Psolus squamatus, Anomiidae, serpulid polychaetes and Munida on Atlantic mid bathyal rock and other hard substrata

Summary

UK and Ireland classification

Description

This biotope consists of cobble, boulder and bedrock areas with the holothurian Psolus being the most conspicuous faunal component. A range of encrusting species are also present but there is no conspicuous presence of lamellate sponges as there is within 'Psolus squamatus and encrusting sponge assemblage'. This assemblage also occurs in the upper bathyal. The characterizing species listed refer to all Psolus squamatus, Anomiidae, serpulid polychaetes and Munida assemblages not just those associated with the zone and substrata specified in this biotope. (Information from JNCC, 2022).

Depth range

600-1300 m

Additional information

This assemblage occurs from the upper to the mid-bathyal (M.AtUB.Ro.SpaEnc.PsoSpo and M.AtMB.Ro.SpaEnc.PsoSpo). It is very similar to other Psolus squamatus dominated biotopes M.AtMB.Ro.SpaEnc.PsoAno and M.AtUB.Ro.SpaEnc.PsoAno that also occur in the upper to mid bathyal or similar substratum. The two groups of biotopes differ in the abundance of encrusting sponges (PsoSpo) and saddle oysters and serpulids (PsoAno). However, the only consistent characteristic species is Psolus squamatus. Therefore, the following review considers the Psolus squamatus dominated, sparse encrusting communities (SpaEnc.PsoSpo and SpaEnc.PsoAno) as a group.

The biotope description is based on habitats mapped by Davies et al. (2015) on the Anton Dohr Seamount and recorded by Howell et al. (2010) from Hatton Bank, the Rosemary Bank Seamount, the Faroe-Shetland Channel and South-West Canyons, and by Weinberg et al. (2008) from the Franken Mound, Rockall Bank. Similar Psolus squamatus dominated habitats were also recorded from the Hatton Bank (Roberts et al., 2008), Sognefjord (Meyer et al., 2020) and Hardangerfjord, Norway (Buhl-Mortensen & Buhl-Mortensen, 2014) and the Oregon continental slope (Hemery et al., 2018). 

Listed By

Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

The SpaEnc.PsoSpo and SpaEnc.PsoAno biotopes are defined by the abundance of Psolus squamatus and hard substrata (bedrock, boulders, and cobbles) and sparse encrusting species; sponges in PsoSpo and saddle oysters and serpulids in PsoAno (JNCC, 2022). Davis et al. (2015) described a Psolus squamatus and sponge dominated community, with caryophyllids and lamellate sponges on mixed substrata with boulders and bedrock, at 854 to 1345 m on the steep escarpment of Anton Dohr Seamount, and another Psolus squamatus, serpulid and encrusting sponge dominated community on mixed substrata at 813 to 1037 m. Davies et al. (2015) noted that the ophiuroid Ophiactis balli, corals (occasional Lophelia, and the antipatharian Leiopathes sp.) were present but sponges were more abundant in the sponge-dominated biotope. Davis et al. (2015) also described a similar biotope along a bedrock escarpment on Rockall Bank at 350 to 600 m, which included large lobose sponges, stylastarid corals and the pencil urchin Cidaris cidaris. The Psolus squamatus and serpulid dominated biotope included encrusting and globose sponges, the ophiuroid Ophiactis abyssicola, and spider crabs. Howell et al. (2010) described a Psolus squamatus assemblage that included Anomiidae, encrusting sponges, serpulids, brachiopods and Munida spp. on cobbles, boulders and bedrock at 332 to 963 m from Hatton Bank, the Rosemary Bank Seamount, the Faroe-Shetland Channel and South-West Canyons. Wienberg et al. (2008) described similar assemblages of Psolus sp., serpulids and bryozoans on gravel to large boulder-sized fields of 'dropstones', mainly on soft sediment on the flanks of the Franken Mound, Rockall Bank. Large pebbles and boulders were also colonized by calcified hydroids, Stylaster sp., actinians (Phelliactis hertwigi), octocorals, and sponges, while the largest boulders sheltered decapods (e.g. Munida sp, and Pagurus sp.) and fish. Wienberg et al. (2008) noted that the eastern flank of Franken Mound was subject to strong but variable bottom currents, indicated by extensive areas of soft sediment, exhumed dropstones, rocky outcrops, and mobile rippled sediment, characteristic of active sediment transport. Roberts et al. (2008) reported that scattered pebbles and larger boulders on sandy sediment were heavily colonized by epifauna, including Psolus sp. (probably Psolus squamatus), at 476 to 539 m on Hatton Bank. Psolus sp. frequently colonized exposed rock in coarse sandy sediments at 518 to 582 m, and exposed rock with a thin layer of sediment at 466 to 482 m (Roberts et al., 2008). Scattered hard substrata (pebbles to large boulders) on sediment were typically colonized by Psolus spp. a species typical of cobbles, dropstones and exposed rock (Roberts et al., 2008). Hemery et al. (2018) described an assemblage dominated by a high density of Psolus squamatus on mixed sediments at 170 to 370 m, with encrusting invertebrates, an unidentified ophiuroid, unidentified small shrimps and the lantern shell Laqueus californicus on the Oregon continental shelf. Psolus squamatus was often observed concentrated at breaks in the slope or on protruding surfaces in areas of mixed substrata (i.e. exposed hard substratum and soft sediment) in the Sognefjord (Meyer et al., 2020). 

The abundance of Psolus squamatus (and other Psolids) depends on the presence of hard substrata, to which they attach using their characteristic 'creeping' sole (Ekman, 1923; Mortensen, 1927; Hyman, 1955; Roberts et al., 2008). Psolus sp. are suspension feeders that capture food particles between their extended tentacles or on adhesive papillae or buds (Fankboner, 1978; Massin, 1982).  Suspension feeding sea cucumbers typically extend their tentacles into flowing water and position themselves on extremities or the top of rocks to position themselves in the current (Massin, 1982). For example, Psolus chitinoides was able to open the tentacular crown into an open-mesh cup and move tentacles towards passing food, or bend the tentacles inwards to form a semi-closed cup-like mesh that promoted the capture of large, inanimate material (Fankboner, 1978; Massin, 1982). This biotope is characterized by suspension feeders such as the encrusting, lamellate and globose spongs, serpulids, anomids, ophiuroids (Ophiactis spp.) and occasional hydroids and corals (Howell et al., 2010; Davis et al., 2015). Suspension feeding sea cumbers such as the Psolids are found in areas of strong currents (Gage et al., 1985; Billett, 1991). 

Psolus squamatus is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight, characterized by egg sizes indicative of lecithotrophic development and near sea bed development (Billett, 1991; Wagstaff et al., 2014). Billett (1991) noted that currents generally ran parallel to the slope contours and could retain larvae and juveniles close to areas suitable for development (see recovery below). This zonation is typical of marine invertebrates in deep-sea habitats and corresponds to the upper boundary biota identified by Carney (2005). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf. Carney (2005) noted that currents tend to flow along the slope due to density stratification and may separate larval transport between the shelf and the upper slope, although internal waves near the edge of the shelf may provide larval transport for near bottom larvae. The transition at about 1000 m is influenced by decreased cooling from the permanent thermocline and decreased food influx and sees the loss of most upper boundary species (Carney, 2005). 

Psolus squamatus is the only species identified consistently in this biotope (habitat). The characteristic encrusting or lamellate sponges, Anomiidae, and serpulids are not identified to genus level (or rarely so), while long-lived species (e.g. corals, globose sponges) occur in small numbers. Other species are mobile and wide-spread e.g. ophiuroids are mobile and aggregate in areas suitable for suspension feeding while e.g. squat lobsters use the habitat for shelter. No information on the ecology of this habitat was found. However, the lack of long-lived species on the available hard substrata suggests the current flow does not support large sponge or cold-water corals and that the habitat is exposed to periodic disturbance, probably due to sediment mobility, scour and burial. The habitat is characterized by exposed rock outcrops amongst sediment, or mixed hard substratum (boulders, pebbles, and gravel etc.) on the surface of potentially mobile sediment (Wienberg et al., 2008; Roberts et al., 2008; Davis et al., 2015). Therefore, the sensitivity of this biotope is based on the abundance of Psolus squamatus and hard substrata, while other species groups are mentioned where relevant. 

Resilience and recovery rates of habitat

Psolus sp. are characterized by their exterior scales that form 'armour plating' around the body and their ventral sole that acts like a powerful sucking disk to attach to rocks and other hard substrata (Mortensen, 1927; Hyman,1955). The species of Psolidae studied are sexual (with one exception) (McEuen & Chia, 1991). The development of 15 species has been studied, of which 11 (73%) are brooders. Late larvae or juveniles are brooded on the upper surface of the adult in spaces under dorsal plates, under its adherent sole, amongst the crown of tentacles or internally, depending on the species (Boolootian, 1966; Young & Chia, 1982; reviewed by McEuen & Chia, 1991; Martinez et al., 2024). Brooding and lecithotrophic pelagic larvae are characteristic of the Psolids (McEuen & Chia, 1991).

