Caryophyllia (Caryophyllia) smithii, sponges and crustose communities on wave-exposed circalittoral rock

Distribution Map

Map Key

  • Orange points: Core Records
  • Pale Blue points: Non-core, certain determination
  • Black points: Non-core, uncertain determination
  • Yellow areas: Predicted habitat extent

Summary

UK and Ireland classification

Description

This biotope typically occurs on the upper and vertical faces of wave-exposed, moderately strong to weakly tide-swept, circalittoral bedrock or boulders, with a water depth range of 20-30m. This often silty biotope has a typically sparse fauna, appearing grazed, and is characterized by common cup corals Caryophyllia smithii, frequent Alcyonium digitatum and occasional urchins Echinus esculentus. There may be occasional large growths of the sponge Cliona celata, Haliclona viscosa, Pachymatisma johnstonia and the axinellid sponge Stelligera stuposa. Echinoderms form a prominent feature of the fauna within this biotope, with species such as Marthasterias glacialis, Asterias rubens, Luidia ciliaris, Henricia oculata, Holothuria forskali, Antedon bifida and Aslia lefevrei present. Bryozoan crusts such as Parasmittina trispinosa and encrusting red algae cover the rock/boulder surface. The bryozoan Porella compressa may also be recorded occasionally. Isolated clumps of hydroids feature species such as Nemertesia antennina, Nemertesia ramosa, Abietinaria abietina, Halecium halecinum and Sertularella gayi. Other species observed include the anemone Corynactis viridis, Urticina felina, Cylista elegans, Calliostoma zizyphinum, Balanus crenatus and Spirobranchus triqueter. Two variants within this biotope have been distinguished: CarSp.PenPcom and CarSp.Bri. While CarSp.PenPcom tends to have the bryozoans Pentapora foliacea and Porella compressa, while CarSp.Bri features a dynamic community of brittlestars covering the seabed in a dense mat. Ophiothrix fragilis is usually the dominant species in shallow water but tends to be replaced by Ophiocomina nigra in deeper water. (Information from Connor et al., 2004; JNCC, 2015).

Depth range

20-30 m, 30-50 m

Additional information

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Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

CR.MCR.EcCr.CarSp, This often silty biotope has a typically sparse fauna, appearing grazed, and is characterized by common cup corals Caryophyllia smithii, frequent Alcyonium digitatum and occasional urchins Echinus esculentus. There may be occasional large growths of the sponges; Cliona celata, Haliclona viscosa, Pachymatisma johnstonia and the axinellid sponge Stelligera stuposa (Connor et al., 2004).

For this sensitivity assessment Alcyonium digitatum, Caryophyllia smithii, Echinus esculentus are primary research focus. Sponges are also an important characterizing component of CR.MCR.EcCr.CarSp. Cliona celata is the most abundant sponge species within CR.MCR.EcCr.CarSp (Connor et al., 2004) and has therefore also been identified as another primary focus of research, however it is important to note there are other important sponge species within CR.MCR.EcCr.CarSp (e.g. Haliclona viscosa, Pachymatisma johnstonia and the axinellid sponge Stelligera stuposa) however they are not subject to specific research.

Resilience and recovery rates of habitat

Alcyonium digitatum is a colonial species of soft coral with a wide distribution in the North Atlantic, recorded from Portugal (41°N) to Northern Norway (70°N) as well as on the east coast of North America (Hartnoll, 1975; Budd, 2008). Colonies consist of stout “finger like” projections (Hartnoll, 1975) which can reach up to 20 cm tall (Budd, 2008) and can dominate circalittoral rock habitats (as in CR.HCR.FaT.CTub.Adig; Connor et al., 2004). Alcyonium digitatum colonies are likely to have a lifespan which exceeds 20 years as colonies have been followed for 28 years in marked plots (Lundälv, pers. comm., in Hartnoll, 1998). Those colonies which are 10-15 cm in height have been aged at between 5 and 10 years old (Hartnoll, unpublished). Most colonies are unisexual, with the majority of individuals being female.  Sexual maturity is predicted to occur, at it’s earliest, when the colony reaches its second year of growth, however the majority of colonies are not predicted to reach maturity until their third year (Hartnoll, 1975).

Alcyonium digitatum spawns from December and January. Gametes are released into the water where fertilization occurs. The embryos are neutrally buoyant and float freely for 7 days, when they give rise to actively swimming lecithotrophic planulae which may have an extended pelagic life before they eventually settle (usually within 1 or 2 further days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975; Budd, 2008). In laboratory experiments, several larvae of Alcyonium digitatum failed to settle within 10 days, presumably finding the conditions unsuitable. These larvae were able to survive 35 weeks as non-feeding planulae. After 14 weeks some were still swimming and after 24 weeks the surface ciliation was still active although they rested on the bottom of the tanks. By the end of the experiment, at 35 weeks the larvae had shrunk to a diameter of 0.3 mm. This ability to survive for long periods in the plankton may favour the dispersal and eventual discovery of a site suitable for settlement (Hartnoll, 1975). The combination of spawning in winter and the long pelagic lifespan may allow a considerable length of time for the planulae to disperse, settle and metamorphose ahead of the spring plankton bloom. Young Alcyonium digitatum will consequently be able to take advantage of an abundant food resource in spring and be well developed before the appearance of other organisms that may otherwise compete for the same substrata. In addition, because the planulae do not feed whilst in the pelagic zone they do not suffer by being released at the time of minimum plankton density. They may also benefit by the scarcity of predatory zooplankton which would otherwise feed upon them (Hartnoll, 1975).

Spirobranchus triqueter and Parasmittina trispinosa are two visually dominant encrusting species within CR.MCR.EcCr.CarSp. Spirobranchus triqueter is a species of serpulid worm which forms encrusting tubes, typically 2-3cm long, on rock and shell surfaces. Once settled onto the substratum the worm forms a temporary delicate semi-transparent tube. Mature tubes are formed by a secretion of calcium carbonate (obtained from seawater). Growth rate has been observed by Dons (1927) to be 1.5 mm per month, although this varies with external conditions. Hayward & Ryland (1995) and Dons (1927) stated that sexual maturity is reached in approximately 4 months. Spirobranchus triqueter is also a visually dominant species within mobile and/or disturbed biotopes e.g. SS.SCS.CCS.SpiB (Connor et al., 2004), indicating this species is either highly resilient to physical disturbance or has a rapid recolonization rate. Hayward & Ryland (1995) noted that individuals lived approximately 1.5 years (Hayward & Ryland, 1995). Spirobranchus triqueter are broadcast spawners and are therefore likely to have large dispersal capacity. Larvae are pelagic for about 2-3 weeks in the summer, however, in the winter this amount of time increases to about 2 months (Hayward & Ryland, 1995). The time of reproduction is variable, Hayward & Ryland (1995) and Segrove (1941) suggested that Spirobranchus triqueter reproduction probably takes place throughout the year, but, peaks in spring and summer. However, Moore (1937) noted Spirobranchus triqueter breeding only took place in April in Port Erin, Isle of Man. Castric-Fey (1983) studied variations in settlement rate and concluded that, although the species settled all year round, very rare settlement was observed during winter and maximum settlement occurred in April, June, August and Sept-Oct. Studies in Bantry Bay  revealed a single peak in recruitment during summer (especially July and August) with very little recruitment at other times of the year (Cotter et al., 2003).

Caryophyllia smithii is a small (max 3cm across) solitary coral common within tide swept sites of the UK (Wood, 2005), distributed from Greece (Koukouras, 2010) to the Shetland Islands and southern Norway (NBN, 2015). It was suggested by Fowler & Laffoley (1993) that Caryophyllia smithii was a slow growing species (0.5-1mm in horizontal dimension of the corallum per year) which in turn suggests that inter-specific spatial competition with colonial faunal or algae species are important factors in determining local abundance of Caryophyllia smithii (Bell & Turner, 2000). Caryophyllia smithii reproduces sexually; sessile polyps discharge gametes typically from January-April, gamete release is most likely triggered by seasonal temperature increases, gametes are fertilised in the water column and develop into a swimming planula, which then settles onto the suitable substrata. The pelagic stage of the larvae may last up to 10 weeks, which provides this species with a good dispersal capability (Tranter et al., 1982).

