MarLIN

information on the biology of species and the ecology of habitats found around the coasts and seas of the British Isles

Kophobelemnon field on Atlantic mid bathyal mud

Summary

UK and Ireland classification

UK and Ireland classification

Description

This biotope is composed of dense aggregations of sea pens of the genus Kophobelemnon (in the UK likely to be Kophobelemnon stelliferum) on mud. Kophobelemnon fields are also found in the upper bathyal zone but the associated infauna are likely to differ. The characterizing species listed refer to all Kophobelemnon stelliferum assemblages not just those found associated with the zone and substrata specified in this biotope. (Information from JNCC, 2015).

Depth range

600-1300 m

Additional information

-

Listed By

- none -

Further information sources

Search on:

Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

Kophobelemnon assemblages are associated with fine mud and muddy sand substrata and occur at a range of depths in the deep-sea. Assemblages occur on mud in the Atlantic upper bathyal zone (M.AtUB.Mu.SpnMeg.KopFie) and also on mud within the mid-bathyal zone (M.AtMB.Mu.SpnMeg.KopFie). The sensitivity of these Kophobelemon dominated biotopes is, therefore, assessed as a group, on the assumption that their sensitivity is very similar in terms of substratum and functional groups present. Any differences in species or biotope response to pressures are highlighted.

The dominant species for the biotopes are in the genus Kophobelemnon and likely to be Kophobelemnon stelliferum in UK waters. Loss of this species may result in loss or degradation of these biotopes, therefore, the sensitivity of these biotopes are dependent on the sensitivity of Kophobelemnon. Other pennatulaceans, such as Pennatula phosphorea and the genus Protoptilum, can be present. Other species that can be found within this biotope are Cerianthidae anemones, the cup coral Flabellum chunii, the polychaetes Lanice conchilega (sand mason worm) and the decapod crustacean Polycheles typhlops. The sensitivity of the other Pennatulid seapens that can occur in this biotope (Pennatula phosphorea and Protoptilum) is likely to be very similar to that of Kophobelemnon. Where information is unavailable for Kophobelemnon, other suitable proxy seapen species will be used.  Polycheles typhlops, Lanice conchilega and Cerianthidae are ubiquitous and are not unique to this biotope. Flabellum chunii similarly occurs on a range of substratum types and is not specifically associated with seapens or Kophobelemnon. Therefore, these species are therefore not considered significant to the assessment of sensitivity. More information on Cerianthidae can be found in other biotope assessments available on this website.

Resilience and recovery rates of habitat

Kophobelemnon stelliferum, like Protoptilum sp. and Pennatula phosphorea, is a sessiflorate pennatulid found on soft mud/sand substratum within a wide depth and geographical range (Baker et al., 2012; Bastari et al., 2018). Kophobelemnon stelliferum is widely distributed along the continental slope of the Atlantic and Pacific Oceans, including in the Mediterranean Sea, off Namibia, Baffin Island, Newfoundland and Labrador, at depths ranging from 40 to 2,500 m (Baker et al., 2012; Bastari et al., 2018; Mastrototaro et al., 2013; Wareham & Edinger, 2007; López-González et al., 2001). Briggs et al. (1996) also recorded Kophobelemnon sp. in the Venezuela Basin at depths of 3934-4095. Kophoblemenon spp. have also been observed in the Whittard Canyon (Hogan et al., 2019), however the depth range of the species is known to be restricted in submarine canyons, most likely due to hydrodynamic controls (Rice et al., 1992). Protoptilum carpenteri was found within the facies of Kophobelemnon stelliferum in the Mediterranean Sea at depths between 240-451 m. Kophobelemnon spp. form erect colonies that can reach 70 cm in length (Rice et al., 1992). The Kophobelemnon stelliferum colonies in the Mediterranean , as sampled by Mastrototaro et al. (2013), are smaller than those reported for Atlantic colonies, possibly exhibiting a case of Mediterranean dwarfism due to the particular hydrological conditions of the basin (Mastrototaro et al., 2013). However, it is still unclear whether the Mediterranean supports a separate endemic species of Kophobelemnon (Mastrototaro et al., 2013).

Seapens have a central primary polyp that forms the axis of the colonies, called the rachis, which has secondary polyps known as autozooids (polyps with well-developed pinnular tentacles), that are responsible for feeding and reproduction (Mastrototaro et al., 2013; Rice et al., 1992). These polyps are also retractable (Rice et al., 1992). Siphonozooids (polyps with a well-developed siphonoglyph and without tentacles), play a role in circulating water through the interior of the colony (Mastrototaro et al., 2013; Rice et al., 1992). The proximal part of the rachis, known as the peduncle, anchors the colony to the substratum. Studies have shown that there is a significant positive correlation between polyp number and colony length (Mastrototaro et al., 2013; Rice et al., 1992). There is also a marked change in the growth form at a total colony length of 250-300 mm, at which the polyp number/colony length ratio increases rather dramatically, with larger colonies having relatively more polyps than smaller ones (Rice et al., 1992). As the polyps have a reproductive role, it is thought that this is a reproductive advantage, whereby the larger number of polyps increases the chances of fertilisation (Levitan, 1996, cited in Pires et al., 2009). As Kophobelemnon stelliferum is gonochoric, with a 1:1 sex ratio, the presence of large aggregations of the species, such as in the cases of the Kophobelemnon fields biotopes further increases the chances of fertilization (Pires et al., 2009).

Kophobelemnon sp. reach sexual maturity with greater colony lengths (25 cm), and gonads are distributed throughout the rachis and the distal part of the peduncle (Rice et al., 1992). There is a maximum oocyte size of 800 µm, and evidence of oocytes in various stages of development within each individual polyp of a colony suggests that fecundity can be high (Rice et al., 1992). Rice et al. (1992) found no obvious synchronization of oocyte development within an individual autozooid polyp or between different autozooids within a colony. Similarly, the testes were also found to be in different stages of development both within an autozooid and within a colony (Rice et al., 1992). High fecundity has also been recorded for the deep-sea seapen species Anthoptilum murrayi (Pires et al., 2009) and Ptilosarcus gurneyi, which can produce >200,000 eggs in one season (Chia and Crawford, 1973).