Young & Chia (1982) and McEuen & Chia (1991) examined larval development and settlement in Psolus chitonoides from the San Juan Archipelago, Washington, USA. The eggs were bright red, 627 um in diameter and the maximum fecundity recorded was 32,700 in the laboratory. Spawning occurred in March and continued into May. In other species of Psolus, egg size ranged from 330 to 800 um. Spawning season varied between species and location but is probably annual (Boolotian, 1966; McEuen & Chia, 1991).  Young & Chia (1982) noted that spawning was initiated by males. Fertilized eggs developed first cleavage within two hours, the blastula stage (18 hours), gastrula (40 hours), doliolaria stage (75 hours), pentacula stage (9 days), settled after 12 days and the sole developed after 28 days in the laboratory. Larvae settled on any hard surfaces. In crowded conditions, larvae aggregated and metamorphosis was delayed up to eight months (McEuen & Chia, 1991). Juveniles supplied with detritus and algae fed from the bottom of dishes. Unfed juveniles survived for one year but were no longer than 1.2 mm. The development stages were similar to those reported in Psolus chitonoides by Young & Chia (1982) and in Psolus phantapus by Thorson (1946). McEuen & Chia (1991) noted that large psolids (7.5 to 15 cm long) tended to have pelagic larvae (e.g. Psolus chitonoides and Psolus phantapus)  while the smaller psolids (0.9 cm to 8.5 cm) were brooders (e.g. Psolus antarcticus and Psolus. patagonicus). 

Young & Chia (1982) reported that the eggs of Psolus chitonoides were spawned in strings that float to the surface. Larvae float until the late gastrula stage when they develop into ciliated larvae, the vitellaria. In the laboratory vitellaria were either swimming or resting on the bottom. These subsequently developed into pentacula which spent most of their time on the bottom, looking for places to settle (Young & Chia,1982).  While vitellaria varied in their response to light, pentacula were negatively phototactic and settled in the darkest areas. Both forms of larvae tended to aggregate in large numbers. In the laboratory, pentacula preferred to settle on adults rather than rock, while in the field, the larvae preferred adults or rock near adults. After settlement, 60-day old juveniles exclusively occupied dark areas. The eggs of other Psolus sp. may not be buoyant. For example, the eggs of Psolus patagonicus are clearly shown on rock next the adult (Martinez et al., 2024; Figure 14.3). 

Juveniles of Psolus chitonoides are capable of moving using ventral tube feet or, more rapidly, by walking on their tentacles up to 12.2 mm/hour (ca 25 body lengths/hour) while adults moved only one body length/hour. After a month, all juveniles had moved away from their settlement site. Adults were able to relocate to their preferred position within days after their boulder had been overturned. Small adults (<2 cm) were capable of moving more than 21 cm in three days while larger adults (>3 cm) moved only 10 cm. Six individuals moved less than 1 cm in two months in the field, except to rotate 30 degrees on the spot. Young & Chia (1982) concluded that adults in suitable living conditions were effectively sessile. They concluded that while juveniles preferred to settle in the vicinity of adults, their eventual distribution and aggregation was due to the common selection of suitable sites by multiple generations of juveniles. 

Psolus squamatus grows up to 6 to 7 cm in length but reaches sexual maturity at ca 3 cm (Mortensen, 1927). Ekman (1923) examined specimens of Psolus squamatus from southern Chile, Tierra del Fuego, and the Falkland Islands. He reported that an adult of 9.25 mm long carried 65 eggs while an adult of 6.5 mm in length carried only 10 eggs. Eggs were ca 0.6 mm in diameter. In two specimens from the Falklands, 18 juveniles (ca 1.8 mm long) were attached to the underside of the adult sole, together with two others of 2.5 and 2.7 mm long. The young attached to the mother's body using their suckers (Ekman, 1923). However, no information on the larval or juvenile development of Psolus squamatus was found. 

Psolus squamatus is widely distributed in the North Atlantic (from Svalbard, Iceland, south through the North Sea, into the Kattegat, and southern England, and Labrador south to the Bay of Fundy) and western Pacific from the Bering Sea, south to Mexico. A few records occur off the coasts of South Africa, Chile, Argentina and the Falkland Islands (OBIS, 2024).  It is recorded from 60 m to ca 1000 m in depth, although most records are from 180 m to ca 1000 m (OBIS, 2024). However, Psolus squamatus is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight, characterized by egg sizes indicative of lecithotrophic development and near sea bed development (Billett, 1991; Wagstaff et al., 2014). Billett (1991) suggested that slope holothurians (such as Psolus) might be expected to have short development close to the sea bed. Billett (1991) noted that near-bed currents generally run parallel to the slope contours and could keep larvae close to suitable habitats. However, Billett (1991) noted that information on the population dynamics of deep-sea holothurians, growth rates or longevity was lacking. 

Resilience assessment. No evidence of the population dynamics, growth rates, mortality rates or longevity of Psolus squamatus and its populations was found. Based on other Psolus sp., Psolus squamatus probably produces short-lived lecithrophic larvae, that aggregate near adults and adults brood either larvae or juveniles until they are large enough to avoid predation. Therefore, they probably have good localized recruitment via larvae or juveniles, with the possibility of more distant dispersal of larvae via near-bed currents.  Assuming annual recruitment, the Psolus squamatus population might be expected to take a few years to recover from disturbance, or possibly 2-10 years ('Medium' resilience) to recover from mortality. However, the assessment is made with 'Low' confidence due to the lack of direct evidence. Recruitment to available hard substrata by epifauna such as serpulids, anomiids and ascidians is probably fairly rapid with sponges and soft corals taking longer to develop (Sebens, 1985, 1986). Bryozoans, hydroids, serpulids and ascidians are opportunistic, grow and colonize space rapidly and will probably develop an epifaunal cover within 1-2 years (for example see Sebens, 1985, 1986). Mobile epifauna and infauna will probably colonize rapidly from the surrounding area. Slow-growing species such as some sponges and anemones (see Sebens, 1985, 1986), will probably take several years to develop significant cover, so the community may also take 2 to 10 years to develop, depending on local conditions. Large globose sponges and solitary cold-water corals would probably take longer to recruit and grow. 

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. Further detail

Evidence

Psolus squamatus is widely distributed in the North Atlantic (from Svalbard, Iceland, south through the North Sea, into the Kattegat, and southern England, and Labrador south to the Bay of Fundy) and western Pacific from the Bering Sea, south to Mexico. A few records occur off the coasts of South Africa, Chile, Argentina and the Falkland Islands (OBIS, 2024). The majority of records have sea surface temperatures of ca 10 to 15°C (OBIS, 2024). However, sea surface temperatures are not relevant to deep water species.

For example, in the Rockall Trough, the deep water temperature and salinity are determined by the ocean currents, such as the Eastern North Atlantic Water, Wyville Thompson Ridge Overflow Water, the Labrador Sea Water and the Antarctic Bottom Water (Gage, 1986; Sherwin et al., 2012). Sherwin et al. (2012) reported that the temperature of the seawater in Rockall Trough was 10°C at ca 500 m and dropped to 5°C at ca 1,500 m (in October 2006). In addition, the North East Atlantic exhibits seasonal thermoclines and winter mixing but a permanent thermocline at ca 500 m (Tyler & Young, 1998). In the Rockall Trough, the seasonal thermocline develops at ca 200 m and winter mixing occurs to about 600 m, while the permanent thermocline extends from ca 800 m to ca 1,000 m (Gage, 1986). 

Psolus squamatus is recorded from 60 m to ca 1000 m in depth, although most records are from 180 m to ca 1000 m (OBIS, 2024) and is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight (Billett, 1991; Wagstaff et al., 2014). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf. The transition at about 1000 m is influenced by decreased cooling from the permanent thermocline and decreased food influx and sees the loss of most upper boundary species (Carney, 2005). 