Cliona celata is considered a hardy sponge, tolerant of environmental stressors such as high nutrient loads, low salinity, and large temperature variation (Duckworth & Peters, 2013). Cliona celata is a physically distinctive species of sponge that can bore into soft rock (e.g. limestone) or in hard rock areas has a massive form (Wood, 2007). The boring form is recognizable as yellow papillae sticking out of limestone (calcareous rock, mollusc shells). The massive form has raised, rounded ridges up to 40 cm across. Large oscules with raised rims are found along the tops of the ridges. It often forms a thick plate-like structure standing on its edge with large specimens growing up to 1 m across and 50 cm high (Snowden, 2007).  According to the World Porifera database, Cliona celata has a relatively cosmopolitan distribution from north of Shetland to the cape of good hope, South Africa, as well as being recorded throughout the Mediterranean (Van Soest, 2001 in Costello et al., 2001). At the time of writing no specific information for Cliona celata longevity was found, however, in general, sponges have a relatively long lifespan (e.g. Ayling (1983) estimated sponge patches in New Zealand were over 70 years old) (Marine institure report). Piscitelli et al. (2011) observed an annual peak in reproductive activity in April-May from individuals in the Mediterranean, and suggested this was a result of a sharp seasonal increase in water temperature. However, Carver et al. (2010) suggested Cliona celata specimens from New Brunswick, Canada spawned from June-July.  Recruitment can occur via larval settlement as well as through transfer/contact (i.e. sponge colonies can spread to new/virgin substrates if they come in contact with existing colonies) (Duckworth & Peterson, 2013). Warburton (1966) documented the spawning of Cliona celata under laboratory conditions, and reported the production of motile larvae that settled after 2 days (Carver et al., 2010). Information concerning colonization rates are scare however, tropical clionid sponges (the same taxonomic family as Cliona celata) can colonize dead coral within “a few weeks” and live coral within 2-3 months (Schönberg & Wilkinson, 2001). Furthermore, the short larval period (2 days) plus observation from Carver et al. (2010) indicate Cliona celata can colonize virgin surfaces within a year. Cliona celata is a pest species in scallop aquaculture, Carver et al. (2010) demonstrated that contact between shells colonized with Cliona celata and those that were not colonized results in rapid spread in Cliona celata throughout scallop farms. Once settled Cliona celata colonies have a rapid growth rate of up to 15cm2/yr (Carver et al., 2010).

Whomersley & Picken (2003) documented epifauna colonization of offshore oil platforms in the North Sea from 1989-2000. On all platforms Mytilus edulis dominated the near surface community. For the first 3 years, hydroids and tubeworms dominated the community below the mussel band. However the hydroid community were later out-competed by other more climax communities. Recruitment of Alcyonium digitatum and Metridium senile began at 2-5 years (dependent on the oil rig). The community structure and zonation differed between the 4 rigs, however generally after4  years Metrdium senile had become the dominant organism below the mussel zone to approximately 60-80 m Below Sea Level (BSL). Zonation differed between oil rigs however, from approximately 60-90 m BSL Alcyonium digitatum was the dominant organism.

The Scylla was intentionally sunk on the 27th March 2004 in Whitsand Bay, Cornwall to act as an artificial reef. Hiscock et al. (2010) recorded the succession of the biological community on the wreck for 5 years following the sinking of the ship. Initially the wreck was colonized by opportunistic species /taxa; filamentous algae, hydroids, serpulid worms and barnacles. Tubularia sp. were early colonizers, appearing within a couple of months after the vessel was sunk. Metridium senile appeared late in the summer of the first year, but didn’t become visually dominant until 2007 (3 years after the vessel was sunk). Sagartia elegens was recorded within the summer of 2005, and by the end of 2006 was well established. Corynactis viridis was first recorded in the summer of the first year and quickly formed colonies via asexual reproduction. Urticina felina was first recorded at the end of august 2006 (2 years after the vessel was sunk), and by summer 2008 had increased in abundance. Alcyonium digitatum was first recorded in early summer 2005, a year after the vessel was sunk. Within 1 year of growth colonies had grown to nearly full size, however, did not become a visually dominant component of the community until 2009 (5 years after the vessel had been sunk). The authors noted that erect branching Bryozoa (such as Securiflustra securifrons) are not a common part of rocky reef communities to the west of Plymouth and at the time of writing had not colonized to any great extent on ‘Scylla’ by the end of the study, although several species were recorded which included Chartella papyracea in 28/08/2006 (2 years after the vessel was sunk). Caryophyllia smithii was noted to colinize the wreck a year after the vessel was sunk (07/09/2005).

Parasmittina trispinosa is an encrusting bryozoan which is described as having a “cosmopolitan” distribution by Powell (1971), in the North East Atlantic recorded from all coasts of the British Isles (NBN, 2015) to the Iberian Peninsula (Ramos, 2010). Parasmittina trispinosa is also recorded from the Panama Cana (Powell (1971) to the Gulf of Alaska (Soule, 2002) in the Pacific ocean. At the time of writing sparse information regarding the life history traits of Parasmittina trispinosa. Eggleston (1972) noted In the Isle of Man, a peak in reproductive and vegetative growth was not well marked in Parasmittina trispinosa, and the number of embryos present is fairly constant throughout the year (Eggleston, 1972). Indicating that Parasmittina trispinosa could potentially reproduces annually within the UK. However, due to the lack of available literature regarding Parasmittina trispinosa it’s resilience cannot be assessed with sufficient confidence in this review.

Echinus esculentus is a sea urchin found within Northeast Atlantic, recorded from Murmansk Coast, Russia to Portugal (Hansson, 1998). Echinus esculentus is an important algal grazer and is thought, combined with low light levels, to control red algal growth within CR.MCR.EcCr.FaAlCr.Adig, CR.MCR.EcCr.FaAlCr.Sec & CR.MCR.EcCr.FaAlCr.Spi (Connor et al., 2004). Echinus esculentus is estimated to have a lifespan of 8-16 years (Nichols, 1979; Gage, 1992) and reach sexual maturity within 1-3 years (Tyler-Walters, 2008). Maximum spawning occurs in spring although individuals may spawn over a protracted period throughout the year. Gonad weight is at it’s maximum in February/March in English Channel (Comely & Ansell, 1989) but decreases during spawning in spring and then increases again through summer and winter until the next spawning season. Spawning occurs just before the seasonal rise in temperature in temperate zones but is probably not triggered by rising temperature (Bishop, 1985). Echinus esculentus is a broadcast spawner, with a complex larval life history which includes a blastula, gastrula and a characteristic 4 armed echinopluteus stage that forms an important component of the zooplankton. MacBride (1914) observed planktonic larval development could take 45-60 days in captivity. Recruitment is sporadic or variable depending on locality, e.g. Millport populations showed annual recruitment, whereas few recruits were found in Plymouth populations during Nichols studies between 1980-1981 (Nichols, 1984). Bishop & Earll (1984) suggested that the population of Echinus esculentus at St Abbs had a high density and recruited regularly whereas the Skomer population was sparse, ageing and had probably not successfully recruited larvae in the previous 6 years (Bishop & Earll, 1984). Comely & Ansell (1988) noted that the largest number of Echinus esculentus occurred below the kelp forest.

Echinus esculentus is a mobile species (Tyler-Walters, 2008) and could therefore migrate and re-populate an area quickly if removed. For example, Lewis & Nichols (1979) found that adults were able to colonize an artificial reef in small numbers within 3 months and the population steadily grew over the following year. If completely removed from a site and local populations are naturally sparse then recruitment may be dependent on larval supply which can be highly variable. As suggested by Bishop & Earll (1984) the Skomer, Wales Echinus esculentus population had most likely not successfully recruited for 6 years which would suggest the mature population would be highly sensitive to removal and may not return for several years. On 19th November 2002 the Prestige oil tanker spilled 63 000t of fuel 130 nautical miles off Galicia, Spain. High wave exposure and strong weather systems increased mixing of the oil to “some” depth within the water column, causing sensitive faunal communities to be effected. Preceding and for nine years following the oil spill, the biological community of Guéthary, France was monitored. Following the oil spill taxonomic richness decreased significantly from 57 recorded species to 41, which included the loss of Echinus esculentus from the site. 2-3 years after the oil spill taxonomic richness had increased to pre-spill levels and Echinus esculentus had returned (Castège et al., 2014).

Resilience assessment. Spirobranchus triqueter can reportedly reach maturity within approximately 4 months and is often a dominant component of physically disturbed habitats, indicating rapid colonization rates (<1 year) and/or physical robustness. Echinus esculentus can reportedly reach sexual maturity within 1-2 years (Tyler-Walters, 2008), however as highlighted by Bishop & Earll (1984) and Castège et al., (2014) recovery may take 2-6 years (possibly more if local recruitment is poor).  Alcyonium digitatum can recruit onto bare surfaces within 2 years, however may take up to 5 years to become a dominant component of the community (Whomersley & Picken, 2003; Hiscock et al., 2010). Caryophyllia smithii colonized the wreck of the Scylla within a year, however this may be due to the time of the vessel sinking and if removed recovery may take up to 2 years. Cliona celata is likely to recover within a year. If the community is completely removed from the habitat (resistance of none or low) resilience has been assessed as ‘Medium’, however if resistance has been assessed as medium or high then resilience will be assessed as ‘High’.

Hydrological Pressures

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ResistanceResilienceSensitivity
Temperature increase (local) [Show more]

Temperature increase (local)

Benchmark. A 5°C increase in temperature for one month, or 2°C for one year (Temperature change pressure definition).

