MarLIN

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

Discrete Lophelia pertusa colonies on Atlantic upper bathyal rock and other hard substrata

Summary

UK and Ireland classification

UK and Ireland classification

Description

This biotope occurs on bedrock, cobbles and boulders and isolated drop stones. Small growths of Lophelia pertusa and often Madrepora occulata are present. A Lophelia pertusa reef framework may also be present adjacent to this biotope in areas where it occurs in an elevated position allowing dead coral framework to accumulate below. This biotope is often observed on the edge of escarpment features. The same assemblage is recorded on Atlantic mid bathyal coarse sediment. (Information from JNCC, 2015).

Depth range

200-600 m

Additional information

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Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

The biotope represents the assemblages occurring on rock in the Atlantic upper bathyal zone. Assemblages occur on rock and other hard substrata within the upper bathyal zone (M.AtUB.Ro.MixCor.DisLop biotope) and the mid bathyal zone (M.AtMB.Ro.MixCor.DisLop biotope), as well as on coarse substrates in both the upper and mid-bathyal zones (M.AtUB.Co.MixCor.DisLop and M.AtMB.Co.MixCor.DisLop respectively). Therefore, the sensitivity of these discrete Lophelia pertusa colony biotopes is 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 predominant species for the biotopes are the scleractinian corals Lophelia pertusa and Madrepora oculata. Loss of these species may result in loss or degradation of the biotope, therefore, the sensitivity of the biotope is dependent on the sensitivity of both these species. Other species that can be found within this biotope include; Actiniaria (including Phelliactis) and Ceriantharia anemones, massive lobose, branching and encrusting (orange, and pale – yellow, white, grey, cream, pink) sponges, cup corals Caryophylliidae, decapods Munida, Paguridae and Pandalus borealis, bivalve molluscs Anomiidae, Ophiuroidea brittle stars, the echinoid Cidaris cidaris, the holothurian Psolus squamatus, Serpulidae and Aberrantidae polychaete worms, the soft coral Anthomastus, and Hydrozoa (including Pliobothrus). Most of these are ubiquitous and not unique to these biotopes. They are, therefore, not considered significant to the assessment of sensitivity. Furthermore, the presence of all these species is not essential for the classification of these biotopes. 

Resilience and recovery rates of habitat

Lophelia pertusa is a cold-water reef forming scleractinian coral with a worldwide distribution. This species is most abundant in deep waters, at high latitudes in the North East Atlantic. Global oceanographic data show that Lophelia pertusa is found at a mean depth of 480 m (Davies et al., 2008). When it occurrs in discrete colonies, it is usually found in either cauliflower-like, or bush-like colonies (Vad et al., 2017). Madrepora oculata is a scleractinian coral, which forms fragile fan-shaped or cauliflower-like colonies of approximately 30 to 50 cm in height (Tsounis et al., 2010). It has small polyps, 3 to 5mm in diameter (Tsounis et al., 2010). Colonial structures or ‘coral bushes’ can also exist where both Madrepora oculata and Lophelia pertusa are growing together, rather than being separate coral bushes (Arnaud-Haond et al., 2017).

Distribution and Habitat. Lophelia pertusa is a cosmopolitan species, with records from north Norwegian waters, the Gulf of Mexico, the eastern Pacific and the Mediterranean (OBIS, 2018). Lophelia pertusa is a reef forming scleractinian coral, although discrete colonies occur, either through the development of Lophelia colonies on vertical substratum, where coral rubble cannot aggregate and therefore reefs are unable to develop, or through the presence of small colonies on boulders, cobbles and isolated drop-stones.  Discrete Lophelia pertusa colonies have been found on bedrock walls of seamounts, canyons, and fjords (Buhl-Mortensen et al., 2017), alongside colonizing oil and gas infrastructure (Gass & Roberts, 2006). On the Mingulay Reef complex colonies of discrete Lophelia pertusa can be found on steep rocky banks, alongside small colonies of Madrepora occulata and the solitary coral Caryophyllia sp. (Howell, 2010).