Sensitivity assessment. No information on the temperature tolerances of adult or larval Psolus squamatus was found. Examples of this biotope were reported from ca 330 m to ca 1,345 m in UK waters (Weinberg et al., 2008; Roberts et al., 2008; Howell et al., 2010; Davies et al., 2015) and a similar biotope was reported at 170 to 370 m on the Oregon continental shelf (Hemery et al., 2018). On the Anton Dohr seamount the biotopes occurred from 4.3 to 7.9°C depending on location (Davies et al., 2015) and at a range of 7 to 10°C at several UK locations (Howell et al., 2010). At depth, and especially below the permanent thermocline, temperatures are likely to be stable and organisms are unlikely to be exposed to the range of temperatures and, in particular, the rapidity of temperature change experienced at the sea surface. Sherwin et al. (2012) reported that the seawater temperature of the upper 800 m of the Rockall Trough had fluctuated between ca 9.0 and 10.5°C from 1948 to 2010. While larvae may be tolerant of a range of temperatures, adults may be more stenothermal, but no direct evidence was found. Hence, while natural temperature changes are unlikely, exposure to localised thermal effluents at the benchmark level (e.g. from deep-sea installations or operations, however unlikely) may be detrimental. Therefore, resistance is assessed as 'Medium' as a precaution, albeit with 'Low' confidence. Resilience is probably 'Medium' so sensitivity is assessed as 'Medium'

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

Temperature decrease (local)

Benchmark. A 5°C decrease in temperature for one month, or 2°C for one year. Further detail

Evidence

Psolus squamatus is widely distributed in the North Atlantic (from Svalbard, Iceland, south through the North Sea, into the Kattegat, and southern England, and Labrador south to the Bay of Fundy) and western Pacific from the Bering Sea, south to Mexico. A few records occur off the coasts of South Africa, Chile, Argentina and the Falkland Islands (OBIS, 2024). The majority of records have sea surface temperatures of ca 10 to 15°C (OBIS, 2024). However, sea surface temperatures are not relevant to deep water species.

For example, in the Rockall Trough, the deep water temperature and salinity are determined by the ocean currents, such as the Eastern North Atlantic Water, Wyville Thompson Ridge Overflow Water, the Labrador Sea Water and the Antarctic Bottom Water (Gage, 1986; Sherwin et al., 2012). Sherwin et al. (2012) reported that the temperature of the seawater in Rockall Trough was 10°C at ca 500 m and dropped to 5°C at ca 1,500 m (in October 2006). In addition, the North East Atlantic exhibits seasonal thermoclines and winter mixing but a permanent thermocline at ca 500 m (Tyler & Young, 1998). In the Rockall Trough, the seasonal thermocline develops at ca 200 m and winter mixing occurs to about 600 m, while the permanent thermocline extends from ca 800 m to ca 1,000 m (Gage, 1986). 

Psolus squamatus is recorded from 60 m to ca 1000 m in depth, although most records are from 180 m to ca 1000 m (OBIS, 2024) and is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight (Billett, 1991; Wagstaff et al., 2014). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf. The transition at about 1000 m is influenced by decreased cooling from the permanent thermocline and decreased food influx and sees the loss of most upper boundary species (Carney, 2005). 

Sensitivity assessment. No information on the temperature tolerances of adult or larval Psolus squamatus was found. Examples of this biotope were reported from ca 330 m to ca 1,345 m in UK waters (Weinberg et al., 2008; Roberts et al., 2008; Howell et al., 2010; Davies et al., 2015) and a similar biotope was reported at 170 to 370 m on the Oregon continental shelf (Hemery et al., 2018). On the Anton Dohr seamount the biotopes occurred from 4.3 to 7.9°C depending on location (Davies et al., 2015) and at a range of 7 to 10°C at several UK locations (Howell et al., 2010). At depth, and especially below the permanent thermocline, temperatures are likely to be stable and organisms are unlikely to be exposed to the range of temperatures and, in particular, the rapidity of temperature change experienced at the sea surface. Sherwin et al. (2012) reported that the seawater temperature of the upper 800 m of the Rockall Trough had fluctuated between ca 9.0 and 10.5°C from 1948 to 2010. While larvae may be tolerant of a range of temperatures, adults may be more stenothermal, but no direct evidence was found. Hence, while natural temperature changes are unlikely, exposure to localised thermal effluents at the benchmark level (e.g. from deep-sea installations or operations, however unlikely) may be detrimental. Therefore, resistance is assessed as 'Medium' as a precaution, albeit with 'Low' confidence. Resilience is probably 'Medium' so sensitivity is assessed as 'Medium'

Medium
Low
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Medium
Low
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Medium
Low
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Salinity increase (local) [Show more]

Salinity increase (local)

Benchmark. A increase in one MNCR salinity category above the usual range of the biotope or habitat. Further detail

Evidence

This biotope is characterized by the sea cucumber Psolus squamatus. Echinoderms are osmoconformers and generally stenohaline due to their lack of an excretory organ, and their poor ability to osmoregulate (Binyon, 1966; Stickle & Diehl, 1987), while several species are recorded from extreme salinities (Stickle & Diehl, 1987; Russell, 2013). However, no information on the salinity tolerance of the characteristic species was found.  

Roberts et al. (2010b) suggested that hypersaline effluent dispersed quickly but was more of a concern at the seabed and in areas of low energy where widespread alternations in the community of soft sediments were observed. In several studies, echinoderms and ascidians were amongst the most sensitive groups examined (Roberts et al., 2010b). Fernández-Torquemada et al. (2013) suggested that echinoderms could be a useful early bioindicator for the effects of increased salinity.  In the Mediterranean, echinoderms were absent within one year in the areas affected by hypersaline effluent from a desalination plant but returned after dilution of the discharge with seawater (Fernández-Torquemada et al., 2013).

However, seawater salinity in the deep sea is more stable than inshore waters. For example, in the Rockall Trough, the deep water temperature and salinity are determined by ocean currents, such as the Eastern North Atlantic Water, Wyville Thompson Ridge Overflow Water, the Labrador Sea Water and the Antarctic Bottom Water (Gage, 1986; Sherwin et al., 2012). Sherwin et al. (2012) reported that the seawater salinity of the upper 800 m of the Rockall Trough had fluctuated between ca 35.25 and 35.45 from 1948 to 2010. Sherwin et al. (2012) reported that the salinity of the seawater in Rockall Trough was 35.4 at ca 500 m and dropped to 35.15 at ca 1,500 m (in October 2006).

Sensitivity assessment. The species that dominate this biotope (e.g. Psolus squamatus, sponges, and serpulid) are probably adapted to stable salinity conditions and have limited tolerance to salinity change. An increase in salinity from full to >40 psu is probably detrimental to the important characteristic species of the biotope. However, it is unlikely that this biotope would be exposed to hypersaline conditions (or effluent) unless from a newly opened brine seep or an unknown deep-sea operation. Although there is no direct evidence of the effects of hypersaline water on the characteristic species, the stenohaline nature of the echinoderms suggests that hypersaline conditions may cause mortality. Therefore, resistance is assessed as 'Low' but at Low confidence. Resilience would probably be 'Low' so sensitivity is assessed as 'High'.

Low
Low
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Low
Low
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High
Low
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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. Further detail

Evidence

This biotope is characterized by the sea cucumber Psolus squamatus. Echinoderms are osmoconformers and generally stenohaline due to their lack of an excretory organ, and their poor ability to osmoregulate (Binyon, 1966; Stickle & Diehl, 1987), while several species are recorded from extreme salinities (Stickle & Diehl, 1987; Russell, 2013). However, no information on the salinity tolerance of the characteristic species was found.  

However, seawater salinity in the deep sea is more stable than inshore waters. For example, in the Rockall Trough, the deep water temperature and salinity are determined by ocean currents, such as the Eastern North Atlantic Water, Wyville Thompson Ridge Overflow Water, the Labrador Sea Water and the Antarctic Bottom Water (Gage, 1986; Sherwin et al., 2012). Sherwin et al. (2012) reported that the seawater salinity of the upper 800 m of the Rockall Trough had fluctuated between ca 35.25 and 35.45 from 1948 to 2010. Sherwin et al. (2012) reported that the salinity of the seawater in Rockall Trough was 35.4 at ca 500 m and dropped to 35.15 at ca 1,500 m (in October 2006). 

Sensitivity assessment. The species that dominate this biotope (e.g. Psolus squamatus, sponges, and serpulid) are probably adapted to stable salinity conditions and have limited tolerance to salinity change. An increase in salinity from full to >40 psu is probably detrimental to the important characteristic species of the biotope. However, it is unlikely that this biotope would be exposed to hypersaline conditions (or effluent) unless from a newly opened brine seep or an unknown deep-sea operation. Although there is no direct evidence of the effects of hypersaline water on the characteristic species, the stenohaline nature of the echinoderms suggests that hyposaline conditions may cause mortality. Therefore, resistance is assessed as 'Low' but at Low confidence. Resilience would probably be 'Low' so sensitivity is assessed as 'High'.

Low
Low
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Low
Low
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High
Low
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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 to 0.2 m/s for more than one year. Further detail

Evidence

Psolus squamatus and the other characteristic species are suspension feeders, dependent on current flow to transport or resuspend organic particulates or phytodetritus. Psolus squamatus is recorded from 60 m to ca 1000 m in depth, although most records are from 180 m to ca 1000 m (OBIS, 2024) and is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight (Billett, 1991; Wagstaff et al., 2014). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf. The transition at about 1000 m is influenced by decreased cooling from the permanent thermocline and decreased food influx and sees the loss of most upper boundary species (Carney, 2005). 