Spawning has never been observed in Kophobelemnon stelliferum, but it is thought to be mechanistically similar to other pennatulids, where the sperm and oocytes leave via the mouth of the autozooid (Rice et al., 1992). No developing embyros were found within the colonies of Kophobelemnon stelliferum, and a period of lecithotrophic development is inferred (Rice et al., 1992). Seasonality has not been observed in the reproductive cycle of Kophobelemnon stelliferum (Rice et al., 1992), so it is likely to be continuous, similar to that of other deep-sea seapens such as Anthoptilym murrayi (Pires et al., 2009) and Pennatula aculeata (Eckelbarger et al., 1998). Another seapen from the Kophobelemnidae family, Malacobelemnon daytoni, which is found in Antarctic shallow-waters, does display seaonality however (Servetto and Sahade, 2016).

Limited information is available on recruitment in seapens. Recruitment in Ptilosarcus gurneyi, a species which occurs in shallow-waters and the deep-sea in the Pacific, is thought to occur either annually or every few years, however it is highly variable (both over time and space; Birkeland, 1974). A morphometry and growth study by Murillo et al. (2018) suggested that recruitment in deep-sea seapen species (Pennatula aculeata, Pennatula grandis, Anthoptilum grandiforum and Halipteris finmarchica) occurs in multi-year pulses, with some periods of limited or no recruitment.

No information was available on the longevity of Kophobelemnon, however other deep-sea seapen species have been aged. De Moura Neves et al. (2018) found that the maximum age of Umbellula encrinus from Baffin Bay (between Greenland and Canada, although the species also occurs in Norwegian waters) was 75 years, and growth rates averaged 0.067 +/- 0.014 mm/year for radial extension and 4.5 +/- 1.2 cm/year for linear extension. Slower growth rates occurred in smaller colonies, whereas larger colonies of Umbellula encrinus had faster growth rates, however the smallest colony indicated that juveniles may potentially exhibit exceptionally fast growth rates (5 cm/yr). Two Halipteris seapen species have been aged according to their growth rings. Halipteris willemoesi, which occurs from the circalittoral to deep-sea, has an estimated longevity of at least 48 years (Wilson et al., 2002), and the deep-sea Halipteris finmarchica has been recorded with a maximum of 22 years (Neves et al., 2015). Based on estimated ages of Halipteris willemoesi, the average growth rate in total length has been estimated to be 3.9, 6.1 and 3.6 cm/yr for small, medium and large-sized colonies (Wilson et al., 2002). In Halipteris finmarchica, linear growth rates averaged 4.9 cm/yr, meaning a colony would take four years to reach sexual maturity (at 18 cm) (Neves et al., 2015). No significant relationships were found between the linear growth rates and environmental variables in Halipteris finmarchica, however when data for both Halipteris finmarchica and Halipteris willemoesi were pooled, diametric growth rates were found to be statistically related to latitude, temperature, chlorophyll a concentration and POC (particulate organic carbon) (Neves et al., 2015). Temperature was negatively related to diametric growth rates, but as this is contrary to the typical positive relationship between these two variables, this was noted with caution (Neves et al., 2015). POC and chlorophyll a are related to food availability (Neves et al., 2015). Colony age was found to be positively related to depth, possibly related to the reduced fishing exposure (Neves et al., 2015). Neves et al. (2015) suggest that recovery rates of Halipteris finmarchica following damage could take over 20 years.

Kophobelemnon stelliferum is known to be a suspension feeder and secondary zooids attached to the central rachis are responsible for feeding (Rice et al., 1992). Kophobelemnon stelliferum is a sessile benthic animal, anchored into the sediment by the peduncle. Kophobelemnon stelliferum, Pennatula phosphorea  and Protoptilum carpenteri are however able to retract into the sediment, likely in response to predation or disturbance (Baker et al., 2012; Chimienti et al., 2018; De Clippele et al., 2015; Greathead et al., 2015;  Langton et al., 1990). Baker et al. (2012), for example, observed rapid retraction of entire colonies of Protoptilum carpenteri into the sediment. Virgularia mirabilis (which occurs from very shallow waters to deep-sea) can withdraw in a few seconds, however a study by Chimienti et al., 2018 found the shallow-water species Pennatula rubra took between 210 and 340 seconds to completely withdraw. Movement by detaching, drifting and re-attaching has also been recorded in the deep-water seapens Umbellula lindahli (Flores, 1999, cited in Wilson et al., 2002) and Ptilosarcus gurneyi (Birkeland, 1974; note this species also occurs in shallow water), as well as some shallow-water species, e.g. Renilla kollikeri (Kastendiek, 1976) and Pennatula rubra (Chimienti et al., 2018). For Pennatula rubra, the seapens were found to inflate themselves with seawater and then get carried by currents, possibly as a displacement strategy or dispersal behaviour (Chimienti et al., 2018). Although there is no evidence for this in Kophobelemnon, Musgrave (1909) suggests that it is extremely probable that many deep-sea Pennatulids do have controlled locomotion abilities. Eno et al. (2001) also noted that the seapen Pennatula phosphorea was able to re-establish itself within 144 hours after removal by divers. However, although Malecha and Stone (2009) found that dislodged seapens were able to re-bury in sediment after disturbance, even without further contact, they eventually became dislodged again.

Biological traits analysis (García-Alegre et al., 2018) was undertaken for eight deep-sea seapen taxa, including Kophobelemnon stelliferum and Pennatula spp. (aculeata/grandis), from specimens taken from the Flemish Cap off Newfoundland. This analysis suggests that all eight taxa have intermediate fragility (not defined), a soft/endoskeleton (non-solid), feed on living planktonic material, prefer mud/sand substratum, have both physical and chemical combat defence mechanisms and are broadcast spawners (Umbellula lindahli is also asexual; Tyler et al., 1995). All are colonial, with five taxa also being gregarious. The majority (except one, unspecified) have a moderate attachment strength, the other (unspecified) has a high attachment strength. All but one taxa have a high degree of flexibility (>45°), with the other taxa (likely to be Kophobelemnon stelliferum) having a flexibility of only 10-45°. López-González et al. (2001) support this, noting that the whole colony of Kophobelemnon stelliferum is very rigid due to the high density of sclerites. Murillo et al. (2018) also state that certain seapen growth forms (such as plumose), may require more rigidity that others (e.g. flagelliform), due to their increased drag.