Madrepora oculata shares this global distribution. Madrepora oculata occurs at a depth range of ca. 50 to >1500 m (Hansson et al., 2009).  However depth is not a principal environmental factor in determining its distribution. Naumann et al. (2014) states that suitable substratum availability, current velocity, food supply and aragonite saturation state are the key factors affecting the occurrence and local abundance of cold-water corals. The gradients of seawater density, as a function of seawater temperature and salinity, was also noted to be important.

Cheung et al. (2005) analysed the distribution of cold-water corals and found that fields of Madrepora oculata usually occurred at the foothills of large seamounts, at a distance of 50-100 km. Madrepora oculata has a preference for hard substratum bottoms (i.e. boulders and hard rock outcrops).  Its presence was one order of magnitude higher in hard substratum areas compared to those with soft bottom substrata (Orejas et al., 2009). In the Gulf of Mexico, Lophelia pertusa has been found to colonize an artificial hard substratumat a depth of 204-292 m (Schroeder et al., 2005), whereas in the North Sea it has been found to colonize oil and gas infrastructure between 47 – 132 m depth (Gass & Roberts, 2006).

Reproduction and Development. Colonial scleractinians corals, such as Madrepora oculata, exhibit both sexual and asexual reproduction (Brooke and Järnegren, 2013). Asexual replication of polyps is important in the development of reefs (Goff-Vitry et al., 2004; Le Goff-Vitry and Rogers, 2005) and is well documented for Lophelia pertusa where is occurs by fragmentation (Dahl et al., 2013, 2012; Larsson et al., 2014), with replication occurring by intratentacular budding (Cairns, 1979, 1994, Roberts et al., 2009, Brooke & Järnegren, 2013). However, studies on the asexual reproduction of Madrepora oculata are limited. Evidence of asexual reproduction was found in large fragments of cold-water corals off New Zealand, where intratentacular budding was evident for Solenosmilia variabilis and extratentacular budding for Goniocorella dumosa. However, their samples of Madrepora oculata were very small (3-15 polyps) and showed no evidence of asexual reproduction.

Lophelia pertusa is gonochoristic and is thought to spawn annually (Waller, 2005). Evidence from the North East Atlantic Lophelia pertusa supports this supposition, and samples collected within this area showed a seasonal reproductive cycle with a single cohort per year, and a spawning event around February (Waller & Tyler, 2005). Larsson et al. (2014) examined embryogenesis and larval development in the laboratory in fragments of live Lophelia pertusa colonies from the Tisler reef and Trondheim Fjord, Norway. Spawning occurred in Jan to March, although spawning was asynchronous depending on site of origin, over a period of two months. They observed that multiple male polyps spawned simultaneously, resulting in a high fertilization efficiency. Spawned oocytes were 160 µm in diameter and resultant embryos were neutral or negatively buoyant and developed into 120-270 µm long ciliated planulae. The planulae were active swimmers (0.5 mm/s) and actively swam upwards into the upper water column. Larsson et al. (2014) estimated that larvae could vertically migrate ca 50 m/day, which would bring them out of the benthic boundary layer in reef conditions. The planulae spent three to five weeks in the water column before the onset of bottom-probing behaviour. Larsson et al. (2014) concluded that the larvae were probably planktotrophic rather than lecithotrophic suggested by Waller (2005). Bottom-probing behaviour became common amongst the larvae studied four to five weeks after fertilization and coincided with the development of nematocysts, which suggested that it had become competent, although settlement was not observed. The planula larvae of Lophelia pertusa require hard substrata for settlement, including rock surfaces, artificial substrata, coral fragments or hydrocarbon seep associated carbonates. In sedimentary areas, Lophelia pertusa may settle on hard substrata as small as a shell, pebble, or worm tube (Rogers, 1999). However, a hard substratum is a pre-requisite for settlement and a layer of sediment may interfere with settlement and hence recruitment.