Deep sea habitats are probably dominated by mass water transport due to oceanic currents. For example, water flow in the Rockall Trough is dominated by the Eastern North Atlantic Water, Wyville Thompson Ridge Overflow Water, the Labrador Sea Water and the Antarctic Bottom Water (Gage, 1986; Sherwin et al., 2012), except near Feni Ridge or Anthon Dohrn seamount which are also influenced by tidal oscillation. Gage (1986) reported Psolus squamatus from the top of the current swept Anton Dohr Seamount and Psolus sp. from Ferni Ridge. Gage (1986) reported maximum flow rates of ca 0.5 m/s within 150 m of the bottom on the Feni Ridge west of the Anthon Dohrn seamount, mostly due to tidal oscillation, and associated with current-moulded bedforms (e.g. ripples). However, lower tidal currents <0.05 m/s with a maximum of ca 0.21 m/s were recorded within 400-500 m of the bottom elsewhere. Wienberg et al. (2008) described assemblages of Psolus sp., serpulids and bryozoans on gravel to large boulder-sized fields of 'dropstones', mainly on soft sediment on the flanks of the Franken Mound, Rockall Bank. Wienberg et al. (2008) noted that the eastern flank of Franken Mound was subject to strong but variable bottom currents, indicated by extensive areas of soft sediment, exhumed dropstones, rocky outcrops, and mobile rippled sediment, characteristic of active sediment transport. Flow speeds in the Rockall Trough are typically 0.15 to 0.3 m/s (Wienberg et al., 2008) but localised flow over mounds, sea mount and other physical features probably affect local currents. 

Sensitivity assessment. These biotopes are characterized by hard substrata (rocky outcrops, boulders, pebbles and gravel) on coarse or mobile sediments. A localized change in water flow of 0.1 to 0.2 m/s (the benchmark) might reduce the suitability of the habitat for the suspension feeders but also might be insignificant in the "strong but variable currents" suggested by Wienberg et al. (2008). The benchmark level of change lies within the range of water flow velocities recorded in parts of the Rockall Trough but no information on flow rates at the seabed was available. Therefore, there is insufficient evidence on which to base an assessment. 

Insufficient evidence (IEv)
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Not relevant (NR)
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Insufficient evidence (IEv)
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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. Further detail

Evidence

This deep-sea biotope is not exposed to changes in emersion. 

Not relevant (NR)
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Not relevant (NR)
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Not relevant (NR)
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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. Further detail

Evidence

These deep-sea biotopes are recorded from the upper and mid bathyal and will not be affected by changes in nearshore wave exposure.

Not relevant (NR)
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Not relevant (NR)
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Not relevant (NR)
NR
NR
NR
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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 contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

Bryan (1984) reported that early work had shown that echinoderm larvae were sensitive to heavy metals, e.g. the intolerance of larvae of Paracentrotus lividus to copper (Cu) had been used to develop a water quality assessment. Kinne (1984) reported developmental disturbances in Echinus esculentus exposed to waters containing 25 µg / l of copper (Cu) and heavy metals caused reproductive anomalies in the starfish Asterias rubens (Besten, et al., 1989, 1991). Sea urchins, especially the eggs and larvae, are used for toxicity testing and environmental monitoring (reviewed by Dinnel et al. 1988). Crompton (1997) reported that mortalities occurred in echinoderms after 4-14 day exposure to above 10-100 µg/l Cu, 1-10 mg/l Zn and 10-100 mg/l Cr but that mortalities occurred in echinoderm larvae above10-100 µg/ l Ni.

Sensitivity assessment. No evidence of the effects of transitional metals or organometal on the characteristic sea cucumbers or other species was found. However, evidence from other echinoderms suggests that the dominant sea cucumber in this biotope is also likely to be adversely affected by transitional metals. Therefore, resistance is assessed as 'Low', so resilience is probably 'Medium' and sensitivity is assessed as 'Medium', albeit with 'Low' confidence due to lack of directly relevant evidence. Further evidence is required for this pressure.

Low
Low
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Medium
Low
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NR
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Medium
Low
NR
NR
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Hydrocarbon & PAH contamination [Show more]

Hydrocarbon & PAH contamination

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

Sea urchins, especially the eggs and larvae are used for toxicity testing and environmental monitoring (reviewed by Dinnel et al. 1988). Therefore, echinoderms, especially their larvae, may be sensitive to a range of contaminants. Echinoderms seem especially intolerant of the toxic effects of oil, likely because of the large amount of exposed epidermis (Suchanek, 1993).

Sea urchin eggs showed developmental abnormalities when exposed to 10-30 mg/l of hydrocarbons and crude oil: Corexit dispersant mixtures have been shown to cause functional loss of tube feet and spines in sea urchins (Suchanek, 1993). Olsgard & Gray (1995) found the brittlestar Amphiura filiformis very sensitive to oil pollution. During monitoring of sediments in the Ekofisk oilfield, Addy et al. (1978) suggest that reduced abundance of Amphiura filiformis within 2-3 km of the site was related to discharges of oil from the platforms and to physical disturbance of the sediment. Although acute toxicity tests showed that drill cuttings containing oil-based muds had a very low toxicity (LC50 52,800 ppm total hydrocarbons in test sediment), Newton & McKenzie (1998) suggest these are poor predictors of chronic response. Chronic sub-lethal effects were detected around the Beryl oil platform in the North Sea where the levels of oil in the sediment were very low (3 ppm) and Amphiura filiformis was excluded from areas nearer the platform with higher sediment oil content. Similarly, in Ophiothrix fragilis, exposure to 30,000 ppm oil reduces its load of symbiotic bacteria by 50% and brittle stars begin to die (Newton & McKenzie, 1995).

Crude oil from the Torrey Canyon and the detergent used to disperse it caused mass mortalities of echinoderms; Asterias rubens, Echinocardium cordatum, Psammechinus miliaris, Echinus esculentus, Marthasterias glacialis and Acrocnida brachiata (Smith, 1968). Large numbers of dead Echinus esculentus were found between 5.5 and 14.5 m in the vicinity of Sennen after the Torrey Canyon oil spill, presumably due to a combination of wave exposure and heavy spraying of dispersants in that area (Smith 1968). Smith (1968) also demonstrated that 0.5 to 1 ppm of the detergent BP1002 resulted in developmental abnormalities in echinopluteus larvae of Echinus esculentusEchinus 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). However, the observed effects may have been due to a single contaminant or synergistic effects of all present.

A study by Brils et al. (2002) into the toxicity of C10-19 hydrocarbons found that the shallow irregular echinoid Echinocardium cordatum was susceptible to oil-contaminated sediments at as low as 190 mg/kg dry weight of Echinocardium cordatum. The high intolerance of Echinocardium cordatum to hydrocarbons was seen by the mass mortality of animals, down to about 20 m, shortly after the Amoco Cadiz oil spill (Cabioch et al., 1978). Reduced abundance of the species was also detectable up to >1,000 m away one year after the discharge of oil-contaminated drill cuttings in the North Sea (Daan & Mulder, 1996).  The polyaromatic hydrocarbon (PAH) fluoranthene was shown to cause mortality in larvae of Arbacia punctulata (purple-spined sea urchin) with 48-hour LC50 of 3.9 µg/l in the presence of UV light (Spehar et al., 1999). 

TBT was shown to inhibit arm regeneration in the brittlestar Ophioderma brevispina, at 10 ng/l and produce significant inhibition at 100 ng/l. It was suggested that TBT acts via the nervous system, although direct action on the tissues at the point of breakage could not be excluded (Bryan & Gibbs, 1991)

Sensitivity assessment. No evidence of the effects of hydrocarbon or PAH contamination on the characteristic echinoderms was found. However, evidence from other echinoderms, suggests that the dominant sea cucumbers in this biotope are also likely to be adversely affected by hydrocarbon or PAH exposure. Therefore, resistance is assessed as 'Low', so resilience is probably 'Medium' and sensitivity is assessed as 'Medium', albeit with 'Low' confidence due to lack of directly relevant evidence. Further evidence is required for this pressure.

Low
Low
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NR
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Medium
Low
NR
NR
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Medium
Low
NR
NR
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Synthetic compound contamination [Show more]

Synthetic compound contamination

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

Little information on the toxicity of synthetic chemicals to holothurians was found. Newton & McKenzie (1995) suggested that echinoderms tend to be very intolerant of various types of marine pollution but gave no detailed information. Cole et al. (1999) reported that echinoderm larvae displayed adverse effects when exposed to 0.15 mg/l of the pesticide Dichlorobenzene (DCB). Smith (1968) demonstrated that 0.5 -1 ppm of the detergent BP1002 resulted in developmental abnormalities in echinopluteus larvae of Echinus esculentus. Therefore, holothurians and their larvae may also be intolerant of synthetic chemicals.

Sensitivity assessment. Little evidence of the effects of synthetic contamination on the characteristic urchins was found. There is 'Insufficient evidence' above to support an assessment and further evidence is required for this pressure.