Evidence

CR.MCR.EcCr.CarSp is distributed across the west coast of Scotland and south west coast of Ireland. Sea surface temperature across this distribution ranges from northern to southern Sea Surface Temperature (SST) of 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013). Alcyonium digitatum is described as a northern species by Hiscock et al. (2004), but is distributed from Northern Norway (70°N) to Portugal (41°N) (Hartnoll, 1975; Budd, 2008). Spirobranchus triqueter is recorded as abundant in sub-tidal habitats of Trondheimsfjord (63°N) (Kukliński & Barnes, 2008), no survey reports could be found further north. The most southerly records are from the Iberian peninsula, Spain (Ramos, 2010) as well from the Alexandria coast of Egypt, Mediterranean Sea (Sarah, 2010). Across this latitudinal gradient Spirobranchus triqueter is likely to experience a range of temperatures from approximately 5-28°C (Seatemperature, 2015), and is therefore unlikely to be affected at the pressure benchmark. Bishop (1985) suggested that Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate and resultant metabolic stress. Ursin (1960) reported Echinus esculentus occurred at temperatures between 0-18°C in Limfjord, Denmark. Bishop (1985) noted that gametogenesis occurred at 11-19°C however, continued exposure to 19°C disrupted gametogenesis. Embryos and larvae developed abnormally after 24hr exposure to 15°C but normally at 4, 7 and 11°C (Tyler & Young 1998). Mature examples of Caryophyllia smithii are recorded in Greece (Koukouras, 2010), and are therefore unlikely to be physically affected at the benchmark. However, Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns.

Seawater temperature is positively correlated to sponge growth (Duckworth & Peters, 2013). Duckworth & Peters (2013) demonstrated that an increase in water temperature to 26 and 31°C had no detectable effect on boring activity of Cliona celata. Whereas as cold winter temperatures (<5°C) can cause boring activity to cease, the incurrent papillae to withdraw and the excurrent papillae constrict (Fell et al., 1984; Carver et al., 2010). Indicating cold temperatures are more limiting to Cliona celata than hot. Piscitelli et al. (2011) observed an annual peak in reproductive activity in April-May from individuals in the Mediterranean, and suggested this was a result of a sharp seasonal increase in water temperature. Which suggests variable temperatures could affect larval recruitment processes, but otherwise not otherwise negatively affect Cliona celata.

Sensitivity assessment. Resistance has been assessed as ‘Medium’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Low’.

Medium
High
High
High
Help		High
High
High
High
Help		Low
High
High
High
Help
Temperature decrease (local) [Show more]

Temperature decrease (local)

Benchmark. A 5°C decrease in temperature for one month, or 2°C for one year (Temperature change pressure definition).

Evidence

CR.MCR.EcCr.CarSp is distributed across the west coast of Scotland and south west coast of Ireland. Sea surface temperature across this distribution ranges from northern to southern Sea Surface Temperature (SST) of 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013). Alcyonium digitatum is described as a northern species by Hiscock et al. (2004), but is distributed from Northern Norway (70°N) to Portugal (41°N) (Hartnoll, 1975; Budd, 2008). Across this latitudinal gradient both species are likely to experience a range of temperatures from approximately 5-18°C. Alcyonium digitatum was also reported to be apparently unaffected by the severe winter of 1962-1963 where air temperature reached -5.8°C (Crisp, 1964). Echinus esculentus has been recorded from the Murmansk Coast, Russia. Due to the high latitude at which Echinus esculentus can occur it is unlikely to be affected at the pressure benchmark. 

Spirobranchus triqueter is described as a temperate species by Kupriyanova & Badyaev (1998). Spirobranchus triqueter is recorded as abundant in sub-tidal habitats of Trondheimsfjord (63°N) (Kukliński & Barnes, 2008), no survey reports could be found further north. Averaged across several years the lowest winter temperature within Trondheimsfjord is 4.9°C (Seatemperature, 2015). Below 7°C Spirobranchus triqueter is unable to build calcareous tubes (Thomas, 1940). Mature adults may survive a decrease at the pressure benchmark however larvae may not be able to attach to the substate (Riley & Ballerstedt, 2005) if a temperature decrease co-occurred with cold winter temperatures in the UK. However, settlement is reportedly low within winter (see resilience section) and, therefore, the effects on recruitment are likely to be minor. However, Caryophyllia smithii has a northern range limit in the Shetland isles and Southern Norway (NBN, 2015) and may, therefore, be negatively affected by cold temperatures in Northern examples of this biotope.

Seawater temperature is positively correlated to sponge growth (Duckworth & Peters, 2013). Duckworth & Peters (2013) demonstrated that an increase in water temperature to 26 and 31°C had no detectable effect on boring activity. Whereas as cold winter temperatures (<5°C) can cause boring activity to cease, the incurrent papillae to withdraw and the excurrent papillae constrict (Fell et al., 1984; Carver et al., 2010). Indicating cold temperatures are more limiting to Cliona celata than hot. Piscitelli et al. (2011) observed an annual peak in reproductive activity in April-May from individuals in the Mediterranean, and suggested this was a result of a sharp seasonal increase in water temperature. Which suggests variable temperatures could affect larval recruitment processes, but not otherwise affect Cliona celata negatively.

Sensitivity assessment. Alcyonium digitatum, Echinus esculentus have northern/boreal distributions and are unlikely to be affected at the benchmark level. Spirobranchus triqueter is unable to build calcareous tubes at low temperatures, however, during winter, this is unlikely to have any significant effects on recruitment. In addition, the depth of the biotope probably protects it from short-term acute decreases in temperature. The important characterizing Caryophyllia smithii is close to its northern distribution limit within the British Isles and a decrease at the benchmark level may result in some mortality in northern examples of the biotope.  Therefore, resistance is therefore ‘Medium’, resilience is ‘High’ and sensitivity is ‘Low’.

Medium
Low
NR
NR
Help		High
High
High
High
Help		Low
Low
Low
Low
Help
Salinity increase (local) [Show more]

Salinity increase (local)

Benchmark. An increase in one MNCR salinity category above the usual range of the biotope or habitat (Salinity regime change pressure definition).

Evidence

Lyster (1965) tested the tolerance of Spirobranchus triqueter larvae to various hyper and hypo salinity treatments. Larvae were placed in cultures ranging from 0-90‰ and notes were made on the time taken for larvae to die or begin displaying abnormal behaviour. Spirobranchus triqueter larvae were tolerant of salinities ranging from 20-50‰, above 50‰ caused high mortality. Spirobranchus triqueter is therefore unlikely to be affected at the pressure benchmark.

Echinoderms are generally stenohaline and possess no osmoregulatory organ (Boolootian, 1966). Therefore an increase in salinity may cause Echinus esculentus mortality. Alcyonium digitatum’ distribution and the depth at which it occurs also suggest it would not likely experience regular salinity fluctuations and therefore tolerate significant increases in salinity.

CR.MCR.EcCr.CarSp is restricted to full salinity (Connor et al., 2004), it therefore seems likely that an increase in salinity to >40‰ may cause a decline in the abundance of characterizing species. Furthermore a reduction in Echinus esculentus may cause an increase in red algae growth, which would change the character of the biotope.

Sensitivity assessment. Resistance has been assessed as ‘Low’, resilience as ‘Medium’. Sensitivity has been assessed as ‘Medium’.

Low
Low
NR
NR
Help		Medium
High
High
High
Help		Medium
Low
Low
Low
Help
Salinity decrease (local) [Show more]

Salinity decrease (local)

Benchmark. A decrease in one MNCR salinity category above the usual range of the biotope or habitat (Salinity regime change pressure definition detail).

Evidence

Alcyonium digitatum does inhabit situations such as the entrances to sea lochs (Budd, 2008) or the entrances to estuaries (Braber & Borghouts, 1977) where salinity may vary occasionally. Furthermore as highlighted the Marine Nature Conservation Review (MNCR) records of 23rd Oct 2014 show Alcyonium digitatum is found within a number of variable salinity biotopes, e.g. MCR.BYH.Flu.Hocu,. However, its distribution and the depth at which it occurs suggest that Alcyonium digitatum would not likely often experience salinity fluctuations and therefore unlikely to survive significant reductions in salinity (Budd, 2008).

Echinoderms are generally unable to tolerate low salinity (stenohaline) and possess no osmoregulatory organ (Boolootian, 1966). At low salinity urchins gain weight, and the epidermis loses its pigment as patches are destroyed; prolonged exposure is fatal. However, within Echinus esculentus there is some evidence to suggest intracellular regulation of osmotic pressure due to increased amino acid concentrations. Furthermore as highlighted the Marine Nature Conservation Review (MNCR) records of 23rd Oct 2014 show Echinus esculentus is found within a number of variable and reduced salinity biotopes, e.g. IR.LIR.KVS.SlatPsaVS.