Evidence suggests that Madrepora oculata has two cohorts of gamete production, with a total fecundity of around 60 oocytes per polyp, however it is uncertain whether they exhibit seasonal reproduction (Lartaud et al., 2014; Waller and Tyler, 2005). Based on the large oocytes of Madrepora oculata (405 µm) and the timing of reproduction, Waller and Tyler (2005) inferred that is had a lecithotrophic larva. Madrepora oculata may, therefore, respond to environmental cues when conditions are suitable for reproduction, thereby by producing and spawning gametes on a seasonal basis (Lartaud et al., 2014). In the Seabight area, reproduction of Madrepora oculata is reported to have a seasonality that fits with the phytodetrital food fall that occurs around July (Waller and Tyler, 2005). A study by Pires et al. (2014) indicated that Madrepora oculata is a broadcast spawner, as no embryos or larvae were observed in their samples, and the species also presented continuous reproduction. Madrepora oculata is thought to be gonochoric (Burgess and Babcock, 2005; Pires et al., 2014), although some colonies off Brazil were found by Pires et al. (2014) to present different hermaphroditism patterns.

In a study on Lophelia pertusa, larvae survived for up to 1 year under laboratory conditions, even without regular feeding or water change, indicating the high potential for long distance dispersal (Strömberg and Larsson, 2017), although migration is not sufficient to counteract reproductive isolation of populations (Dr Alex Rogers, 2005 pers comm.). Molecular genetic data indicates that Beryl oil fields samples of Lophelia are closely related to northern Rockall Trough populations but that there is strong genetic differentiation (population sub-division), with very low gene flow between areas (Goff-Vitry et al., 2004; Dr Alex Rogers, 2005 pers comm.). Molecular genetic studies of the population of Lophelia pertusa in the North East Atlantic showed that it was not a panmictic population but composed of genetically distinct offshore and fjordic subpopulations from the Iberian margin to the Scandinavian fjords (Le Goff-Vitry & Rogers, 2005). Also, inbreeding was also observed in some subpopulations that indicated self-recruitment in those sites. In addition, there was high variation in the degree of genetic variation between subpopulations. For example, the Darwin mounds exhibited a high proportion of clones and low genetic diversity.  In particular, the fjordic populations were highly differentiated genetically, for example, the Osterfjord subpopulation showed very low genetic diversity.  Morrison et al. (2011) also found genetic differentiation between populations of Lophelia pertusa in the Gulf of Mexico, coastal southeast United States, New England seamounts and the eastern North Atlantic. They concluded that, while some larvae were dispersed over large geographic distances, gene flow between the ocean regions was restricted. The evidence suggests that asexual reproduction predominates in reef growth and that the contribution from larvae may be limited (Dr Alex Rogers, 2005 pers comm.), leading to the conclusion that gene flow along the continental margin iss sporadic and that recolonization of disturbed coral reefs through larval dispersal is likely to take long periods of time (Goff-Vitry et al., 2004, Le Goff-Vitry & Rogers, 2005).

Gass & Roberts (2006) examined 14 oil and gas platforms within the North Sea and found Lophelia pertusa to be growing on 13 of them.  Two of the platforms were examined more closely and 947 individual colonies were found, the largest of which was 132 cm in diameter (Gass & Roberts, 2006). Prior to the oil and gas platforms in the North Sea, there were no known records of live Lophelia pertusa.  Larvae recruited to these North Sea platforms were probably transported in the North Atlantic water mass entering the North Sea.  The nearest known Lophelia pertusa colonies to the North Sea are from the west coast of Scotland. Lophelia pertusa larvae are most likely to have reached the North Sea via the substantial inflow of Atlantic water flowing southwards east of Shetland from the Atlantic shelf edge current and the Fair Isle Current (Roberts, 2002, taken from Gass & Roberts, 2006).