Insufficient evidence (IEv)
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Not relevant (NR)
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Insufficient evidence (IEv)
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Radionuclide contamination [Show more]

Radionuclide contamination

Benchmark. An increase in 10µGy/h above background levels. Further detail

Evidence

No evidence was found.

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

Introduction of other substances

Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

Evidence

No evidence was found.

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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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). Further detail

Evidence

Lawrence (1996) reported mass mortality of echinoderms in the Gulf of Trieste due to hypoxia caused by a strong thermocline combined with high pelagic productivity and eutrophication. The brittlestar Ophiura quinquemaculata was killed within a few days, and holothurians including Ocnus planci (as Cucumaria planci), starfish Asteropecten sp. and the remaining brittlestars were killed within a week. In experiments, Amphiura filiformis only left its protected position in the sediment when oxygen levels fell below 0.85mg/l (Rosenberg et al., 1991). Mass mortality of Amphiura filiformis was observed during severely low oxygen events (<0.7 mg/l) (Nilsson, 1999). However, at oxygen concentrations between 0.85 mg/l and 1.0 mg/l Rosenberg et al. (1991) observed the species survived for several weeks. Echinoderms were shown to be intolerant of the effects of algal blooms, resulting in mortalities of the sea urchins Echinus esculentus and Paracentrotus lividus, and the holothurian Labidoplax digitata amongst other echinoderms, probably due to hypoxia caused by death of the algal bloom algae (Boalch, 1979; Forster, 1979; Griffiths et al., 1979; Lawrence, 1996). Vaquer-Sunyer & Duarte (2008) suggested a median sublethal oxygen concentration of 1.22 mg O2/l (± 0.25) for a number of echinoderms reviewed in their study. Echinoderms were neither the most nor the least sensitive of the taxonomic groups examined.  Riedel et al. (2012) examined the effects of hypoxia and anoxia on macrofauna in the North Adriatic, using in situ experimental apparatus.  They concluded that decapods, echinoderms and polychaetes were among the most sensitive to hypoxia while ascidians and anthozoans were among the most resistant of the species encountered in their study. 

Sensitivity assessment. Overall, the above evidence suggests echinoderms are intolerant of hypoxic conditions. However, In the deep sea oxygen levels are probably determined by mass transport of water by deep currents. For example, Ellett & Martin (1973) reported that the dissolved oxygen levels in the Rockall Trough varied between ca 5 and 6 ml/l (ca 7 and 8.4 mg/l) between the surface and ca 2,000 m in depth. In addition, the short-term acute hypoxia, represented by the benchmark, may also be mitigated by the large water masses and strong currents typical of areas in which these biotopes occur. Therefore, resistance is assessed as ‘Medium’ to represent some mortality under the worst-case scenario. Hence, resilience is assessed as ‘Medium' and sensitivity as 'Medium' but with 'Low' confidence. 

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

Nutrient enrichment

Benchmark. Compliance with WFD criteria for good status. Further detail

Evidence

Psolus squamatus is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight (Billett, 1991; Wagstaff et al., 2014). This zonation is typical of marine invertebrates in deep-sea habitats and corresponds to the upper boundary biota identified by Carney (2005). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf.  These biotopes are probably dependent on seasonal phytodetritus and suspended organic particulates. For example, Maier et al. (2023) concluded that local hydrography produces periodic pulses of food due to internal waves operating on seasonal, multi-year, decadal or millennial cycles, with currents that interact with the deep-sea topography (such as sea mounts, continental shelf margins, fjord sills) or cold-water reefs to form internal waves, hydraulics jumps and trapped waves. The resultant downwelling can rapidly transport surface productivity (such as plankton or POM) to the reef (Maier et al., 2023). For example, fresh organic matter can be transported from the surface in less than one hour to 140 m on Mingulay Reef (reviewed by Maier et al., 2023). Internal waves also resuspend deposited organic matter into the bottom or intermediate layers (Maier et al., 2023). Similarly, the nutrient levels (e.g. nitrates, phosphates, and ammonia) and inorganic carbon in the vicinity of cold-water coral reefs in the North East Atlantic vary with the tidal cycle and depth (Findlay et al., 2014). 

However, no evidence of the nutrient levels typical of these biotopes was found and no information on the effect of nutrient enrichment on the characteristic species was found. Therefore, 'No evidence' is recorded. 

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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Organic enrichment [Show more]

Organic enrichment

Benchmark. A deposit of 100 gC/m2/yr. Further detail

Evidence

Psolus squamatus is typical of the upper slope (200 m to 1000 m) holothurian fauna in the Porcupine Seabight (Billett, 1991; Wagstaff et al., 2014). This zonation is typical of marine invertebrates in deep-sea habitats and corresponds to the upper boundary biota identified by Carney (2005). Carney (2005) suggested that the upper slope from the shelf break to ca 500 m is an area of vertical food influx and possibly upwelling (depending on location) or influx from the adjacent shelf.  These biotopes are probably dependent on seasonal phytodetritus and suspended organic particulates. For example, Maier et al. (2023) concluded that local hydrography produces periodic pulses of food due to internal waves operating on seasonal, multi-year, decadal or millennial cycles, with currents that interact with the deep-sea topography (such as sea mounts, continental shelf margins, fjord sills) or cold-water reefs to form internal waves, hydraulics jumps and trapped waves. The resultant downwelling can rapidly transport surface productivity (such as plankton or POM) to the reef (Maier et al., 2023). For example, fresh organic matter can be transported from the surface in less than one hour to 140 m on Mingulay Reef (reviewed by Maier et al., 2023). Internal waves also resuspend deposited organic matter into the bottom or intermediate layers (Maier et al., 2023). Similarly, the nutrient levels (e.g. nitrates, phosphates, and ammonia) and inorganic carbon in the vicinity of cold-water coral reefs in the North East Atlantic vary with the tidal cycle and depth (Findlay et al., 2014). 

However, no evidence of the organic carbon levels typical of these biotopes was found and no information on the effect of organic enrichment on the characteristic species was found. Therefore, 'No evidence' is recorded. 

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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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. Further detail

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 available habitat (resilience is ‘Very low’). Therefore, the biotopes are considered to have ‘High’ sensitivity to this pressure.

None
High
High
High
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Very Low
High
High
High
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High
High
High
High
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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. Further detail

Evidence

The loss of hard substrata and its replacement with sediment would cause the biotope to be lost. In addition, the mechanical removal of hard substratum would destroy any characterizing organisms present and ultimately result in the loss and reclassification of the biotope. Therefore, resistance is assessed as 'None'. As this pressure is considered a permanent change, resilience is assessed as 'Very Low', and sensitivity is assessed as 'High'.

None
High
High
High
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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). Further detail

Evidence

The loss of hard and mixed substrata and its replacement with sediment alone would cause the biotope to be lost. In addition, the mechanical removal of hard substratum would destroy any characterizing organisms present and ultimately result in the loss and reclassification of the biotope. Therefore, resistance is assessed as 'None'. As this pressure is considered a permanent change, resilience is assessed as 'Very Low', and sensitivity is assessed as 'High'.

None
High
High
High
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Very Low
High
High
High
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High
High
High
High
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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). Further detail

Evidence

The important characteristic epifauna (Psolus squamatus) position themselves on stones, pebbles, shells, and other hard substrata to avail themselves of the passing current. the other suspension-feeding brittlestars may do the same. The extraction of sediment to a depth of 30 cm would remove the infauna, surface hard substrata such as dropstones, cobbles and pebbles, remove or relocate boulders and, hence,  the characteristics surface epifauna of sea cucumbers, ophiuroids, sponges, serpulids, anomiids in the affected area.  Therefore, a resistance of 'None' is suggested based on expert judgment. Resilience is probably 'Medium' so the sensitivity is assessed as 'Medium'

None
Low
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Medium
Low
NR
NR
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Medium
Low
NR
NR
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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). Further detail

Evidence

Erect epifaunal species are particularly vulnerable to physical disturbance (Jennings & Kaiser, 1998). Veale et al. (2000) reported that the abundance, biomass, and production of epifaunal assemblages decreased with increasing fishing effort. Mobile gears also result in modification of the substratum, including the removal of shell debris, cobbles and rocks, and the movement of boulders (Bullimore, 1985; Jennings & Kaiser, 1998) on which many of the species in this community depend. Picton & Goodwin (2007) noted that an area of boulders with a rich fauna of sponges and hydroids on the east coast of Rathlin Island, Northern Ireland was significantly altered since the 1980s.  Scallop dredging had begun in 1989 and boulders were observed to have been turned and the gravel harrowed. In addition, many of the boulders had disappeared and rare hydroid communities were greatly reduced (Picton & Goodwin, 2007). Prior records indicated the presence of large sponges, mainly Axinella infundibuliformis (Picton & Goodwin, 2007). Freese (2001) studied deep cold-water sponges in Alaska a year after a trawl event;  46.8% of sponges exhibited damage with 32.1% having been torn loose.  None of the damaged sponges displayed signs of regrowth or recovery.  This was in stark contrast to early work by Freese et al. (1999) on shallow sponge communities, with impacts of trawling activity being much more persistent due to the slower growth/regeneration rates of deep, cold-water sponges. 