On the Norwegian continental margin, Kophobelemnon stelliferum was found to play a key role as a shelter against predators, with Munida sp. frequently observed underneath and near to the species (De Clippele et al., 2015). Kophobelemnon stelliferum has stinging cells and emits light, which is likely to scare away potential predators of organisms sheltering under the seapen (De Clippele et al., 2015), and it is thought that the squat lobsters use the seapen as a base station for scavenging and active hunting (De Clippele et al., 2015).

Resilience assessment. Where resistance is ‘None’ or ‘Low’, and an element of habitat recovery is required, resilience is assessed as ‘Low’ (10-25 years). The evidence suggests that recovery to maximum sizes (i.e. pre-disturbance conditions) could take 13-22 years, based on other deep-sea seapen growth rates (de Moura Neves et al., 2018; Wilson et al., 2002). Recovery from physical impacts has been shown to take >4 years (Lindholm et al., 2008) for the proxy species Halipeteris willemoesi and Pennatularia showed no recovery after 0, 0.5, 3 or 7 years after intensive ploughing (Bluhm, 2001; Simon-Lledó et al., 2019). The confidence associated with this score are ‘Medium’ for Quality of Evidence (proxy used and some expert judgement on recovery time), ‘Medium’ for Applicability of Evidence (studies from Gulf of Alaska/Bering Sea, California, New Zealand and Peru) and ‘Medium’ for Degree of Concordance.  Where resistance of the characterizing species is ‘Medium’ or ‘High’ and the habitat has not been altered, resilience is assessed as ‘Low’ (10-25 years). Kophobelemnon stelliferum is likely to only reach sexual maturity at 5-8 years (based on size, using growth rates from other species; de Moura Neves et al., 2018; Rice et al., 1992; Wilson et al., 2002), with fewer and smaller colonies, which  reduces the chance of fertilization (Pires et al., 2009). Furthermore, recruitment is likely to occur in multi-year pulses, with some periods of limited or no recruitment (Murillo et al., 2018). As mentioned above, recovery to pre-disturbance sizes could also take 13-22 years (de Moura Neves et al., 2018; Wilson et al., 2002). The confidences associated with this score are ‘Medium’ for Quality of Evidence (proxies used), ‘Medium’ for Applicability of Evidence (studies from across Atlantic and Pacific) and ‘High’ for Degree of Concordance. An exception is made for permanent or ongoing (long-term) pressures where recovery is not possible as the pressure is irreversible, in which case resilience is assessed as ‘Very low’ by default.

Climate Change Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
High High Not sensitive
Q: Medium
A: Low
C: High
Q: High
A: High
C: High
Q: Medium
A: Low
C: High

There is little direct evidence on the resistance of Kophobelemnon stelliferum to temperature changes. In the absence of this information, the temperature tolerances of Kophobelemnon stelliferum (based on their current geographical range and environmental preferences) have been used as a proxy.

Yesson et al. (2012) found that temperature was the most important factor in habitat suitability modelling for Pennatulacea, which included records of Kophobelemnidae, Protoptilidae and Pennatulidae. Similarly, Georgian et al. (2019) found temperature to be the most important for modelling Pennatulacea. However, they had one of the largest observed temperature ranges of all taxa studied. Modelling work was undertaken by Gormley et al. (2015) under an increased ocean temperature scenario of 4°C by 2100. The modelling suggested that sea pens would experience a significant loss of suitable habitat. In the UK, this is predicted to be a loss of 15,706 km2. However, this value represents all sea pens and burrowing megafauna communities under the Priority Marine Habitats definition (i.e. water depths from 15-200m), so is not specific to Kophobelemnon stelliferum.

The sea pen Virgularia mirabilis is known to experience annual temperature variations of 10°C in Scottish coastal waters (Hughes, 1998a), however in the more stable mid to upper bathyal waters where Kophobelemnon stelliferum is found, temperature variation is likely to be narrower (Gage & Tyler, 1991, cited in Neves et al., 2015). The wide distribution of Kophobelemnon stelliferum (Mastrototaro et al., 2013; Rice et al., 1992) suggests it is eurythermal. Kophobelemnon stelliferum assemblages were reported by Howell et al. (2010) to occur within the temperature range of 7-12°C (average 9.78°C) in the North-east Atlantic. Kophobelemnon stelliferum also occurs off Newfoundland where average bottom temperatures are ~4.4°C (>1,100 m) and ~5°C (<1,100 m; Baker et al., 2012). Kophobelemnon spp. have also been recorded in the Venezuela Basin, where the bottom water temperature is relatively constant at 3.83 to 3.86°C (Briggs et al., 1996).

Sensitivity assessment. As Kophobelemnon stelliferum naturally occurs within a range of bottom water temperatures, it is not likely to be affected by a change in bottom temperature at the middle, high or extreme scenario pressure benchmarks (all relating to a 1°C rise in temperature in the deep-sea). Therefore, resistance is assessed as ‘High’, resistance as ‘High’ and the biotopes are considered to be ‘Not sensitive’ at the pressure benchmarks for extreme, high and middle scenarios.

High High Not sensitive
Q: Medium
A: Low
C: High
Q: High
A: High
C: High
Q: Medium
A: Low
C: High

There is little direct evidence on the resistance of Kophobelemnon stelliferum to temperature changes. In the absence of this information, the temperature tolerances of Kophobelemnon stelliferum (based on their current geographical range and environmental preferences) have been used as a proxy.