Lyster (1965) tested the tolerance of Spirobranchus triqueter larvae to various hyper and hypo salinity treatments. Larvae were placed in cultures ranging from 0-90‰ and notes were made on the time taken for larvae to die or begin displaying abnormal behaviour. Spirobranchus triqueter larvae can survive very well in salinities down to 20‰, and can tolerate salinities down to 10‰. Adults are tolerant of salinities as low as 3‰, and can be found in areas were salinity ranges from 18-23‰ (Alexander et al., 1935).

Carver et al. (2010) suggested Cliona celata can function efficiently in salinities as low as 20‰, and can withstand exposure to 15‰ for brief periods, but prevailing salinities less than 10–15‰ are likely to be lethal (Hartman 1958, Hopkins 1962).

Sensitivity review. CR.MCR.EcCr.CarSp are recorded exclusively in full marine conditions (30-40 ‰) (Connor et al., 2004). The lack of records within “Reduced” salinity (18-30‰) suggests the community would not persist/be recognisable if salinity was reduced. Spirobranchus triqueter is likely to be able to tolerate reduced salinity, Records from the MNCR suggest Alcyonium digitatum, Caryophyllia smithii & Echinus esculentus can occur in reduced salinity habitats, however the general evidence suggests that these species would decrease in abundance. Resistance has been assessed as ‘Low’, Resilience as ‘Medium’. Sensitivity has been assessed as ‘Medium’.

Low
Low
NR
NR
Help		Medium
High
High
High
Help		Medium
Low
Low
Low
Help
Water flow (tidal current) changes (local) [Show more]

Water flow (tidal current) changes (local)

Benchmark. A change in peak mean spring bed flow velocity of between 0.1 m/s and 0.2 m/s for more than one year (Water flow pressure definition). 

Evidence

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are suspension feeders, relying on water currents to supply food (Hiscock, 1983). These taxa therefore thrive in conditions of vigorous water flow e.g. around Orkney and St Abbs, Scotland, where Alcyonium digitatum dominated biotopes may experience tidal currents of 3 and 4 knots (approximately 1.5 m/sec) during spring tides (Kluijver, 1993). Caryophyllia smithii in particular is described as favouring sites with high tidal flow (Bell & Turner, 2000; Wood, 2005).

Bell (2002) documented Cliona celata regeneration, following artificial damage, at two sites within Lough Hyne, Ireland. The study demonstrated 80% of Cliona celata were fully regenerated at the site which received high water flow (2m/sec) within 100 days, whereas no Cliona celata found in slower water velocities (0.1 m/sec) had fully regenerated. The author suggested an increase in water flow may increase food availability and therefore be of benefit to Cliona celata, whereas slow water flow (0.1 m/sec) may limit food supply and therefore slow regeneration if damaged.

Spirobranchus triqueter has been recorded in areas with very sheltered to exposed water flow rates (Price et al., 1980). Wood (1988) observed Spirobranchus sp. in strong tidal streams and Hiscock (1983) found that in strong tidal streams or strong wave action where abrasion occurs, fast growing species such as Spirobranchus triqueter occur.

Echinus esculentus occurred in kelp beds on the west coast of Scotland in currents of about 0.5 m/sec. Outside the beds specimens were occasionally seen being rolled by the current (Comely & Ansell, 1988), which may have been up to 1.4 m/sec. Urchins are removed from the stipe of kelps by wave and current action. Echinus esculentus are also displaced by storm action. After disturbance Echinus esculentus migrates up the shore, an adaptation to being washed to deeper water by wave action (Lewis & Nichols, 1979). Therefore, increased water flow may remove the population from the affected area; probably to deeper water although individuals would probably not be killed in the process and could recolonize the area quickly.

Sensitivity assessment. CR.MCR.EcCr.CarSp is recorded from moderately strong to negligible tidal currents (<1.5m/sec) (Connor et al., 2004). The abundance of the characterizing species may be affected by a large scale increase/decrease in water flow (e.g. >1m/sec), however a change in tidal velocity of 0.1-0.2 m/s is not likely to have a significant effect. Resistance has been assessed as ‘High’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Not sensitive’.

High
Medium
High
High
Help		High
High
High
High
Help		Not sensitive
Medium
High
High
Help
Emergence regime changes [Show more]

Emergence regime changes

Benchmark.  1) A change in the time covered or not covered by the sea for a period of ≥1 year, or 2) an increase in relative sea level or decrease in high water level for ≥1 year. (Emergence regime change pressure definition).

Evidence

Changes in emergence are 'Not relevant' to CR.MCR.EcCr.FaAlCr.Adig, CR.MCR.EcCr.FaAlCr.Pom & CR.MCR.EcCr.FaAlCr.Sec, which are restricted to fully subtidal/circalittoral conditions-The pressure benchmark is relevant only to littoral and shallow sublittoral fringe biotopes.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Wave exposure changes (local) [Show more]

Wave exposure changes (local)

Benchmark. A change in near shore significant wave height of >3% but <5% for more than one year (Wave action pressure definition). 

Evidence

CR.MCR.EcCr.CarSp is recorded from extremely wave exposed-moderately wave exposed sites (Connor et al., 2004). Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueteri and sponges are suspension feeders relying on water currents to supply food. These taxa therefore thrive in conditions of vigorous water flow.

Echinus esculentus occurred in kelp beds on the west coast of Scotland in currents of about 0.5 m/sec. Outside the beds specimens were occasionally seen being rolled by the current (Comely & Ansell, 1988), which may have been up to 1.4 m/sec. Urchins are removed from the stipe of kelps by wave and current action. Echinus esculentus are also displaced by storm action. After disturbance Echinus esculentus migrates up the shore, an adaptation to being washed to deeper water by wave action (Lewis & Nichols, 1979). Keith Hiscock (pers. comm.) reported Echinus esculentus occurred in significant numbers as shallow as 15m below low water at the extremely wave exposed site of Rockall, Scotland. Therefore, localised increases in wave height may remove the population from the affected area; probably to deeper water although individuals would probably not be killed in the process and could recolonize the area quickly.

Sensitivity assessment. Wave action is a fundamental environmental variable controlling the biological community of sublittoral biotopes. A large and significant change in wave height may fundamentally alter the character of CR.MCR.EcCr.CarSp. However a change in near shore significant wave height of 3-5% is not likely to have a significant effect on the biological community. Resistance has been assessed as ‘High’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Not sensitive’.

High
Low
NR
NR
Help		High
High
High
High
Help		Not sensitive
Low
Low
Low
Help

Chemical Pressures

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ResistanceResilienceSensitivity
Transition elements & organo-metal contamination [Show more]

Transition elements & organo-metal contamination

Benchmark. Exposure of marine species or habitat to one or more relevant Transitional metal or organometal (e.g. TBT) contaminants via uncontrolled releases or incidental spills (Transitional metals and organometals pressure definition). 

Evidence

This pressure is Not assessed but evidence is presented where available.

No information on the direct biological effects of heavy metal contamination on Alcyonium digitatum. Possible sub-lethal effects of exposure to heavy metals, may result in a change in morphology, growth rate or disruption of reproductive cycle. The vulnerability of this species to concentrations of pollutants may also depend on variations in other factors e.g. temperature and salinity conditions outside the normal range.

Based on the available evidence for several species Bryan (1984) suggested that polychaetes are fairly resistant to heavy metals.

Little is known about the effects of heavy metals on echinoderms. Bryan (1984) reported that early work had shown that echinoderm larvae were sensitive to heavy metals contamination, for example Migliaccio et al. (2014) reported exposure of Paracentrotus lividis larvae to increased levels of cadmium and manganese caused abnormal larval development and skeletal malformations. Kinne (1984) reported developmental disturbances in Echinus esculentus exposed to waters containing 25 µg / l of copper (Cu).

At the time of writing no information could be found relating to the sensitivity of Caryophyllia smithii to heavy metal contamination.

Cliona celata is known to be a hardy sponge, tolerant to a number of abiotic stressors (Duckworth & Peters, 2013), however at the time of writing no information could be gathered concerning the effects of heavy metal contamination of Cliona celata.

Not Assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help
Hydrocarbon & PAH contamination [Show more]

Hydrocarbon & PAH contamination

Benchmark. Exposure of marine species or habitat to one or more relevant hydrocarbon or polyaromatic hydrocarbon (PAH) contaminants via uncontrolled releases or incidental spills (Hydrocarbon & PAH pressure definition).

Evidence

This pressure is Not assessed but evidence is presented where available.

CR.MCR.EcCr.CarSp is a sub-tidal biotope (Connor et al., 2004). Oil pollution is mainly a surface phenomenon its impact upon circalittoral turf communities is likely to be limited. However, as in the case of the Prestige oil spill off the coast of France, high swell and winds can cause oil pollutants to mix with the seawater and potentially negatively affect sub-littoral habitats (Castège et al., 2014). Smith (1968) reported dead colonies of Alcyonium digitatum at a depth of 16m in the locality of Sennen Cove, Cornwall which was likely a result of toxic detergents sprayed along the shoreline to disperse oil from the Torrey Cannon tanker spill (Budd, 2008).

