depth. As
recent records have shown that northern sea fan communities can occur far beyond
their typical depth range (Davies, 1999; Malthus et al., 2006) it is thought that depth
can greatly influence the faunal composition. Differences between the faunal
communities found by the Firth of Lorn Seasearch surveys (2005, 2007-2009) and the
deepwater community recorded near the Garvellachs (Davies, 1999) are probably
attributable to depth. Davies (1999) mentions that the sea bed was dominated by
hydroids, bryozoans, large numbers of sponges and, most notably, dense aggregations
of L. celtica. In contrast the Seasearch surveys (2005, 2007-2009) describe a more
heterogeneous community (including the European spiny lobster Palinurus elephas
and the rare sea fan anemone Amphianthus dohrnii) despite L. celtica not being
recorded. The northern feather star (L. celtica) is listed as an associated species of
northern sea fan communities but never in such abundance as recorded by Davies
(1999). L. celtica is a deepwater species therefore it is not surprising that it is found in
such densities in the deeper localities, however, it does highlight how the northern sea
fan community can vary.
Viability
Although no work specific to the viability of northern sea fan communities has been
carried out, a review by Hill et al. (2010) was carried out on S. pallida. The work
indicated large gaps in the biology and ecology of S. pallida, however, they suggest
that the very limited and site-specific distribution of S. pallida indicates that site-based
conservation measures would be appropriate. It is generally considered that
populations of S. pallida are self-sustaining due to their short-lived larvae and limited
potential for larval dispersal (Hill et al., 2010).
6
A large number of species within northern sea fan communities are sessile encrusting
fauna and so the area needed to protect their home ranges is small. Many sponges
reproduce asexually by fragmentation and budding, so some local recruitment would
be protected by a small area; however, sexual reproduction also occurs, often
seasonally, e.g. most shallow water sponges produce a planktonic larvae. The
dispersal duration for sponge larvae ranges from a few hours to a few days with large
dispersal distance. The reproductive strategies of the other species vary widely, e.g.
within the hydroids, individual species can be either free spawning with external
fertilisation or brooders and produce both planktotrophic and lecithotrophic larvae.
Asexual vegetative reproduction is also known in some hydroids, although the overall
dispersal potential of hydroids is thought to be small (Hill et al., 2010). The ascidians
found within northern sea fan communities also have a relatively short dispersal
potential, e.g. C. lepadiformis is a free spawning species which produces planktonic
lecithotrophic larvae which settle within a few hours (Hill et al., 2010). According to Hill
et al. (2010) an area larger than any recorded examples of northern sea fan
communities in Scottish waters, would be needed to incorporate the complete life
cycle of the sponges and many of the other species present. In light of the uncertainty
around reproduction, viable area size and dispersal of S. palida, it is recommended that
a precautionary approach should be taken to protect northern sea fan communities (Hill
et al., 2010). Therefore it is recommended that the entire, known extent of particular
examples are protected. The boundary of any MPA should also be drawn so as to
encompass not only the known distribution of northern sea fan communities in an area,
but also adjacent areas that support associated or overlapping biotopes.
Longevity
No information was found directly on the longevity of northern sea fan communities.
They are dominated by large, slow growing species such as branching sponges and
sea fans for example, the characteristic species C. smithii and S. pallida have
estimated life spans of approximately 11-20 years (Gregory, 2008; Wilding & Wilson,
2009), and hence are considered relatively stable..
Fragmentation
There is little information relating to typical levels of fragmentation for northern sea fan
communities; however, populations of S. pallida are thought to be self-sustaining
despite their limited and site specific distribution (Hiscock et al., 2001). The reliance of
S. pallida on a suitable substratum for attachment (i.e. bedrock and boulders) is
possibly the major contributor to fragmentation.
7
Indicators of Least Damaged/More Natural
Up to date information on the sensitivity of Northern sea fan communties to pressures
associated with human activities are included, alongside other components, in the
Northern sea fan and sponge communities search feature assessment in the Feature
Activity Sensitivity Tool (FeAST; Marine Scotland, 2013). Below, information on
indicators of naturalness and damage are taken from MarLIN sensitivity data on key
species - namely S. pallida (Wilding & Wilson, 2009) and C. smithii (Gregory, 2008).