Yesson et al. (2012) found that temperature was the most important factor in habitat suitability modelling for Pennatulacea, which included records of Kophobelemnidae, Protoptilidae and Pennatulidae. Similarly, Georgian et al. (2019) found temperature to be the most important for modelling Pennatulacea. However, they had one of the largest observed temperature ranges of all taxa studied. Modelling work was undertaken by Gormley et al. (2015) under an increased ocean temperature scenario of 4°C by 2100. The modelling suggested that sea pens would experience a significant loss of suitable habitat. In the UK, this is predicted to be a loss of 15,706 km2. However, this value represents all sea pens and burrowing megafauna communities under the Priority Marine Habitats definition (i.e. water depths from 15-200m), so is not specific to Kophobelemnon stelliferum.

The sea pen Virgularia mirabilis is known to experience annual temperature variations of 10°C in Scottish coastal waters (Hughes, 1998a), however in the more stable mid to upper bathyal waters where Kophobelemnon stelliferum is found, temperature variation is likely to be narrower (Gage & Tyler, 1991, cited in Neves et al., 2015). The wide distribution of Kophobelemnon stelliferum (Mastrototaro et al., 2013; Rice et al., 1992) suggests it is eurythermal. Kophobelemnon stelliferum assemblages were reported by Howell et al. (2010) to occur within the temperature range of 7-12°C (average 9.78°C) in the North-east Atlantic. Kophobelemnon stelliferum also occurs off Newfoundland where average bottom temperatures are ~4.4°C (>1,100 m) and ~5°C (<1,100 m; Baker et al., 2012). Kophobelemnon spp. have also been recorded in the Venezuela Basin, where the bottom water temperature is relatively constant at 3.83 to 3.86°C (Briggs et al., 1996).

Sensitivity assessment. As Kophobelemnon stelliferum naturally occurs within a range of bottom water temperatures, it is not likely to be affected by a change in bottom temperature at the middle, high or extreme scenario pressure benchmarks (all relating to a 1°C rise in temperature in the deep-sea). Therefore, resistance is assessed as ‘High’, resistance as ‘High’ and the biotopes are considered to be ‘Not sensitive’ at the pressure benchmarks for extreme, high and middle scenarios.

High High Not sensitive
Q: Medium
A: Low
C: High
Q: High
A: High
C: High
Q: Medium
A: Low
C: High

There is little direct evidence on the resistance of Kophobelemnon stelliferum to temperature changes. In the absence of this information, the temperature tolerances of Kophobelemnon stelliferum (based on their current geographical range and environmental preferences) have been used as a proxy.

Yesson et al. (2012) found that temperature was the most important factor in habitat suitability modelling for Pennatulacea, which included records of Kophobelemnidae, Protoptilidae and Pennatulidae. Similarly, Georgian et al. (2019) found temperature to be the most important for modelling Pennatulacea. However, they had one of the largest observed temperature ranges of all taxa studied. Modelling work was undertaken by Gormley et al. (2015) under an increased ocean temperature scenario of 4°C by 2100. The modelling suggested that sea pens would experience a significant loss of suitable habitat. In the UK, this is predicted to be a loss of 15,706 km2. However, this value represents all sea pens and burrowing megafauna communities under the Priority Marine Habitats definition (i.e. water depths from 15-200m), so is not specific to Kophobelemnon stelliferum.

The sea pen Virgularia mirabilis is known to experience annual temperature variations of 10°C in Scottish coastal waters (Hughes, 1998a), however in the more stable mid to upper bathyal waters where Kophobelemnon stelliferum is found, temperature variation is likely to be narrower (Gage & Tyler, 1991, cited in Neves et al., 2015). The wide distribution of Kophobelemnon stelliferum (Mastrototaro et al., 2013; Rice et al., 1992) suggests it is eurythermal. Kophobelemnon stelliferum assemblages were reported by Howell et al. (2010) to occur within the temperature range of 7-12°C (average 9.78°C) in the North-east Atlantic. Kophobelemnon stelliferum also occurs off Newfoundland where average bottom temperatures are ~4.4°C (>1,100 m) and ~5°C (<1,100 m; Baker et al., 2012). Kophobelemnon spp. have also been recorded in the Venezuela Basin, where the bottom water temperature is relatively constant at 3.83 to 3.86°C (Briggs et al., 1996).

Sensitivity assessment. As Kophobelemnon stelliferum naturally occurs within a range of bottom water temperatures, it is not likely to be affected by a change in bottom temperature at the middle, high or extreme scenario pressure benchmarks (all relating to a 1°C rise in temperature in the deep-sea). Therefore, resistance is assessed as ‘High’, resistance as ‘High’ and the biotopes are considered to be ‘Not sensitive’ at the pressure benchmarks for extreme, high and middle scenarios.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Marine heatwaves caused by increased air-sea flux of heat are only expected to penetrate surface waters (≤ 50 m) (Cerrano et al., 2000, Garrabou et al., 2009; Dan Smale, pers. comms.). The Kophobelemnon field biotopes occur in the Atlantic mid and upper bathyal zone, at depths of 200 – 1300 m. Therefore, the biotope will not be affected by changes arising from marine heatwaves, and the assessment at the pressure benchmark is ‘Not relevant’.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: 
C: NR
Q: NR
A: NR
C: NR

Marine heatwaves caused by increased air-sea flux of heat are only expected to penetrate surface waters (≤ 50 m) (Cerrano et al., 2000, Garrabou et al., 2009; Dan Smale, pers. comms.). The Kophobelemnon field biotopes occur in the Atlantic mid and upper bathyal zone, at depths of 200 – 1300 m. Therefore, the biotope will not be affected by changes arising from marine heatwaves, and the assessment at the pressure benchmark is ‘Not relevant’.