Large numbers of dead polychaetes and other fauna were washed up at Rulosquet marsh near Isle de Grand following the Amoco Cadiz oil spill in 1978 (Cross et al., 1978). However, no information was found relating to Spirobranchus triqueter in particular.

Echinus esculentus is subtidal and unlikely to be directly exposed to oil spills. However, as with the ‘Prestige’ oil spill rough seas can cause mixing with the oil and the seawater, and therefore sub-tidal habitats can be affected by the oil spill. Castège et al., (2014) recorded the recovery of rocky shore communities following the Prestige oil spill which impacted the French Atlantic coast. Rough weather at the time of the spill increased mixing between the oil and seawater, causing sub-tidal communities/habitats to be affected. The urchin Echinus esculentus was reported absent after the oil spill however returned after 2-5 years. Large numbers of dead Echinus esculentus were found between 5.5 and 14.5 m in the vicinity of Sennen cove, presumably due to a combination of wave exposure and heavy spraying of dispersants following the ‘Torrey canyon’ oil spill (Smith 1968). Smith (1968) also demonstrated that 0.5 -1ppm of the detergent BP1002 resulted in developmental abnormalities in its echinopluteus larvae. Echinus esculentus populations in the vicinity of an oil terminal in A Coruna Bay, Spain, showed developmental abnormalities in the skeleton. The tissues contained high levels of aliphatic hydrocarbons, naphthalenes, pesticides and heavy metals (Zn, Hg, Cd, Pb, and Cu) (Gomez & Miguez-Rodriguez 1999).

Cliona celata is known to be a hardy sponge, tolerant to a number of abiotic stressors (Duckworth & Peters, 2013). Bustamante et al. (2010) suggested intertidal Cliona celata were negatively affected by the Prestige’ oil spill. However at the time of writing no other information could be gathered on the effects of oil contamination on Cliona celata.

At the time of writing no information could be found relating to the sensitivity of Caryophyllia smithii to hydrocarbon contamination.

Not Assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help
Synthetic compound contamination [Show more]

Synthetic compound contamination

Benchmark. Exposure of marine species or habitat to one or more synthetic compound contaminants via uncontrolled releases or incidental spills (Synthetic compound contamination pressure definition).

Evidence

This pressure is Not assessed but evidence is presented where available.

Smith (1968) reported dead colonies of Alcyonium digitatum at a depth of 16 m in the locality of Sennen Cove, Cornwall resulting from the offshore spread and toxic effect of detergents (a mixture of a surfactant and an organic solvent) e.g. BP 1002 sprayed along the shoreline to disperse oil from the Torrey Canyon tanker spill. Possible sub-lethal effects of exposure to synthetic chemicals, may result in a change in morphology, growth rate or disruption of reproductive cycle. The vulnerability of this species to concentrations of pollutants may also depend on variations in other factors e.g. temperature and salinity conditions outside the normal range (Budd, 2008).

Large numbers of dead Echinus esculentus were found between 5.5 and 14.5 m in the vicinity of Sennen, presumably due to a combination of wave exposure and heavy spraying of dispersants in that area following the Torrey Canyon oil spill (Smith 1968). Smith (1968) also demonstrated that 0.5 -1ppm of the detergent BP1002 resulted in developmental abnormalities in echinopluteus larvae of Echinus esculentus. Echinus esculentus populations in the vicinity of an oil terminal in A Coruna Bay, Spain, showed developmental abnormalities in the skeleton. The tissues contained high levels of aliphatic hydrocarbons, naphthalenes, pesticides and heavy metals (Zn, Hg, Cd, Pb, and Cu) (Gomez & Miguez-Rodriguez 1999).

At the time of writing no information could be found relating to the sensitivity of Caryophyllia smithii to heavy metal contamination.

Cliona celata is known to be a hardy sponge, tolerant to a number of abiotic stressors (Duckworth & Peters, 2013), however at the time of writing no information could be gathered concerning the effects of synthetic compound contamination of Cliona celata.

Not Assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help
Radionuclide contamination [Show more]

Radionuclide contamination

Benchmark. An increase in 10µGy/h above background levels (Radionuclides contamination pressure definition).

Evidence

'No Evidence'.

No evidence (NEv)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		No evidence (NEv)
NR
NR
NR
Help
Introduction of other substances [Show more]

Introduction of other substances

Benchmark. Exposure of marine species or habitat to one or more relevant "other" substances (solid, liquid or gas) contaminants via uncontrolled releases or incidental spills (Introduction of other substances pressure definition). 

Evidence

This pressure is Not assessed.

Not Assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help
De-oxygenation [Show more]

De-oxygenation

Benchmark. Exposure to dissolved oxygen concentration of less than or equal to 2 mg/l for one week (a change from WFD poor status to bad status) (deoxygenation pressure definition).

Evidence

There is anecdotal evidence to suggest that Alcyonium digitatum, Caryophyllia smithii and Echinus esculentus are sensitive to hypoxic events.

In general, respiration in most marine invertebrates do not appear to be significantly affected until extremely low concentrations are reached. For many benthic invertebrates this concentration is about 2 ml l-1, or even less (Herreid, 1980; Rosenberg et al., 1991; Diaz & Rosenberg, 1995).

Alcyonium digitatum mainly inhabits environments in which the oxygen concentration usually exceeds 5 ml l-1 and respiration is aerobic (Budd, 2008). In August 1978 a dense bloom of a dinoflagellate, Gyrodinium aureolum occurred surrounding Geer reef in Penzance Bay, Cornwall and persisted until September that year. Observations by local divers indicated a decrease in underwater visibility (<1 m) from below 8 m Below Sea Level. It was also noted that many of the faunal species appeared to be affected, e.g. no live Echinus esculentus were observed whereas on surveys prior to August were abundant. Alcyonium sp. and Bryozoans were also in an impoverished state. Caryophyllia smithii were also in a contracted state, apparently dead, and with Echinus esculentus were the worst affected species during the bloom. During follow up surveys conducted in early September Alcyonium sp. were noted to be much healthier and feeding. It was suggested the decay of Gyrodinium aureolum either reduced oxygen levels or physically clogged faunal feeding mechanisms. Adjacent reefs where also surveyed during the same time period and the effects of the Gyrodinium aureolum bloom were less apparent. It was suggested that higher water agitation in shallow water on reefs more exposed to wave action were less effected by the phytoplankton bloom (Dennis, 1979).

CR.MCR.EcCr.CarSp is recorded from negligible-weak tidal streams (<0.5 m/sec) but can occur at wave exposed sites (Connor et al., 2004). Therefore, water movement (through wave action and/or tidal flow) could potentially cause mixing with surrounding oxygenated water (Dennis, 1979) and may therefore decrease the effects of de-oxygenation rapidly.

Sensitivity assessment. Resistance has been assessed as ‘Low’, Resilience as ‘Medium’. Sensitivity as ‘Medium’.

Low
Low
NR
NR
Help		Medium
Low
NR
NR
Help		Medium
Low
Low
Low
Help
Nutrient enrichment [Show more]

Nutrient enrichment

Benchmark. Increased levels of the elements nitrogen, phosphorus, silicon, and iron in the marine environment compared to background concentrations (Nutrient enrichment pressure definition).

Evidence

This biotope is considered to be 'Not sensitive' at the pressure benchmark that assumes compliance with good status as defined by the WFD.

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are suspension feeders. Nutrient enrichment of coastal waters that enhances the population of phytoplankton may be beneficial to Alcyonium digitatum, Caryophyllia smithii and Spirobranchus triqueter in terms of an increased food supply but the effects are uncertain (Hartnoll, 1998). The survival of Alcyonium digitatum, Caryophyllia smithii and Spirobranchus triqueter may be influenced indirectly. High primary productivity in the water column combined with high summer temperature and the development of thermal stratification (which prevents mixing of the water column) can lead to hypoxia of the bottom waters which faunal species are likely to be highly intolerant of (see de-oxygenation pressure).

Cliona celata is considered a hardy sponge, tolerant of environmental stressors such as high nutrient loads, low salinity, and large temperature variation (Duckworth & Peters, 2013). Carballo et al. (1994) suggested Cliona celata was a good indicator species of pollution, and noted it’s abundance at polluted sites which receive sewage discharge within Algeciras Bay, Spain.

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types.

It was suggested by Comely & Ansell (1988) that Echinus esculentus could absorb dissolved organic material for the purposes of nutrition. Nutrient enrichment may encourage the growth of ephemeral and epiphytic algae and therefore increase sea-urchin food availability. Lawrence (1975) reported that sea urchins had persisted over 13 years on barren grounds near sewage outfalls, presumably feeding on dissolved organic material, detritus, plankton and microalgae, although individuals died at an early age.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not sensitive
NR
NR
NR
Help
Organic enrichment [Show more]

Organic enrichment

Benchmark. A deposit of 100 gC/m2/yr (Organic enrichment pressure definition).