Sensitivity assessment. No evidence of the effects of fishing activities on these deep-sea biotopes was found. However, the above evidence from shallow sea demonstrates that bottom trawling can relocate and turn boulders, and relocate or remove cobbles, pebbles and presumably 'dropstones' that are required for the development of this community. The resultant abrasion and disturbance may kill or break up sponges, in particular, although it is unclear if Psolus squamatus would survive or be displaced. Nevertheless, the removal of hard substratum (boulders, pebbles etc.) would result in the loss of suitable habitat. Therefore, resistance is assessed as 'Low'. Resilience is probably 'Medium', so sensitivity is assessed as 'Medium'

Low
Low
NR
NR
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Medium
Low
NR
NR
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Medium
Low
NR
NR
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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). Further detail

Evidence

The effects of penetrative fishing gear on the biotope are probably at least as severe as surface abrasion above.  Therefore, resistance is assessed as 'Low'. Resilience is probably 'Medium', so sensitivity is assessed as 'Medium'

Low
Low
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Medium
Low
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Medium
Low
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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. Further detail

Evidence

The SpaEnc.PsoSpo and SpaEnc.PsoAno biotopes are defined by the abundance of Psolus squamatus and hard substrata (bedrock, boulders, and cobbles) and sparse encrusting species; sponges in PsoSpo and saddle oysters and serpulids in PsoAno (JNCC, 2022). No information on the ecology of this habitat was found. However, the lack of long-lived species on the available hard substrata suggests the current flow does not support large sponge or cold-water corals and that the habitat is exposed to periodic disturbance, probably due to sediment mobility, scour and burial. The habitat is characterized by exposed rock outcrops amongst sediment, or mixed hard substratum (boulders, pebbles, and gravel etc.) on the surface of potentially mobile sediment (Wienberg et al., 2008; Roberts et al., 2008; Davis et al., 2015). 

Sensitivity assessment. The characteristic species are suspension feeders so a decrease in suspended particulates could be detrimental but possibly seasonal (see Maier et al., 2023). However, the physical nature of the habitat suggests that it could be subject to periodic sediment resuspension, mobility and burial.  Therefore, resistance is assessed as 'Medium' assuming that a reduction in suspended particulates may reduce the food supply for the characteristic species but with 'Low' confidence due to the lack of evidence. Hence, resilience is assessed as 'Medium' and sensitivity as 'Medium'. 

Medium
Low
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NR
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Medium
Low
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NR
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Medium
Low
NR
NR
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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. Further detail

Evidence

The SpaEnc.PsoSpo and SpaEnc.PsoAno biotopes are defined by the abundance of Psolus squamatus and hard substrata (bedrock, boulders, and cobbles) and sparse encrusting species; sponges in PsoSpo and saddle oysters and serpulids in PsoAno (JNCC, 2022). No information on the ecology of this habitat was found. However, the lack of long-lived species on the available hard substrata suggests the current flow does not support large sponge or cold-water corals and that the habitat is exposed to periodic disturbance, probably due to sediment mobility, scour and burial. The habitat is characterized by exposed rock outcrops amongst sediment, or mixed hard substratum (boulders, pebbles, and gravel etc.) on the surface of potentially mobile sediment (Wienberg et al., 2008; Roberts et al., 2008; Davis et al., 2015). 

Sensitivity assessment. The physical nature of the habitat suggests that it could be subject to periodic sediment resuspension, mobility and burial.  In areas of strong current flow, burial may be short-term. If this assumption is correct, and the habitat is structured by periodic smothering, then resistance is assessed as 'High' but with 'Low' confidence due to the lack of evidence. Hence, resilience is assessed as 'High' and sensitivity as 'Not sensitive'. 

High
Low
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NR
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High
High
High
High
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Not sensitive
Low
NR
NR
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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. Further detail

Evidence

The SpaEnc.PsoSpo and SpaEnc.PsoAno biotopes are defined by the abundance of Psolus squamatus and hard substrata (bedrock, boulders, and cobbles) and sparse encrusting species; sponges in PsoSpo and saddle oysters and serpulids in PsoAno (JNCC, 2022). No information on the ecology of this habitat was found. However, the lack of long-lived species on the available hard substrata suggests the current flow does not support large sponge or cold-water corals and that the habitat is exposed to periodic disturbance, probably due to sediment mobility, scour and burial. The habitat is characterized by exposed rock outcrops amongst sediment, or mixed hard substratum (boulders, pebbles, and gravel etc.) on the surface of potentially mobile sediment (Wienberg et al., 2008; Roberts et al., 2008; Davis et al., 2015). 

Sensitivity assessment. The physical nature of the habitat suggests that it could be subject to periodic sediment resuspension, mobility and burial.  In areas of strong current flow, burial may be short-term. If this assumption is correct, and the habitat is structured by periodic smothering, then resistance is assessed as 'High' but with 'Low' confidence due to the lack of evidence. Hence, resilience is assessed as 'High' and sensitivity as 'Not sensitive'. 

High
Low
NR
NR
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High
High
High
High
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Not sensitive
Low
NR
NR
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Litter [Show more]

Litter

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

Evidence

No evidence could be found regarding the introduction of litter on this biotope

No evidence (NEv)
NR
NR
NR
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Not relevant (NR)
NR
NR
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No evidence (NEv)
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Electromagnetic changes [Show more]

Electromagnetic changes

Benchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT. Further detail

Evidence

No evidence was found.

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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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

This biotope is characterized by invertebrates with no known means to detect noise, which will not be affected by changes in underwater noise, as defined under this pressure.

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

Introduction of light or shading

Benchmark. A change in incident light via anthropogenic means. Further detail

Evidence

This biotope is characterized by invertebrates with limited ability to detect light and is aphotic.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
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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. Further detail

Evidence

The benthic lifestyle and widespread deep-sea habitat of the characteristic species mean they will not be affected by barriers as defined under this pressure. Physical and hydrographic barriers may limit the dispersal of larvae but larval dispersal is not considered under the pressure definition and benchmark.

Not relevant (NR)
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NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
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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. Further detail

Evidence

This biotope is characterized by benthic invertebrates that are not at risk of collision with artificial structures. It might be affected adversely by falling marine debris such as barrels, containers, and even shipwrecks but the effects are probably addressed under 'abrasion' above. 

Not relevant (NR)
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NR
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Not relevant (NR)
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Not relevant (NR)
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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. Further detail

Evidence

This biotope is characterized by invertebrates that do not rely on visual cues and will not be affected by visual disturbance, as defined under this pressure.

Not relevant (NR)
NR
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Not relevant (NR)
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NR
NR
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Not relevant (NR)
NR
NR
NR
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Biological Pressures

Use [show more] / [show less] to open/close text displayed

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 that may result in changes in the genetic structure of local populations, hybridization, or change in community structure. Further detail

Evidence

No evidence was found to suggest that any of the characteristic species were subject to translocation or genetic modification, nor the introduction of genetically distinct organisms. 

Not relevant (NR)
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Not relevant (NR)
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Not relevant (NR)
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Introduction or spread of invasive non-indigenous species [Show more]

Introduction or spread of invasive non-indigenous species

Benchmark. The introduction of one or more invasive non-indigenous species (INIS). Further detail

Evidence

No information on the effect of the introduction of one or more invasive non-indigenous species on this biotope was found.

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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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). Further detail

Evidence

No information on the effect of pathogens or disease on the characterizing species was found.

No evidence (NEv)
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Not relevant (NR)
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No evidence (NEv)
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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. Further detail

Evidence

The characterizing species of these biotopes are not targeted by commercial or recreational fisheries.

Not relevant (NR)
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Not relevant (NR)
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Not relevant (NR)
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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. Further detail

Evidence

The physical effects of fisheries or dredging activities are addressed under abrasion, penetration and extraction pressures above. No clear biological relationships between the important characteristic species were found. Therefore, the removal of any one species may not affect other members of the community adversely.  However, if the important characterizing species were removed as by-catch, the character of the biotope would change. A significant decline in the abundance of Psolus squamatus would result in the loss of the biotope as recognised by the habitat classification. Therefore, a resistance of 'Medium' is recorded, albeit at Low confidence. As resilience is probably 'Medium' sensitivity is assessed as 'Medium'.

Medium
Low
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Medium
Low
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Medium
Low
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Bibliography

  1. Addy, J.M., Levell, D. & Hartley, J.P., 1978. Biological monitoring of sediments in the Ekofisk oilfield. In Proceedings of the conference on assessment of ecological impacts of oil spills. American Institute of Biological Sciences, Keystone, Colorado 14-17 June 1978, pp.514-539.