High High Not sensitive
Q: Medium
A: Medium
C: Low
Q: High
A: High
C: High
Q: Medium
A: Medium
C: Low

Unlike scleractinians, octocorals do not have a rigid calcium carbonate exoskeleton, however they do form discrete sclerites consisting of high magnesium calcite (Schubert et al., 2017). Kophobelemnon stelliferum, for example, has a calcified axis (López-González et al., 2009) and a high density of sclerites (López-González et al., 2001). This magnesium calcite is more soluble than the aragonite precipitated by scleractinians (Schubert et al., 2017), putting octocorals at risk from ocean acidification. However, the sclerites in octocorals are loosely dispersed in their fleshy tissue, so are not in direct contact with surrounding seawater. This tissue may therefore provide some protection against ocean acidification, as suggested in a study by Gabay et al. (2014), on a shallow water zooxanthellate octocoral, Ovabunda macrospiculata. At a reduced pH of 7.6 and 7.3, when exposed for 31 - 42 days, no dissolution occurred to the sclerites which were protected by tissue. In comparison, isolated sclerites underwent microstructural changes indicating dissolution, which caused more than 60% damage over the exposure period (Gabay et al., 2014). Despite the results of this study, experiments on other octocorals showed differing effects of ocean acidification, ranging from no response to a decrease in calcification, varying by species (see Schubert et al., 2017; note that no Pennatulacea studies were included).

An ongoing study looking at the effects of ocean acidification on the shallow water Antarctic seapen Malacobelemnon daytoni, which is in the same family as Kophobelemnon stelliferum (Kophobelemnidae), has indicated that several stress responses occur under acidic conditions, along with up-regulated genes (Natalia Servetto, 2019, pers. comm., 15th October). Kophobelemnon stelliferum however is known to occur naturally in the slightly acidic waters around mud volcanoes (Rueda et al., 2016; Sitjà et al., 2019). For example, in the Spanish mud volcano fields, pH was measured by Rueda et al. (2012) to vary from 7.4 to 6.8. Kita et al. (2015) studied the effects of CO2 gas released from the seabed at a depth of 12 m in Scotland. The seapen Virgularia mirabilis (which also occurs in the deep-sea) was frequently observed in the study area, however no abnormal behaviour was observed in response to the CO2, either during or after release. The authors note that the elevated pCO2 concentrations (<1599 µatm, at 30 cm above the seabed) returned to background levels (360-370 µatm) very quickly. Therefore, Virgularia mirabilis was not necessarily exposed to continuous low pH conditions.

As the oceans absorb carbon dioxide from the atmosphere, leading to a decrease in pH and an increase in acidity, there is a further concern; the shoaling of the aragonite saturation horizon (ASH). The ASH is defined as the depth in the oceans at which aragonite saturation equals 1. Below this depth, aragonite saturation will fall below 1, and dissolution of calcified structures may occur. Currently the depth of the ASH in the North Atlantic is approximately 2000m (Jiang et al., 2015). This depth has already become 80-150m shallower over the past two centuries (Chung et al., 2003, Feely et al., 2004). By the end of this century, the depth of the ASH is expected to become shallower still, reaching depths of up to 400 m under the high emission scenario (RCP 8.5) and 600 m for the middle emission scenario (RCP 4.5) (Zheng & Long, 2014).

In addition to the shallowing of the aragonite saturation horizon, the calcite saturation horizon will also become shallower as ocean acidification occurs (Feely et al., 2004). The calcite saturation horizon is more relevant for octocorals, as they require calcium carbonate for structural development and calcite is essential in the skeletal structure (specifically for their sclerites; (Bayer and Macintyre, 2001). Yesson et al. (2012) found that a 12% of records of octocorals occurred in water undersaturated with aragonite (<1). For Sessiliflorae (seapens), 15-20% of observations were below the aragonite saturation horizon.  Calcite saturation state was the second most important factor determining the model for Pennatulacea (after temperature, although it is noted that all environmental variables are intercorrelated). For the Sessiliflorae, 6% were found in areas undersaturated for calcite, however this was affected by a shallow water sampling bias.

Sensitivity assessment. Kophobelemnon stelliferum has a fleshy structure where its sclerites are protected by the surrounding tissue. It is therefore possible that the species can tolerate a decrease in pH. This is supported by evidence showing that the species occurs in the acidic waters associated with mud volcanoes (pH 7.4 to 6.8; Rueda et al., 2016, 2012; Sitjà et al., 2019). However, there is some evidence that suggests that another (shallow-water) species in the Kophobelemnidae family shows stress responses and up-regulated genes in response to ocean acidification (Natalia Servetto, 2019, pers. comm., 15th October). Under the mid and high emission scenarios, resistance is assessed as ‘High’, as no mortality has been observed. Resilience is assessed as ‘High’ and the biotopes are assessed as ‘Not sensitive’ at the middle and high emmission scenario pressure benchmarks.

High High Not sensitive
Q: Medium
A: Medium
C: Low
Q: High
A: High
C: High
Q: Medium
A: Medium
C: Low

Unlike scleractinians, octocorals do not have a rigid calcium carbonate exoskeleton, however they do form discrete sclerites consisting of high magnesium calcite (Schubert et al., 2017). Kophobelemnon stelliferum, for example, has a calcified axis (López-González et al., 2009) and a high density of sclerites (López-González et al., 2001). This magnesium calcite is more soluble than the aragonite precipitated by scleractinians (Schubert et al., 2017), putting octocorals at risk from ocean acidification. However, the sclerites in octocorals are loosely dispersed in their fleshy tissue, so are not in direct contact with surrounding seawater. This tissue may therefore provide some protection against ocean acidification, as suggested in a study by Gabay et al. (2014), on a shallow water zooxanthellate octocoral, Ovabunda macrospiculata. At a reduced pH of 7.6 and 7.3, when exposed for 31 - 42 days, no dissolution occurred to the sclerites which were protected by tissue. In comparison, isolated sclerites underwent microstructural changes indicating dissolution, which caused more than 60% damage over the exposure period (Gabay et al., 2014). Despite the results of this study, experiments on other octocorals showed differing effects of ocean acidification, ranging from no response to a decrease in calcification, varying by species (see Schubert et al., 2017; note that no Pennatulacea studies were included).