Evidence

Alcyonium digitatum, Caryophyllia smithii and Spirobranchus triqueter are suspension feeders of phytoplankton and zooplankton. Organic enrichment of coastal waters that enhances the population of phytoplankton may be beneficial to Alcyonium digitatum, Caryophyllia smithii and Spirobranchus triqueter in terms of an increased food supply but the effects are uncertain (Hartnoll, 1998). The survival of Alcyonium digitatum, Caryophyllia smithii and Spirobranchus triqueter may be influenced indirectly. High primary productivity in the water column combined with high summer temperature and the development of thermal stratification (which prevents mixing of the water column) can lead to hypoxia of the bottom waters which faunal species are likely to be highly intolerant of (see de-oxygenation pressure).

Cliona celata is considered a hardy sponge, tolerant of environmental stressors such as high nutrient loads, low salinity, and large temperature variation (Duckworth & Peters, 2013). Carballo et al. (1994) suggested Cliona celata was a good indicator species of pollution, and noted it’s abundance at polluted sites which receive sewage discharge within Algeciras Bay, Spain.

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types.

It was suggested by Comely & Ansell (1988) that Echinus esculentus could absorb dissolved organic material for the purposes of nutrition. Organic enrichment may encourage the growth of ephemeral and epiphytic algae and therefore increase sea-urchin food availability. Lawrence (1975) reported that sea urchins had persisted over 13 years on barren grounds near sewage outfalls, presumably feeding on dissolved organic material, detritus, plankton and microalgae, although individuals died at an early age.

Sensitivity assessment. Organic enrichment is not likely to directly negatively affect the characterizing species within this biotope, however chronic organic enrichment may cause secondary effects such as hypoxia (refer to de-oxygenation pressure). Resistance has been assessed as ‘Low’, Resilience as ‘Medium’. Sensitivity as ‘Medium’.

Low
Low
NR
NR
Help		Medium
Low
NR
NR
Help		Medium
Low
Low
Low
Help

Physical Pressures

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ResistanceResilienceSensitivity
Physical loss (to land or freshwater habitat) [Show more]

Physical loss (to land or freshwater habitat)

Benchmark. A permanent loss of existing saline habitat within the site (Physical loss pressure definition). 

Evidence

All marine habitats and benthic species are considered to have a resistance of ‘None’ to this pressure and to be unable to recover from a permanent loss of habitat (resilience is ‘Very Low’). Sensitivity within the direct spatial footprint of this pressure is therefore ‘High’. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.

None
High
High
High
Help		Very Low
High
High
High
Help		High
High
High
High
Help
Physical change (to another seabed type) [Show more]

Physical change (to another seabed type)

Benchmark. Permanent change from sedimentary or soft rock substrata to hard rock or artificial substrata, or vice versa (Physical change in subtratum type pressure definition).

Evidence

If rock were replaced with sediment, this would represent a fundamental change to the physical character of the biotope and the species would be unlikely to recover. The biotope would be lost.

Sensitivity assessment. Resistance to the pressure is considered ‘None’, and resilience ‘Very low’. Sensitivity has been assessed as ‘High’.

None
High
High
High
Help		Very Low
High
High
High
Help		High
High
High
High
Help
Physical change (to another sediment type) [Show more]

Physical change (to another sediment type)

Benchmark. Permanent change in one Folk class (based on UK SeaMap simplified classification) (Physical change in sediment type pressure definition). 

Evidence

Not relevant

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Habitat structure changes - removal of substratum (extraction) [Show more]

Habitat structure changes - removal of substratum (extraction)

Benchmark. The extraction of substratum to 30 cm (where substratum includes sediments and soft rock but excludes hard bedrock) (Removal of substratum pressure definition). 

Evidence

The species characterizing this biotope are epifauna or epiflora occurring on rock and would be sensitive to the removal of the habitat. However, extraction of rock substratum is considered unlikely and this pressure is considered to be ‘Not relevant’ to hard substratum habitats.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Abrasion / disturbance of the surface of the substratum or seabed [Show more]

Abrasion / disturbance of the surface of the substratum or seabed

Benchmark. Damage to surface features (e.g. species and physical structures within the habitat) (Surface abrasion/disturbance pressure definition).

Evidence

CR.MCR.EcCr.CarSp is a subtidal biotope (Connor et al., 2004). Therefore abrasion is most likely to be a result of bottom or pot fishing gear, cable laying etc. which may cause localised mobility of the substrata and mortality of the resident community. The effect would be situation dependent however if bottom fishing gear were towed over a site it may mobilise a high proportion of the rock substrata and cause high mortality in the resident community.

Alcyonium digitatum, Caryophyllia smithii, Echinus esculentus and Spirobranchus triqueter are sedentary or slow moving species that might be expected to suffer from the effects of dredging. Boulcott & Howell (2011) conducted experimental Newhaven scallop dredging over a circalittoral rock habitat in the sound of Jura, Scotland and recorded the damage to the resident community. The results indicated that the sponge Pachymatisma johnstoni was highly damaged by the experimental trawl. However, only 13% of photographic samples showed visible damage to Alcyonium digitatum. Where Alcyonium digitatum damage was evident it tended to be small colonies that were ripped off the rock. The authors highlight physical damage to faunal turfs (erect bryozoans and hydroids) was difficult to quantify in the study. However, the faunal turf communities did not show large signs of damage and were only damaged by the scallop dredge teeth which was often limited in extent (approximately. 2cm wide tracts). The authors indicated that species such as Alcyonium digitatum and faunal turf communities were not as vulnerable to damage through trawling as sedimentary fauna and whilst damage to circalittoral rock fauna did occur it was of an incremental nature, with loss of species such as Alcyonium digitatum and faunal turf communities increasing with repeated trawls.

Species with fragile tests, such as Echinus esculentus were reported to suffer badly as a result of scallop or queen scallop dredging (Bradshaw et al., 2000; Hall-Spencer & Moore, 2000). Kaiser et al. (2000) reported that Echinus esculentus were less abundant in areas subject to high trawling disturbance in the Irish Sea. Jenkins et al. (2001) conducted experimental scallop trawling in the North Irish sea and recorded the damage caused to several conspicuous megafauna species, both when caught as bi-catch and when left on the seabed. The authors predicted 16.4% of Echinus esculentus were crushed/dead, 29.3% would have >50% spine loss/minor cracks, 1.1% would have <50% spine loss and the remaining 53.3% would be in good condition. Sea urchins can rapidly regenerate spines, e.g. Psammechinus miliaris were found to re-grow all spines within a period of 2 months (Hobson, 1930).  The trawling examples mentioned above were conducted on sedimentary habitats and thus the evidence is not directly relevant to the rock based biotopes- CR.MCR.EcCr.CarSp however does indicate the likely effects of abrasion on Echinus esculentus.

Sensitivity assessment. Resistance has been assessed ‘Medium’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Low’

Please note Boulcott & Howell (2011) did not mention the abrasion caused by fully loaded collection bags on the new haven dredges. A fully loaded Newhaven dredge may cause higher damage to community as indicated in their study.

Medium
High
High
High
Help		High
High
High
High
Help		Low
High
High
High
Help
Penetration or disturbance of the substratum subsurface [Show more]

Penetration or disturbance of the substratum subsurface

Benchmark. Damage to sub-surface features (e.g. species and physical structures within the habitat) (Sub-surface penetration pressure definition).

Evidence

The species characterizing this biotope group are epifauna or epiflora occurring on rock which is resistant to subsurface penetration.  The assessment for abrasion at the surface only is therefore considered to equally represent sensitivity to this pressure. This pressure is 'Not relevant' to hard rock biotopes.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Changes in suspended solids (water clarity) [Show more]

Changes in suspended solids (water clarity)

Benchmark. A change in one rank on the WFD (Water Framework Directive) scale, e.g. from clear to intermediate for one year (Suspended sediment pressure definition).

Evidence

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are not thought highly susceptible to changes in water clarity due to the fact they are suspension feeding organisms and are not directly dependent on sunlight for nutrition. Alcyonium digitatum has been shown to be tolerant of high levels of suspended sediment. Hill et al. (1997) demonstrated that Alcyonium digitatum sloughed off settled particles with a large amount of mucous. Alcyonium digitatum is also known to inhabit the entrances to sea lochs (Budd, 2008) or the entrances to estuaries (Braber & Borghouts, 1977) where water clarity is likely to be highly variable.

Moore (1977) suggested that Echinus esculentus was unaffected by turbid conditions. Echinus esculentus is an important grazer of red macro-algae within CR.MCR.EcCr. Increased turbidity and resultant reduced light penetration is likely to negatively affect algal growth. However, Echinus esculentus can feed on alternative prey, detritus or dissolved organic material (Lawrence, 1975, Comely & Ansell, 1988).

Increased turbidity will reduce light penetration and hence phytoplankton productivity. Small phytoplankton are probably an important food source in the shallow subtidal, although, Flustra foliacea is also found at greater depths, where organic particulates (detritus) are probably more important.