Growth rates and longevity. Orejas et al. (2011) investigated the growth rates of Madrepora oculata and Lophelia pertusa under control conditions (12°C in the dark, being fed five times a week) over eight months. In terms of total weight increase, they found that Madrepora oculata had a mean growth rate of 0.11% per day (± 0.04 SD).  This was significantly (p < 0.0001) higher than that of Lophelia pertusa, measured at 0.02% (± 0.01 SD). Linear growth was also recorded by Orejas et al. (2011) and was found to be 5.11(± 2.56 SD) and 8.76 (± 6.57 SD) mm/ yr for Madrepora oculata and Lophelia pertusa, respectively. In an in situ growth rate study in the Lacaze-Duthiers canyon in the northwestern Mediterranean Sea (at 520 m depth), Madrepora oculata had a faster growth rate in summer (5.8 mm per year) than in winter (4.1 mm per year, whereas Lophelia pertusa had slightly higher growth rates in winter (8.4 mm per year) than summer (7.3 mm per year) (Lartaud et al., 2014). The budding rate (the rate of new polyp addition per mother polyp per year) in winter/spring for Madrepora oculata was 45% and for Lophelia pertusa was 48%, but whilst Lophelia pertusa had a slightly elevated budding rate in summer (58%), budding rate was significantly lower for Madrepora oculata in summer (14%). This was thought to be due to the sensitivity of Madrepora oculata to the variability of food supply resulting from seasonal sinking of dense water masses, bring organic matter from surface waters. On oil and gas infrastructure, the growth rate of Lophelia pertusa colonies was calculated to be up to 33 mm/ yr (Gass & Roberts, 2006). In addition to food supply, the growth of deep-water corals is thought to be influenced by a variety of biotic and abiotic factors, including turbidity, temperature, hydrography and seawater chemistry (Lartaud et al., 2014). It is thought that these factors have given rise to the range of growth rates that have been recorded for Madrepora oculata (3-18 mm/year; Lartaud et al., 2014) and Lophelia pertusa (2.4 – 35 mm/ year; Brooke & Young, 2009).

Feeding behaviour. Madrepora oculata and Lophelia pertusa are filter feeders (Duineveld et al., 2007), which feed primarily on zooplankton (mainly phrosinid and platyscelid Amphipods) or phytodetritus (Carlier et al., 2009; Lartaud et al., 2014). However, Lophelia pertusa has also been noted to be a generalist feeder, taking any nutritious particles available (Duineveld et al., 2007) and Duineveld et al. (2004) found no clear-cut single food source for Madrepora oculata and Lophelia pertusa. Madrepora oculata was found by Tsounis et al. (2010) to have a much lower prey capture rate compared to Lophelia pertusa, for both larger plankton (2.38 ±2.31 and 7.82 ±2.49 ind./polyp/hour, respectively) and small plankton (47.91 ±33.29 and 283.73 ±130.09 ind./polyp/hour, respectively). Madrepora oculata often failed to capture adult zooplankton. These differences may be due to differences in feeding strategy (Kiriakoulakis et al., 2005).    

Resilience assessment. From experiments within controlled aquaria, there is evidence that Lophelia pertusa can recover from very small fragments (Maier, 2008), whilst colonization of oil and gas platforms provide evidence that the larvae of Lophelia pertusa have the potential to establish and grow to considerable sizes (≤ 118 cm) within 20-25 years (Gass & Roberts, 2006). It is now widely accepted that anthropogenic pressures are having a negative effect on cold-water corals, including Lophelia pertusa (Roberts & Cairns, 2014). However, the limited knowledge regarding the worldwide distribution of the cold-water coral habitats makes it very difficult to determine how much habitat has been lost to anthropogenic pressures. Lophelia pertusa has also been identified within the by-catch of deep-water fishing vessels trawling off the west coast of Ireland (Hall-Spencer et al., 2002).  In addition to deep water fisheries, the hydrocarbon industry, mining, and ocean acidification have all been found to degrade the health of cold-water corals (Roberts et al., 2009). Huvenne et al. (2016) found that eight years after bottom trawling impacts, minimal live coral was seen in an area of the Darwin Mounds in UK waters, with the exception of a few small colonies, suggesting slow recovery rates even for small colonies of the species. For a 45.5 cm long branch of Madrepora occulata, Sabatier et al. (2012) determined an age of 31 years. Growth rates of Madrepora oculata range between 3-18 mm/year (Lartaud et al., 2014) and 2.4 – 35 mm/ year for Lophelia pertusa (Brooke & Young, 2009).  These growth rates suggest that a colony could grow to between 60 – 875 mm in 25 years. Whilst recovery of this biotope is possible, it is likely to take up to 25 years to recover structure and function, hence where resistance is ‘None’, ‘Low’, ‘Medium’, resilience is assessed ‘Low’. 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: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