An ongoing study looking at the effects of ocean acidification on the shallow water Antarctic seapen Malacobelemnon daytoni, which is in the same family as Kophobelemnon stelliferum (Kophobelemnidae), has indicated that several stress responses occur under acidic conditions, along with up-regulated genes (Natalia Servetto, 2019, pers. comm., 15th October). Kophobelemnon stelliferum however is known to occur naturally in the slightly acidic waters around mud volcanoes (Rueda et al., 2016; Sitjà et al., 2019). For example, in the Spanish mud volcano fields, pH was measured by Rueda et al. (2012) to vary from 7.4 to 6.8. Kita et al. (2015) studied the effects of CO2 gas released from the seabed at a depth of 12 m in Scotland. The seapen Virgularia mirabilis (which also occurs in the deep-sea) was frequently observed in the study area, however no abnormal behaviour was observed in response to the CO2, either during or after release. The authors note that the elevated pCO2 concentrations (<1599 µatm, at 30 cm above the seabed) returned to background levels (360-370 µatm) very quickly. Therefore, Virgularia mirabilis was not necessarily exposed to continuous low pH conditions.

As the oceans absorb carbon dioxide from the atmosphere, leading to a decrease in pH and an increase in acidity, there is a further concern; the shoaling of the aragonite saturation horizon (ASH). The ASH is defined as the depth in the oceans at which aragonite saturation equals 1. Below this depth, aragonite saturation will fall below 1, and dissolution of calcified structures may occur. Currently the depth of the ASH in the North Atlantic is approximately 2000m (Jiang et al., 2015). This depth has already become 80-150m shallower over the past two centuries (Chung et al., 2003, Feely et al., 2004). By the end of this century, the depth of the ASH is expected to become shallower still, reaching depths of up to 400 m under the high emission scenario (RCP 8.5) and 600 m for the middle emission scenario (RCP 4.5) (Zheng & Long, 2014).

In addition to the shallowing of the aragonite saturation horizon, the calcite saturation horizon will also become shallower as ocean acidification occurs (Feely et al., 2004). The calcite saturation horizon is more relevant for octocorals, as they require calcium carbonate for structural development and calcite is essential in the skeletal structure (specifically for their sclerites; (Bayer and Macintyre, 2001). Yesson et al. (2012) found that a 12% of records of octocorals occurred in water undersaturated with aragonite (<1). For Sessiliflorae (seapens), 15-20% of observations were below the aragonite saturation horizon.  Calcite saturation state was the second most important factor determining the model for Pennatulacea (after temperature, although it is noted that all environmental variables are intercorrelated). For the Sessiliflorae, 6% were found in areas undersaturated for calcite, however this was affected by a shallow water sampling bias.

Sensitivity assessment. Kophobelemnon stelliferum has a fleshy structure where its sclerites are protected by the surrounding tissue. It is therefore possible that the species can tolerate a decrease in pH. This is supported by evidence showing that the species occurs in the acidic waters associated with mud volcanoes (pH 7.4 to 6.8; Rueda et al., 2016, 2012; Sitjà et al., 2019). However, there is some evidence that suggests that another (shallow-water) species in the Kophobelemnidae family shows stress responses and up-regulated genes in response to ocean acidification (Natalia Servetto, 2019, pers. comm., 15th October). Under the mid and high emission scenarios, resistance is assessed as ‘High’, as no mortality has been observed. Resilience is assessed as ‘High’ and the biotopes are assessed as ‘Not sensitive’ at the middle and high emmission scenario pressure benchmarks.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: 
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

The Kophobelemnon field biotopes occur in the Atlantic mid and upper bathyal zone, at depths of 200 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

The Kophobelemnon field biotopes occur in the Atlantic mid and upper bathyal zone, at depths of 200 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

The Kophobelemnon field biotopes occur in the Atlantic mid and upper bathyal zone, at depths of 200 – 1300 m. Therefore, the biotope will not be affected by changes arising from sea-level rise, and the assessment at the pressure benchmark is ‘Not relevant’.

Hydrological Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Chemical Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Not assessed

Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not assessed

Physical Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Biological Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Bibliography

  1. Musgrave, E.M., 1909. Memoirs: Experimental Observations on the Organs of Circulation and the Power of Locomotion in Pennatutlids. Journal of Cell Science, 2 (215), 443-481.
  2. Baker, K.D., Wareham, V.E., Snelgrove, P.V.R., Haedrich, R.L., Fifield, D.A., Edinger, E.N., Gilkinson, K.D., 2012. Distributional patterns of deep-sea coral assemblages in three submarine canyons off Newfoundland, Canada. Marine Ecology Progress Series, 445, 235–249. https://doi.org/10.3354/meps09448

  3. Bastari, A., Pica, D., Ferretti, F., Micheli, F. & Cerrano, C., 2018. Sea pens in the Mediterranean Sea: habitat suitability and opportunities for ecosystem recovery. ICES Journal of Marine Science, 75 (5), 1722-1732. DOI https://doi.org/10.1093/icesjms/fsy010

  4. Bayer, F.M., Macintyre, I.G., 2001. The mineral component of the axis and holdfast of some gorgonacean octocorals (Coelenterata: Anthozoa), with special reference to the family Gorgoniidae. Proceeding of the Biological Society of Washington, 114, 309–345.

  5. Birkeland, C., 1974. Interactions between a seapen and seven of its predators. Ecological Monographs, 44, 211-232. DOI https://doi.org/10.2307/1942312

  6. Bluhm, H., 2001. Re-establishment of an abyssal megabenthic community after experimental physical disturbance of the seafloor. Deep Sea Research Part II: Topical Studies in Oceanography, 48 (17), 3841-3868. DOI https://doi.org/10.1016/S0967-0645(01)00070-4

  7. Briggs, K.B., Richardson, M.D. & Young, D.K., 1996. The classification and structure of megafaunal assemblages in the Venezuela Basin, Caribbean Sea. Journal of Marine Research, 54 (4), 705-730. DOI https://doi.org/10.1357/0022240963213736

  8. Chia, F.S. & Crawford, B.J., 1973. Some observations on gametogenesis, larval development and substratum selection of the sea pen Ptilosarcus guerneyi. Marine Biology, 23, 73-82. DOI https://doi.org/10.1007/BF00394113

  9. Chimienti, G., Angeletti, L. & Mastrototaro, F., 2018. Withdrawal behaviour of the red sea pen Pennatula rubra (Cnidaria: Pennatulacea). The European Zoological Journal, 85 (1), 64-70. DOI https://doi.org/10.1080/24750263.2018.1438530