According to Bacescu (1972), sabellids are accustomed to turbidity and silt. Spirobranchus triqueter has also recently been recorded by De Kluijver (1993) from Scotland in the aphotic zone, indicating that the species would not be sensitive to an increase in turbidity.

Sensitivity assessment. Resistance has been assessed as ‘High’, Resilience as ‘High’. Sensitivity has been assessed as ‘Not Sensitive’.

High
High
High
High
Help		High
High
High
High
Help		Not sensitive
High
High
High
Help
Smothering and siltation rate changes (light) [Show more]

Smothering and siltation rate changes (light)

Benchmark. ‘Light’ deposition of up to 5 cm of fine material added to the seabed in a single discrete event (Smothering pressure definition).

Evidence

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are not thought highly susceptible to changes in water clarity due to the fact they are suspension feeding organisms and are not directly dependent on sunlight for nutrition. Alcyonium digitatum has been shown to be tolerant of high levels of suspended sediment. Hill et al. (1997) demonstrated that Alcyonium digitatum sloughed off settled particles with a large amount of mucous. Alcyonium digitatum is also known to inhabit the entrances to sea lochs (Budd, 2008) or the entrances to estuaries (Braber & Borghouts, 1977) where water clarity is likely to be highly variable.

Moore (1977) suggested that Echinus esculentus was unaffected by turbid conditions. Echinus esculentus is an important grazer of red macro-algae within CR.MCR.EcCr. Increased turbidity and resultant reduced light penetration is likely to negatively affect algal growth. However, Echinus esculentus can feed on alternative prey, detritus or dissolved organic material (Lawrence, 1975, Comely & Ansell, 1988).

Increased turbidity will reduce light penetration and hence phytoplankton productivity. Small phytoplankton are probably an important food source in the shallow subtidal, although, Flustra foliacea is also found at greater depths, where organic particulates (detritus) are probably more important.

According to Bacescu (1972), sabellids are accustomed to turbidity and silt. Spirobranchus triqueter has also recently been recorded by De Kluijver (1993) from Scotland in the aphotic zone, indicating that the species would not be sensitive to an increase in turbidity.

Sensitivity assessment. Resistance has been assessed as ‘High’, Resilience as ‘High’. Sensitivity has been assessed as ‘Not Sensitive’.

High
Low
NR
NR
Help		High
High
High
High
Help		Not sensitive
Low
Low
Low
Help
Smothering and siltation rate changes (heavy) [Show more]

Smothering and siltation rate changes (heavy)

Benchmark. ‘Heavy’ deposition of up to 30 cm of fine material added to the seabed in a single discrete event (Smothering pressure definition).

Evidence

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are sessile and thus would be unable to avoid the deposition of a smothering layer of sediment. Some Alcyonium digitatum colonies can attain a height of up to 20 cm (Edwards, 2008), so would still be able to feed in the event of sediment deposition. However, Spirobranchus triqueter is an encrusting species and would thus likely be smothered, and depending on sediment retention could block larval settlement. Cliona celata can either have a boring life form (it bores into rock) or a massive form (grows ontop of rock). With the boring form only inhalant and exhalent papillae are visible just above the rock surface (Wood, 2007). The massive form has raised, rounded ridges with large specimens growing up to 1 m across and 50 cm high (Snowden, 2007). Haliclona viscosa is described as a cushion forming species, which forms a thick crust which may reach a diameter of 30-40 cm and a height of 1.5-5cm (Topsent, 1888). Pachymatisma johnstonia can reach up to 15 cm in diameter and height up to 10 cm (Neish, 2007).

 

Comely & Ansell (1988) recorded large Echinus esculentus from kelp beds on the west coast of Scotland in which the substratum was seasonally covered with "high levels" of silt. This suggests that Echinus esculentus is unlikely to be killed by smothering, however, smaller specimens and juveniles may be less resistant. A layer of sediment may interfere with larval settlement.  If retained within the host biotope for extended periods a layer of 5cm of the sediment may negatively affect successive recruitment events.

Caryophyllia smithii is small (approx. <3 cm height from the seabed) and would therefore likely be inundated in a “light” sedimentation event. However Bell & Turner (2000) reported Caryophyllia smithii was abundant at sites of “moderate” sedimentation (7mm ± 0.5mm) in Lough Hyne. It is therefore likely that Caryophyllia smithii would be resistant to periodic sedimentation. If 5cm of sediment were removed rapidly, via tidal currents, Caryophyllia smithii would likely remain within the biotope.

It is likely that 30cm of deposited sediment may inundate a significant proportion of the encrusting community within CR.MCR.EcCr.CarSp. However, CR.MCR.EcCr.CarSp is recorded from moderately strong to negligible tidal currents (<1.5m/sec) (Connor et al., 2004). Therefore, water movement (through wave action and/or tidal flow) would be expected to clear deposited sediment.

Sensitivity assessment. Resistance has been assessed as ‘Medium’, resilience as ‘High’. Sensitivity has therefore been assessed as ‘Low’.

Medium
Low
NR
NR
Help		High
High
High
High
Help		Low
Low
Low
Low
Help
Litter [Show more]

Litter

Benchmark. The introduction of man-made objects able to cause physical harm (surface, water column, seafloor or strandline) (Litter pressure definition). 

Evidence

Not assessed.

Not Assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help		Not assessed (NA)
NR
NR
NR
Help
Electromagnetic changes [Show more]

Electromagnetic changes

Benchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT (Electromagnetic pressure definition).

Evidence

'No evidence' was found.

No evidence (NEv)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		No evidence (NEv)
NR
NR
NR
Help
Underwater noise changes [Show more]

Underwater noise changes

Benchmark. MSFD indicator levels (SEL or peak SPL) exceeded for 20% of days in a calendar year. Further detail

Evidence

Alcyonium digitatum, Caryophyllia smithii, Echinus esculentus, Spirobranchus triqueter and sponges have no hearing perception but vibrations may cause an impact, however no studies exist to support an assessment (where relevant).

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Introduction of light or shading [Show more]

Introduction of light or shading

Benchmark. A change in incident light via anthropogenic means (Introduced light or shade pressure definition).

Evidence

There was no evidence to suggest that If exposed to anthropogenic light sources algal species would benefit. CR.MCR.EcCr.CarSp is also a circalittoral biotope and are thus by definition naturally shaded environments with low light levels. Increased shading (e.g. by construction of a pontoon, pier etc) could be beneficial to the characterizing species within these biotopes.

Sensitivity assessment. Resistance is probably 'High', with a 'High' resilience and a sensitivity of 'Not Sensitive'.

High
High
High
High
Help		High
High
High
High
Help		Not sensitive
High
High
High
Help
Barrier to species movement [Show more]

Barrier to species movement

Benchmark. A permanent or temporary barrier to species movement over ≥50% of water body width or a 10% change in tidal excursion (Barrier to species movement pressure definition).

Evidence

Barriers and changes in tidal excursion are 'Not relevant' to biotopes restricted to open waters.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Death or injury by collision [Show more]

Death or injury by collision

Benchmark. Injury or mortality from collisions of biota with both static or moving structures due to 0.1% of tidal volume on an average tide, passing through an artificial structure (Death for collision pressure definition).

Evidence

'Not relevant' to seabed habitats.  NB. Collision by grounding vessels is addressed under ‘surface abrasion’.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Visual disturbance [Show more]

Visual disturbance

Benchmark. The daily duration of transient visual cues exceeds 10% of the period of site occupancy by the feature (Visual disturbance pressure definition). 

Evidence

'Not relevant'

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help

Biological Pressures

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ResistanceResilienceSensitivity
Genetic modification & translocation of indigenous species [Show more]

Genetic modification & translocation of indigenous species

Benchmark. Translocation of indigenous species or the introduction of genetically modified or genetically different populations of indigenous species may result in changes in the genetic structure of local populations, hybridization, or a change in community structure (Translocation pressure definition).

Evidence

Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges are not commercially cultivated within the UK or likely to be translocated. Xavier et al. (2010) suggested Cliona celtata was in fact a species complex of potentially four morphologically indistinct species. Cliona celata is also a “pest” species within scallop mariculture (Carver et al., 2010), where Cliona celata bores into reared scallop shells. It is therefore conceivable that separate species within the Cliona celata species complex could be transported outside of it’s traditional range, however at the time of writing there is no evidence to suggest translocation would negatively affect CR.MCR.EcCr.CarSp. Echinus esculentus was identified by Kelly & Pantazis (2001) as a species suitable for culture for the urchin Roe industry. However, at the time of writing no evidence could be found to suggest that significant Echinus esculentus mariculture was present in the UK. If industrially cultivated it is feasible that Echinus esculentus individuals could be translocated. This pressure is therefore considered ‘Not relevant’ at the time of writing.

Translocation also has the potential to transport pathogens to uninfected areas (see pressure ‘introduction of microbial pathogens’). The sensitivity of the ‘donor’ population to harvesting to supply stock for translocation is assessed for the pressure ‘removal of target species’.