  2. Besten, P.J. den, Donselaar, E.G. van, Herwig, H.J., Zandee, D.I. & Voogt, P.A., 1991. Effects of cadmium on gametogenesis in the seastar Asterias rubens L. Aquatic Toxicology, 20, 83-94.

  3. Besten, P.J. den, Herwig, H.J., Zandee, D.I. & Voogt, P.A., 1989. Effects of Cd and PCBs on reproduction in the starfish Asterias rubens: aberrations in early development. Ecotoxicology and Environmental Safety, 18, 173-180.

  4. Billett, D.S.M., 1991. Deep-sea holothurians. Oceanography and Marine Biology: an Annual Review, 29, 259-317. 

  5. Binyon, J., 1966. Salinity tolerance and ionic regulation. In Physiology of Echinodermata (ed. R.A. Boolootian), pp. 359-377. New York: John Wiley & Sons.

  6. Boalch, G.T., 1979. The dinoflagellate bloom on the coast of south-west England, August to September 1978. Journal of the Marine Biological Association of the United Kingdom, 59, 515-517.

  7. Boolootian, R.A.,1966. Physiology of Echinodermata. (Ed. R.A. Boolootian), pp. 822. New York: John Wiley & Sons.

  8. Boolukos, C.M., Lim, A., O'Riordan, R. M. & Wheeler, A.J., 2019. Cold-water corals in decline - A temporal (4 year) species abundance and biodiversity appraisal of complete photomosaiced cold-water coral reef on the Irish Margin. Deep-Sea Research Part I-Oceanographic Research Papers, 146, 44-54. DOI https://doi.org/10.1016/j.dsr.2019.03.004

  9. Brils, J.M., Huwer, S.L., Kater, B.J., Schout, P.G., Harmsen, J., Delvigne, G.A.L. & Scholten, M.C.T., 2002. Oil effect in freshly spiked marine sediment on Vibrio fischeri, Corophium volutator, and Echinocardium caudatum. Environmental Toxicology and Chemistry, 21, 2242-2251.

  10. Bryan, G.W. & Gibbs, P.E., 1991. Impact of low concentrations of tributyltin (TBT) on marine organisms: a review. In: Metal ecotoxicology: concepts and applications (ed. M.C. Newman & A.W. McIntosh), pp. 323-361. Boston: Lewis Publishers Inc.

  11. Bryan, G.W., 1984. Pollution due to heavy metals and their compounds. In Marine Ecology: A Comprehensive, Integrated Treatise on Life in the Oceans and Coastal Waters, vol. 5. Ocean Management, part 3, (ed. O. Kinne), pp.1289-1431. New York: John Wiley & Sons.

  12. Bullimore, B., 1985. An investigation into the effects of scallop dredging within the Skomer Marine Reserve. Report to the Nature Conservancy Council by the Skomer Marine Reserve Subtidal Monitoring Project, S.M.R.S.M.P. Report, no 3., Nature Conservancy Council.

  13. Cabioch, L., Dauvin, J.C. & Gentil, F., 1978. Preliminary observations on pollution of the sea bed and disturbance of sub-littoral communities in northern Brittany by oil from the Amoco Cadiz. Marine Pollution Bulletin, 9, 303-307.

  14. Carney, R.S., 2005. Zonation of deep biota on continental margins. Oceanography and Marine Biology: an Annual Review, 43, 211-278. 

  15. Cole, S., Codling, I.D., Parr, W. & Zabel, T., 1999. Guidelines for managing water quality impacts within UK European Marine sites. Natura 2000 report prepared for the UK Marine SACs Project. 441 pp., Swindon: Water Research Council on behalf of EN, SNH, CCW, JNCC, SAMS and EHS. [UK Marine SACs Project.]. Available from: http://ukmpa.marinebiodiversity.org/uk_sacs/pdfs/water_quality.pdf

  16. Crompton, T.R., 1997. Toxicants in the aqueous ecosystem. New York: John Wiley & Sons.

  17. Daan, R. & Mulder, M., 1996. On the short-term and long-term impact of drilling activities in the Dutch sector of the North Sea ICES Journal of Marine Science, 53, 1036-1044.

  18. Davey, Nicola & Whitfield, Emily, 2013. The Psolidae of New Zealand and some additions to the Macquarie Ridge fauna (Echinodermata: Holothuroidea: Psolidae). Memoirs of Museum Victoria, 70, 51-66.

  19. Davies, J.S., Stewart, H.A., Narayanaswamy, B.E., Jacobs, C., Spicer, J., Golding, N. & Howell, K.L., 2015. Benthic Assemblages of the Anton Dohrn Seamount (NE Atlantic): defining deep-sea biotopes to support habitat mapping and management efforts with a focus on Vulnerable Marine Ecosystems. PLOS ONE, 10 (5), e0124815. DOI https://doi.org/10.1371/journal.pone.0124815

  20. Dinnel, P.A., Pagano, G.G., & Oshido, P.S., 1988. A sea urchin test system for marine environmental monitoring. In Echinoderm Biology. Proceedings of the Sixth International Echinoderm Conference, Victoria, 23-28 August 1987, (R.D. Burke, P.V. Mladenov, P. Lambert, Parsley, R.L. ed.), pp 611-619. Rotterdam: A.A. Balkema.

  21. Duineveld, G.C.A., Lavaleye, M.S.S., Bergman, M.J.N., de Stigter, H. & Mienis, F., 2007b. Trophic structure of a cold-water coral mound community (Rockall Bank, NE Atlantic) in relation to the near-bottom particle supply and current regime. Bulletin of Marine Science, 81 (3), 449-467

  22. Ekman, S., 1923. Uber Psolus squamatus und verwandte Arten. Zugleich ein Beitrag zu Bipolaritatsfrage. Arkiv För Zoologi, Band 15, 1-59.

  23. Ellett, D.J. & Martin, J.H.A., 1973. The physical and chemical oceanography of the Rockall channel. Deep Sea Research and Oceanographic Abstracts, 20 (7), 585-625. DOI https://doi.org/10.1016/0011-7471(73)90030-2

  24. Fankboner, P.V., 1971. The ciliary currents associated with feeding, digestion, and sediment removal in Adula (Botula) falcata Gould 1851. Biological Bulletin, 140, 28-45.

  25. Fernández-Torquemada, Y., González-Correa, J.M. & Sánchez-Lizaso, J.L., 2013. Echinoderms as indicators of brine discharge impacts. Desalination and Water Treatment, 51 (1-3), 567-573. DOI https://doi.org/10.1080/19443994.2012.716609

  26. Findlay, H.S., Hennige, S.J., Wicks, L.C., Navas, J.M., Woodward, E.M.S. & Roberts, J.M., 2014. Fine-scale nutrient and carbonate system dynamics around cold-water coral reefs in the northeast Atlantic. Scientific Reports, 4, 3671. DOI https://doi.org/10.1038/srep03671 Available from  https://www.nature.com/articles/srep03671#supplementary-information

  27. Forster, G.R., 1979. Mortality of the bottom fauna and fish in St Austell Bay and neighbouring areas. Journal of the Marine Biological Association of the United Kingdom, 59, 517-520.

  28. Freese, J.L., 2001. Trawl-induced damage to sponges observed from a research submersible. Marine Fisheries Review, 63 (3), 7-13.

  29. Freese, L., Auster, P.J., Heifetz, J. & Wing, B.L., 1999. Effects of trawling on seafloor habitat and associated invertebrate taxa in the Gulf of Alaska. Marine Ecology Progress Series, 182, 119-126.

  30. Gage, J.D., 1986. The benthic fauna of the Rockall Trough: regional distribution and bathymetric zonation. Proceedings of the Royal Society of Edinburgh. Section B. Biological Sciences, 88, 159-174. DOI https://doi.org/10.1017/S026972700000453X

  31. Griffiths, A.B., Dennis, R. & Potts, G., 1979. Mortality associated with a phytoplankton bloom off Penzance in Mounts Bay. Journal of the Marine Biological Association of the United Kingdom, 59 (2), 520-521

  32. Gutt, J., 1991. Investigations on brood protection in Psolus dubiosus (Echinodermata: Holothuroidea) from Antarctica in spring and autumn. Marine Biology, 111 (2), 281-286. DOI https://doi.org/10.1007/BF01319710

  33. Hemery, L.G., Henkel, S.K. & Cochrane, G.R., 2018. Benthic assemblages of mega epifauna on the Oregon continental margin. Continental Shelf Research, 159, 24-32. DOI https://doi.org/10.1016/j.csr.2018.03.004

  34. Howell, K.L., Davies, J.S. & Narayanaswamy, B.E., 2010. Identifying deep-sea megafaunal epibenthic assemblages for use in habitat mapping and marine protected area network design. Journal of the Marine Biological Association of the United Kingdom, 90 (01), 33. DOI https://doi.org/10.1017/S0025315409991299

  35. Hyman, L.V., 1955. The Invertebrates: Vol. IV. Echinodermata. The coelomate Bilateria. New York: McGraw Hill.

  36. Jennings, S. & Kaiser, M.J., 1998. The effects of fishing on marine ecosystems. Advances in Marine Biology, 34, 201-352.