  10. De Clippele, L.H., Buhl-Mortensen, P. & Buhl-Mortensen, L., 2015. Fauna associated with cold water gorgonians and sea pens. Continental Shelf Research, 105, 67-78. DOI https://doi.org/10.1016/j.csr.2015.06.007

  11. de Moura Neves, B., Edinger, E., Hayes, V.W., Devine, B., Wheeland, L. & Layne, G., 2018. Size metrics, longevity, and growth rates in Umbellula encrinus (Cnidaria: Pennatulacea) from the eastern Canadian Arctic. Arctic Science, 4 (4), 722-749. DOI https://doi.org/10.1139/as-2018-0009

  12. Eckelbarger, K.J., Tyler, P.A. & Langton, R.W., 1998. Gonadal morphology and gametogenesis in the sea pen Pennatula aculeata (Anthozoa: Pennatulacea) from the Gulf of Maine. Marine Biology, 132 (4), 677-690. DOI https://doi.org/10.1007/s002270050432

  13. Eno, N.C., MacDonald, D. & Amos, S.C., 1996. A study on the effects of fish (Crustacea/Molluscs) traps on benthic habitats and species. Final report to the European Commission. Study Contract, no. 94/076.

  14. Feely, R.A., Sabine, C.L., Lee, K., Berelson, W., Kleypas, J., Fabry, V.J. & Millero, F.J., 2004. Impact of Anthropogenic CO2 on the CaCO3 System in the Oceans. Science, 305 (5682), 362-366. DOI https://doi.org/10.1126/science.1097329

  15. Gabay, Y., Fine, M., Barkay, Z. & Benayahu, Y., 2014. Octocoral Tissue Provides Protection from Declining Oceanic pH. PLoS ONE, 9 (4), e91553. DOI https://doi.org/10.1371/journal.pone.0091553

  16. García-Alegre A, Murillo FJ, Sacau M, Kenchington E, Serrano A, Durán Muñoz P., 2018 (in press). Trait-based approach on deep-sea corals in the high-seas of the Flemish Cap and Flemish Pass (northwest Atlantic).  PeerJ Preprints 6:e26698v1 https://doi.org/10.7287/peerj.preprints.26698v1
  17. Georgian, S.E., Anderson, O.F. & Rowden, A.A., 2019. Ensemble habitat suitability modeling of vulnerable marine ecosystem indicator taxa to inform deep-sea fisheries management in the South Pacific Ocean. Fisheries Research, 211, 256-274. DOI https://doi.org/10.1016/j.fishres.2018.11.020

  18. Gormley, K.S.G., Hull, A.D., Porter, J.S., Bell, M.C. & Sanderson, W.G., 2015. Adaptive management, international co-operation and planning for marine conservation hotspots in a changing climate. Marine Policy, 53, 54-66. DOI https://doi.org/10.1016/j.marpol.2014.11.017

  19. Greathead, C., González-Irusta, J.M., Clarke, J., Boulcott, P., Blackadder, L., Weetman, A. & Wright, P.J., 2015. Environmental requirements for three sea pen species: relevance to distribution and conservation. ICES Journal of Marine Science: Journal du Conseil, 72 (2), 576-586.

  20. Hogan, R.I., Hopkins, K., Wheeler, A.J., Allcock, A.L. & Yesson, C., 2019. Novel diversity in mitochondrial genomes of deep-sea Pennatulacea (Cnidaria: Anthozoa: Octocorallia). Mitochondrial DNA Part A, 30 (6), 764-777. DOI https://doi.org/10.1080/24701394.2019.1634699

  21. Hughes, D.J., 1998a. Sea pens & burrowing megafauna (volume III). An overview of dynamics and sensitivity characteristics for conservation management of marine SACs. Natura 2000 report prepared for Scottish Association of Marine Science (SAMS) for the UK Marine SACs Project., Scottish Association for Marine Science. (UK Marine SACs Project). Available from:  http://www.ukmarinesac.org.uk/publications.htm

  22. JNCC, 2015. The Marine Habitat Classification for Britain and Ireland Version 15.03. (20/05/2015). Available from https://mhc.jncc.gov.uk/

  23. Kastendiek, J., 1976. Behavior of the sea pansy Renilla kollikeri pfeffer (Coelenterata: Pennatulacea) and its influence on the distribution and biological interactions of the species. The Biological Bulletin, 151 (3), 518-537. DOI https://doi.org/10.2307/1540503
  24. Kita, J., Stahl, H., Hayashi, M., Green, T., Watanabe, Y. & Widdicombe, S., 2015. Benthic megafauna and CO2 bubble dynamics observed by underwater photography during a controlled sub-seabed release of CO2. International Journal of Greenhouse Gas Control, 38, 202-209. DOI https://doi.org/10.1016/j.ijggc.2014.11.012
  25. López-González, P.J., Gili, J.-M. & Fuentes, V., 2009. A new species of shallow-water sea pen (Octocorallia: Pennatulacea: Kophobelemnidae) from Antarctica. Polar Biology, 32 (6), 907-914. DOI https://doi.org/10.1007/s00300-009-0591-8
  26. López-González, P.J., Gili, J.M. & Williams, G.L., 2001. New records of Pennatulacea (Anthozoa: Octocorallia) from the African Atlantic coast, with description of a new species and a zoogeographic analysis. Scientia Marina, 65 (1), 59-74. DOI https://doi.org/10.3989/scimar.2001.65n159
  27. Langton, R., W. Langton, E., B. Theroux, R. & R. Uzmann, J., 1990. Distribution, behavior and abundance of sea pens, Pennatula aculeata, in the Gulf of Maine. Marine Biology, 107, 463-469. DOI https://doi.org/10.1007/BF01313430

  28. Levitan, D.R., 1996. Effects of gamete traits on fertilization in the sea and the evolution of sexual dimorphism. Nature, 382 (6587), 153. DOI https://doi.org/10.1038/382153a0