Table 1. Indicators of damage and naturalness
Indicators of Naturalness
Indicators of Change/Damage
Potential Sources of Damage
Community mainly comprising of
sessile animals with little algal
cover.
Elevated levels of epiphytic algal
cover.
Eutrophication
Low levels of silt present. Filter
feeds un-smothered.
Elevated levels of
smothering of filer
organisms.
siltation,
feeding
Siltation / smothering
Intact fragile species (such as
sea fans and corals). Few bare
patches on rocks.
Evidence of damaged or broken
sea fans (including detached
broken sections). Bare patches
on rock where sessile animals
such as anemones have been
detached.
Increases in abundance of
southern species and declines in
northern species e.g. increase in
abundance C. smithii and a
decline in the abundance of S.
pallida, which is at its southern
limit in Scotland.
Physical disturbance.
Community
composition
comprising of typical species, see
biological diversity section. Few
‘southern species’ present.
Increase in ambient water
temperature possibly as a result
of climate change.
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
Very little evidence was available on the potential of northern sea fan communities to
recover from elevated levels of epiphytic algal cover, elevated levels of siltation and
physical damage. However, the presence of large, slow growing organisms may mean
that full recovery from damage or loss of such individuals could take many years. Some
sensitivity data are available for S. pallida specifically.
Physical damage
Based on findings from a related species (Eunicella verrucosa) MarLIN lists S. pallida
as having a moderate ability to recover from abrasion and physical disturbance
(Hiscock et al., 2004). Some sea fan colonies returned to an upright position
immediately after impact from lobster pots, while others were permanently bent, which
would reduce feeing efficiency (Eno et al., 1996). Tinsley (2006) observed flattened sea
fans which had continued growing, with new growth being aligned perpendicular to the
current.
8
Siltation
Swiftia pallida is found on rocks covered with a fine layer of silt (Mitchell et al., 1983).
While siltation may inhibit feeding, colonies of the sea fan Eunicella verrucosa produce
mucus to clear themselves of silt (Hiscock, pers. comm.). It is, however, thought that
smothering causes mortality (Hiscock et al., 2004). Therefore MarLIN lists S. pallida as
having a low ability to recover from smothering, but tolerant of increases in suspended
sediment (Hiscock et al., 2004).
Geographical Variation
The majority of British records of the species S. pallida and the biotopes listed within
northern sea fan communities are located in Scotland. These are all recorded along the
west coast and the Outer Hebrides. In the last decade the majority of .CarSwi and
.SwiLgAs records are from the Firth of Lorn (Seasearch survey, 2005) and Loch Sunart
(Dipper & Johnston, 2000). There are also recent records off the east coast of North
Uist for .CarSwi and .SwiLgAs and a single record for .CarSwi off the northern coast of
Skye (Covey et al., 1998; Thorpe et al., 1999). Along the east coast of the Outer
Hebrides are a number of records from more than 20 years ago for .CarSwi. Around
this period there are also many records for .SwiLgAs from Mull, Loch Sunart and the
Firth of Lorn, with individual records from the Ardnamurchan peninsula and the east
coast of North Uist; and more recent records in the Sound of Harris (Malthus et al.,
2006)
Geographical context
Little information on the distribution of northern sea fan communities was found from
outside the UK. Research into the distribution of the characterising species reveals that
C. smithii is present around south-west Europe and the Mediterranean and also found
in Denmark (Hayward & Ryland, 1995; Tendal & Nielsen, 1997). S. pallida occurs on
the west coasts of Norway and Sweden, in south-west Ireland (the Kenmare River, the
only place where its distribution overlaps the pink sea fan Eunicella verrucosa) and in
deep water from the Bay of Biscay, the Mediterranean, Madeira and Morocco (Wilding
& Wilson, 2009).
9
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