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Introduction of microbial pathogens [Show more]

Introduction of microbial pathogens

Benchmark. The introduction of relevant microbial pathogens or metazoan disease vectors to an area where they are currently not present (e.g. Martelia refringens and Bonamia, Avian influenza virus, viral Haemorrhagic Septicaemia virus) (pathogen or disease pressure definition).

Evidence

‘No evidence’ was found to suggest that any of the characterizing species within CR.MCR.EcCr.CarSp are sensitive to current/known microbial pathogens.

Alcyonium digitatum acts as the host for the endoparasitic species Enalcyonium forbesiand and Enalcyonium rubicundum (Stock, 1988). Parasitisation may reduce the viability of a colony but not to the extent of killing them but no further evidence was found to substantiate this suggestion. Thomas (1940) recorded parasites of Spirobranchus triqueter. Trichodina pediculus (a ciliate) was observed in high numbers moving over the branchial crown. However, this relationship is symbiotic, not parasitic. Parasites found in the worm include gregarines & ciliated protozoa and parasites that had the appearance of sporozoan cysts. However, no information was found about the effects of microbial pathogens on Spirobranchus triqueter. Echinus esculentus is susceptible to 'Bald-sea-urchin disease', which causes lesions, loss of spines, tube feet, pedicellariae, destruction of the upper layer of skeletal tissue and death. It is thought to be caused by the bacteria Vibrio anguillarum and Aeromonas salmonicida. Bald sea-urchin disease was recorded from Echinus esculentus on the Brittany Coast. Although associated with mass mortalities of Strongylocentrotus franciscanus in California and Paracentrotus lividus in the French Mediterranean it is not known if the disease induces mass mortality (Bower, 1996). At the time of writing, there was no evidence concerning microbial pathogens that may affect Cliona celata.

No evidence (NEv)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		No evidence (NEv)
NR
NR
NR
Help
Removal of target species [Show more]

Removal of target species

Benchmark. Removal of species targeted by fishery, shellfishery or harvesting at a commercial or recreational scale (targeted removal pressure definition).

Evidence

At the time of writing none of the characterizing species within CR.MCR.EcCr.CarSp are commercially exploited. This pressure is considered ‘Not Relevant’.

Echinus esculentus was identified by Kelly & Pantazis (2001) as a species suitable for culture for the urchin Roe industry. However, at the time of writing no evidence could be found to suggest that significant Echinus esculentus mariculture was present in the UK. Removal of Echinus esculentus from CR.MCR.EcCr.CarSp could cause an increase in algal growth which may limit the growth of faunal species (Bell & Turner, 2000; Connor et al., 2004).

Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help
Removal of non-target species [Show more]

Removal of non-target species

Benchmark. Removal of features or incidental non-targeted catch (by-catch) through targeted fishery, shellfishery or harvesting at a commercial or recreational scale (non-targeted removed pressure definition).

Evidence

Faunal turf communities are probably resistant to abrasion through bottom fishing (see abrasion pressure).

Alcyonium digitatum goes through an annual cycle, From Febuary to July all Alcyonium digitatum colonies are feeding, from July to November an increasing number of colonies stop feeding. During this period a large number of polyps can retracts and a variety of filamentous algae, hydroids and amphipods can colonize the surface of colonies epiphytically. From December-February the epiphytic community is however sloughed off (Hartnoll, 1975). If Alcyonium digitatum were removed the epiphytic species would likely colonize rock surfaces and are therefore not dependant on Alcyonium digitatum.

 

Within CR.MCR.EcCr.CarSp Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges spatially compete, however at the time of writing there isn’t any evidence to suggest other interspecific relationships or dependencies between these species. Therefore removal of 1 or a number of these species would provide colonization space and most likely benefit the species with rapid colonization rates (e.g. Spirobranchus triqueter). Echinus esculentus is an important red algae grazer within CR.MCR.EcCr (Connor et al., 2004), without which the abundance of red algae may increase and possibly displace some of the faunal turf species. If Alcyonium digitatum, Caryophyllia smithii, Spirobranchus triqueter and sponges were removed this would alter the character of the biotope.

Sensitivity assessment. Resistance has been assessed as ‘Medium’, resilience has been assessed as ’High’. Sensitivity has been assessed as ‘Low’.

Medium
High
High
High
Help		High
High
High
High
Help		Low
High
High
High
Help

Introduction or spread of invasive non-indigenous species (INIS) Pressures

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ResistanceResilienceSensitivity
Other INIS [Show more]

Other INIS

Evidence

Crepidula fornicata larvae require hard substrata for settlement. It prefers muddy gravelly, shell-rich, substrata that include gravel, or shells of other Crepidula, or other species e.g., oysters, and mussels. It is highly gregarious and seeks out adult shells for settlement, forming characteristic ‘stacks’ of adults. But it also recorded from rock, artificial substrata, and Sabellaria alveolata reefs (Blanchard, 1997, 2009; Bohn et al., 2012, 2013a, 2013b, 2015; De Montaudouin et al., 2018; Hinz et al., 2011; Helmer et al., 2019; Powell-Jennings & Calloway, 2018; Preston et al., 2020; Tillin et al., 2020). Close examination of the literature (2023) shows that evidence of its colonization and density on bedrock in the infralittoral or circalittoral was lacking. Tillin et al. (2020) suggested that Crepidula could colonize circalittoral rock due to its presence on tide-swept rough grounds in the English Channel (Hinz et al., 2011). However, Hinz et al. (2011) reported that Crepidula fornicata only dominated one assemblage (with an average of 181 individuals per trawl) on gravel substratum with boulders. Bohn et al. (2015) noted that Crepidula occurred at low density or was absent in areas dominated by boulders, and Bohn et al. (2013a, 2013b, 2015) and Preston et al. (2020) showed that while Crepidula could settle on slate panels or ‘stone’ it preferred shell, especially that of conspecifics. In addition, no evidence was found of the effect of Crepidula populations on faunal turf-dominated habitats. It was only recorded at low density (0.1-0.9/m2) in one faunal turf biotope (CR.MCR.CFaVS.CuSpH.As) (JNCC, 2015). Faunal turfs are dominated by suspension feeders so larval predation is probably high, which may prevent colonization by Crepidula. Also, faunal turf species actively compete for space and many are fast growing and opportunistic, so may out-compete Crepidula for space even if it gained a foothold in the community. 

Didemnum vexillum is an invasive colonial sea squirt native to Asia which was first recorded in the UK in Darthaven Marina, Dartmouth in 2005. Didemnum vexillum can form extensive mats over the substrata it colonizes; binding boulders and cobbles and altering the host habitat (Griffith et al., 2009). Didemnum vexillum can also grow over and smother the resident biological community. Recent surveys within Holyhead Marina, North Wales have found Didemnum vexillum growing on and smothering native tunicate communities (Griffith et al., 2009). Due to the rapid-re-colonization of Didemnum vexillum eradication attempts have to date failed. 

Didemnum vexillum is isolated to several sheltered locations in the UK (NBN, 2015), however, Didemnum vexillum has successfully colonized the offshore location of the Georges Bank, USA (Lengyel et al., 2009) which is more exposed than the locations which Didemnum vexillum have colonized in the UK. It is therefore possible that Didemnum vexillum could colonize more exposed locations within the UK and could therefore pose a threat to CR.MCR.EcCr.CarSp.

Sensitivity assessment. The circalittoral rock characterizing this biotope is likely to be unsuitable for the colonization by Crepidula fornicata due to the extremely wave exposed to moderately wave exposed conditions, in which wave action and storms may mitigate or prevent the colonization by Crepidula at high densities, although Crepidula has been recorded from areas of strong tidal streams (Hinz et al., 2011). Crepidula has been recorded from the lower intertidal to ca 160 m in depth but is most common in the shallow subtidal above 50 m (Blanchard, 1997; Thieltges et al., 2003; Bohn et al., 2012, 2015; Hinz et al., 2011; OBIS, 2023; Tillin et al., 2020), therefore, colonization of Crepidula would be limited to low densities in deeper examples of the biotope. In addition, no evidence was found of the effect of Crepidula populations on faunal turf-dominated habitats or infralittoral or circalittoral rock habitats. At present, there is 'Insufficient evidence' to suggest that the circalittoral rock biotopes are sensitive to colonization by Crepidula fornicata or other invasive species; further evidence is required. 

Insufficient evidence (IEv)
NR
NR
NR
Help		Not relevant (NR)
NR
NR
NR
Help		Insufficient evidence (IEv)
NR
NR
NR
Help

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This review can be cited as:

Stamp, T.E., Lloyd, K.A., & Watson, A.J., 2023. Caryophyllia (Caryophyllia) smithii, sponges and crustose communities on wave-exposed circalittoral rock. In Tyler-Walters H. Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 14-05-2026]. Available from: https://www.marlin.ac.uk/habitat/detail/6

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