  37. King, William, 1846. XXVI.—An account of some shells and other invertebrate forms found on the coast of Northumberland and of Durham. Annals and Magazine of Natural History, 18 (119), 233-251. DOI https://doi.org/10.1080/037454809494420

  38. Kinne, O. (ed.), 1984. Marine Ecology: A Comprehensive, Integrated Treatise on Life in Oceans and Coastal Waters.Vol. V. Ocean Management Part 3: Pollution and Protection of the Seas - Radioactive Materials, Heavy Metals and Oil. Chichester: John Wiley & Sons.

  39. Lawrence, J.M., 1996. Mass mortality of echinoderms from abiotic factors. In Echinoderm Studies Vol. 5 (ed. M. Jangoux & J.M. Lawrence), pp. 103-137. Rotterdam: A.A. Balkema.

  40. Maier, S.R., Brooke, S., De Clippele, L.H., de Froe, E., Van der Kaaden, A-S., Kutti, T., Mienis, F. & Van Oevelen, D., 2023. On the paradox of thriving cold-water coral reefs in the food-limited deep sea. Biological Reviews, 98 (5), 1768-1795. DOI https://doi.org/10.1111/brv.12976

  41. Martinez, M.I., Martínez-Salinas, A.P. & Moura, R.B., 2024. Chapter 14 - Knowledge of biodiversity and reproduction in sea cucumbers from southern South America to the Antarctic Peninsula. In Mercier, A., Hamel, J-F., Suhrbier, A.D. & Pearce, C.M. (eds.). The World of Sea Cucumbers. Academic Press, pp. 201-220. DOI https://doi.org/10.1016/B978-0-323-95377-1.00006-0

  42. McEuen, F.S. & Chia, F.S., 1991. Development and metamorphosis of two psolid sea cucumbers, Psolus chitonoides andPsolidium bullatum, with a review of reproductive patterns in the family Psolidae (Holothuroidea: Echinodermata). Marine Biology, 109 (2), 267-279. DOI https://doi.org/10.1007/BF01319395

  43. Meyer, H.K., Roberts, E.M., Mienis, F. & Rapp, H.T., 2020. Drivers of Megabenthic Community Structure in One of the World’s Deepest Silled-Fjords, Sognefjord (Western Norway). Frontiers in Marine Science, 7. DOI https://doi.org/10.3389/fmars.2020.00393

  44. Mortensen, T.H., 1927. Handbook of the echinoderms of the British Isles. London: Humphrey Milford, Oxford University Press.

  45. Newton, L.C. & McKenzie, J.D., 1998. Brittlestars, biomarkers and Beryl: Assessing the toxicity of oil-based drill cuttings using laboratory, mesocosm and field studies. Chemistry and Ecology, 15, 143-155.

  46. O’Loughlin, P.M., 2020. Brood-protecting and fissiparous cucumariids (Echinodermata, Holothurioidea). In Guille, A., Feral, J.-P. and Roux, M. (eds.). Echinoderms through time, Rotterdam, Netherlands: CRC Press, pp. 539-547. [Proceedings of the eighth international echinoderm conference, Dijon, France, 6-10 September 1993].

  47. OBIS (Ocean Biodiversity Information System),  2024. Global map of species distribution using gridded data. Available from: Ocean Biogeographic Information System. www.iobis.org. Accessed: 2024-12-27

  48. Olsgard, F. & Gray, J.S., 1995. A comprehensive analysis of the effects of offshore oil and gas exploration and production on the benthic communities of the Norwegian continental shelf. Marine Ecology Progress Series, 122, 277-306.

  49. Picton, B. & Goodwin, C., 2007. Sponge biodiversity of Rathlin Island, Northern Ireland. Journal of the Marine Biological Association of the United Kingdom, 87 (06), 1441-1458.

  50. Riedel, B., Zuschin, M. & Stachowitsch, M., 2012. Tolerance of benthic macrofauna to hypoxia and anoxia in shallow coastal seas: a realistic scenario. Marine Ecology Progress Series, 458, 39-52.

  51. Roberts, D.A., Johnston, E.L. & Knott, N.A., 2010b. Impacts of desalination plant discharges on the marine environment: A critical review of published studies. Water Research, 44 (18), 5117-5128.

  52. Roberts, J.M., Henry, L.A., Long, D. & Hartley, J.P., 2008. Cold-water coral reef frameworks, megafaunal communities and evidence for coral carbonate mounds on the Hatton Bank, north east Atlantic. Facies, 54 (3), 297-316. DOI https://doi.org/10.1007/s10347-008-0140-x

  53. Rosenberg, R., Hellman, B. & Johansson, B., 1991. Hypoxic tolerance of marine benthic fauna. Marine Ecology Progress Series, 79, 127-131. DOI https://dx.doi.org/10.3354/meps079127

  54. Russell, M., 2013. Echinoderm Responses to Variation in Salinity. Advances in Marine Biology, 66, 171-212. DOI http://dx.doi.org/10.1016/B978-0-12-408096-6.00003-1

  55. Sherwin, T.J., Read, J.F., Holliday, N.P. & Johnson, C., 2012. The impact of changes in North Atlantic Gyre distribution on water mass characteristics in the Rockall Trough. ICES Journal of Marine Science, 69 (5), 751-757. DOI https://doi.org/10.1093/icesjms/fsr185

  56. Smith, J.E. (ed.), 1968. 'Torrey Canyon'. Pollution and marine life. Cambridge: Cambridge University Press.

  57. Southward, A.J., Southward, E.C. & Cooper, L.H.N., 1958. On the occurrence and behaviour of two little-known barnacles, Hexelasma hirsutum and Verruca recta, from the continental slope. Journal of the Marine Biological Association of the United Kingdom, 37 (3), 633-647. DOI https://doi.org/10.1017/S0025315400005683

  58. Spehar, R.L., Poucher, S., Brooke, L.T., Hansen, D.J., Champlin, D. & Cox, D.A., 1999. Comparative Toxicity of Fluoranthene to Freshwater and Saltwater Species Under Fluorescent and Ultraviolet Light. Archives of Environmental Contamination and Toxicology, 37 (4), 496-502. DOI https://doi.org/10.1007/s002449900544

  59. Stickle, W.B. & Diehl, W.J., 1987. Effects of salinity on echinoderms. In Echinoderm Studies, Vol. 2 (ed. M. Jangoux & J.M. Lawrence), pp. 235-285. A.A. Balkema: Rotterdam.

  60. Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523. DOI https://doi.org/10.1093/icb/33.6.510

  61. Thorson, G., 1946. Reproduction and larval development of Danish marine bottom invertebrates, with special reference to the planktonic larvae in the Sound (Øresund). Meddelelser fra Kommissionen for Danmarks Fiskeri- Og Havundersögelser, Serie: Plankton, 4, 1-523.

  62. Tyler, P.A. & Young, C.M., 1998. Temperature and pressure tolerances in dispersal stages of the genus Echinus (Echinodermata: Echinoidea): prerequisites for deep sea invasion and speciation. Deep Sea Research II, 45 (1), 253-277. DOI https://doi.org/10.1016/S0967-0645(97)00091-X

  63. Veale, L.O., Hill, A.S., Hawkins, S.J. & Brand, A.R., 2000. Effects of long term physical disturbance by scallop fishing on subtidal epifaunal assemblages and habitats. Marine Biology, 137, 325-337.

  64. Wagstaff, M.C., Howell, K.L., Bett, B.J., Billett, D.S.M., Brault, S., Stuart, C.T. & Rex, M.A., 2014. β-diversity of deep-sea holothurians and asteroids along a bathymetric gradient (NE Atlantic). Marine Ecology Progress Series, 508, 177-185. DOI https://doi.org/10.3354/meps10877

  65. Wienberg, C., Beuck, L., Heidkamp, S., Hebbeln, D., Freiwald, A., Pfannkuche, O. & Monteys, X., 2008. Franken Mound: facies and biocoenoses on a newly-discovered “carbonate mound” on the western Rockall Bank, NE Atlantic. Facies, 54 (1), 1-24. DOI https://doi.org/10.1007/s10347-007-0118-0

  66. Young, C.M. & Chia, F.S., 1982. Factors controlling spatial distribution of the sea cucumber Psolus chitonoides: Settling and post-settling behavior. Marine Biology, 69 (2), 195-205. DOI https://doi.org/10.1007/BF00396899

Citation

This review can be cited as:

Tyler-Walters, H., 2024. Psolus squamatus, Anomiidae, serpulid polychaetes and Munida on Atlantic mid bathyal rock and other hard substrata. 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 27-12-2024]. Available from: https://marlin.ac.uk/habitat/detail/1316

Last Updated: 29/11/2024