  29. Lindholm, J., Kelly, M., Kline, D. & de Marignac, J., 2008. Patterns in the Local Distribution of the Sea Whip, Halipteris willemoesi, in an Area Impacted by Mobile Fishing Gear. Marine Technology Society Journal, 42 (4), 64-68.
  30. Mastrototaro, F., Chimienti, G., Capezzuto, F., Carlucci, R. & Williams, G., 2015. First record of Protoptilum carpenteri (Cnidaria: Octocorallia: Pennatulacea) in the Mediterranean Sea. Italian Journal of Zoology, 82 (1), 61-68. DOI https://doi.org/10.1080/11250003.2014.982218
  31. Mastrototaro, F., Maiorano, P., Vertino, A., Battista, D., Indennidate, A., Savini, A., Tursi, A. & D'Onghia, G., 2013. A facies of Kophobelemnon (Cnidaria, Octocorallia) from Santa Maria di Leuca coral province (Mediterranean Sea). Marine Ecology, 34 (3), 313-320. DOI https://doi.org/10.1111/maec.12017
  32. Murillo, F.J., MacDonald, B.W., Kenchington, E., Campana, S.E., Sainte-Marie, B. & Sacau, M., 2018. Morphometry and growth of sea pen species from dense habitats in the Gulf of St. Lawrence, eastern Canada. Marine Biology Research, 14 (4), 366-382. DOI https://doi.org/10.1080/17451000.2017.1417604

  33. Neves, B.d.M., Edinger, E., Layne, G.D. & Wareham, V.E., 2015. Decadal longevity and slow growth rates in the deep-water sea pen Halipteris finmarchica (Sars, 1851) (Octocorallia: Pennatulacea): implications for vulnerability and recovery from anthropogenic disturbance. Hydrobiologia, 759 (1), 147-170. DOI https://doi.org/10.1007/s10750-015-2229-x

  34. Pires, D., Castro, C. & Silva, J., 2009. Reproductive biology of the deep-sea pennatulacean Anthoptilum murrayi (Cnidaria, Octocorallia). Marine Ecology Progress Series, 397, 103-112. DOI https://doi.org/10.3354/meps08322
  35. Rice, A.L., Tyler, P.A. & Paterson, G.J.L., 1992. The Pennatulid Kophobelemnon stelliferum (Cnidaria: Octocorallia) in the Porcupine Seabight (north-east Atlantic Ocean). Journal of the Marine Biological Association of the United Kingdom, 72 (02), 417. DOI https://doi.org/10.1017/S0025315400037796

  36. Rueda, J.L., González-García, E., Krutzky, C., López-Rodriguez, F.J., Bruque, G., López-González, N., Palomino, D., Sánchez, R.F., Vázquez, J.T., Fernández-Salas, L.M. & Díaz-del-Río, V., 2016. From chemosynthesis-based communities to cold-water corals: Vulnerable deep-sea habitats of the Gulf of Cádiz. Marine Biodiversity, 46 (2), 473-482. DOI https://doi.org/10.1007/s12526-015-0366-0
  37. Rueda, J.L., Urra, J., Gofas, S., Lopez-Gonzalez, N., Fernandez-Salas, L.M. & Diaz-Del-Rio, V., 2012. New records of recently described chemosymbiotic bivalves for mud volcanoes within the European waters (Gulf of Cádiz). Mediterranean Marine Science, 13 (2), 262-267. DOI https://doi.org/10.12681/mms.307
  38. Schubert, N., Brown, D. & Rossi, S., 2017. Symbiotic Versus Nonsymbiotic Octocorals: Physiological and Ecological Implications. In Rossi, S., et al. (eds.). Marine Animal Forests: The Ecology of Benthic Biodiversity Hotspots, Cham: Springer International Publishing, pp. 887-918.
  39. Servetto, N. & Sahade, R., 2016. Reproductive Seasonality of the Antarctic Sea Pen Malacobelemnon daytoni (Octocorallia, Pennatulacea, Kophobelemnidae). PLOS ONE, 11 (10). DOI https://doi.org/10.1371/journal.pone.0163152

  40. Simon-Lledó, E., Bett, B.J., Huvenne, V.A.I., Köser, K., Schoening, T., Greinert, J. & Jones, D.O.B., 2019. Biological effects 26 years after simulated deep-sea mining. Scientific reports, 9 (1). DOI https://doi.org/10.1038/s41598-019-44492-w
  41. Sitjà, C., Maldonado, M., Farias, C. & Rueda, J.L., 2019. Deep-water sponge fauna from the mud volcanoes of the Gulf of Cadiz (North Atlantic, Spain). Journal of the Marine Biological Association of the United Kingdom, 99 (4), 807-831. DOI https://doi.org/10.1017/S0025315418000589
  42. Tyler, P.A., Bronsdon, S.K., Young, C.M. & Rice, A.L., 1995. Ecology and Gametogenic Biology of the Genus Umbellula (Pennatulacea) in the North Atlantic Ocean. Internationale Revue der gesamten Hydrobiologie und Hydrographie, 80 (2), 187-199.
  43. Wareham, V.E. & Edinger, E.N., 2007. Distribution of deep-sea corals in the Newfoundland and Labrador region, Northwest Atlantic Ocean. Bulletin of Marine Science, 81 (3), 289-313.

  44. Wilson, M.T., Andrews, A.H., Brown, A.L. & Cordes, E.E., 2002. Axial rod growth and age estimation of the sea pen, Halipteris willemoesi Kölliker Hydrobiologia, 471, 133-142.

  45. Yesson, C., Taylor, M.L., Tittensor, D.P., Davies, A.J., Guinotte, J., Baco, A., Black, J., Hall-Spencer, J.M. & Rogers, A.D., 2012. Global habitat suitability of cold-water octocorals: global distribution of deep-sea octocorals. Journal of Biogeography, 39 (7), 1278-1292. DOI https://doi.org/10.1111/j.1365-2699.2011.02681.x

Citation

This review can be cited as:

Last, E.K., Baron-Cohen, L. & Robson, L.M. 2019. [Kophobelemnon] field on Atlantic mid bathyal mud. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 24-01-2020]. Available from: https://www.marlin.ac.uk/habitat/detail/1197

Last Updated: 22/11/2019