Deep waters off the continental shelf (200 – 2,500 m) are expected to see a lower temperature rise (≈ 1oC) than shallow water habitats by the end of this century, regardless of scenario (FAO (Fisheries and Aquaculture Organisation), 2019).  Lophelia pertusa is typically found in areas where temperatures range from 4 to 12°C (Davies et al., 2008, Lunden et al., 2014),  with UK reefs and colonies generally in water temperatures of 6-10°C (Frederiksen et al., 1992, Duineveld et al., 2007, Roberts et al., 2009). The upper lethal limit for this species is thought to be 15°C (Brooke et al., 2013). Madrepora oculata is found in temperatures varying from approximately 5°C in the NE Atlantic (Wild et al., 2008) to 13.9°C in the Mediterranean (Freiwald et al., 2009).

Whilst temperatures are thought to generally be very stable in Lophelia pertusa environments (Dullo et al., 2008), there are occasions when substantial increases in temperature can be observed. For example on the Tisler Reef, in Norway, in 2006 and 2008 the temperature rose by approximately 4°C in 24 hours, spiked at 12°C and remained above 10°C for approximately 30 days (Guihen et al., 2012). No Lophelia pertusa mortality was observed during this time, although mass mortality of the deepwater sponge, Geodia baretti, occurred (Guihen et al., 2012). Furthermore, over coral mounds off the coast of North Carolina, US, incursion of the Gulf stream  can lead to temperatures fluctuating greatly, reaching 15°C on a regular basis, from a mean bottom water temperature of 9.9°C (Brooke et al., 2013). ). In the Gulf of Mexico, Lophelia pertusa occurred in areas where temperatures were 8.5-10.6°C (Davies et al., 2010). Temperature variations of 0.8°C were observed over 5-11 hr periods, associated with internal waves. Furthermore, high-frequency temperature variability over 20-30-minute periods was also recorded at one of coral sites (476m depth), where a temperature rise of 0.5°C occurred, followed by a slower temperature decline. These fluctuations exhibit 5-11 hour cycles. These observations suggest that this species can tolerate increases in temperature.

Results of the effect of short-term (≤6 months) experimental increases in temperature on Lophelia pertusa have been somewhat mixed, with some finding this species sensitive to increased temperatures (Dodds et al., 2007, Lunden et al., 2014), and others showing positive effects of increases in temperature (Büscher et al., 2017). Buscher et al. (2017) found an increase in growth rates in Norwegian populations in response to a six month increase in temperature from 8-12°C. In the study by Lunden et al. (2013) a 2°C increase in temperature (from 8-10°C) saw a 10% decrease in survivorship, although some genotypes were more sensitive than others to the increase in temperature. Naumann et al. (2014) examined the respiration rate and calcification rates of Lophelia pertusa and Madrepora oculata collected from the Mediterranean at 12, 9 and 6°C after acclimation for one month. Lophelia pertusa was found to acclimate to lower temperatures (9 and 6°C) and maintained a constant respiration rate although calcification rates were reduced by 58% at 12°C. On the other hand, Madrepora oculata respiration decreased at the lower temperatures. Madrepora oculata appears to exhibit greater growth at temperatures towards the upper end of its temperature range (Naumann et al., 2014), which may be why this species is more abundant in the Mediterranean (Arnaud-Haond et al., 2017).

Sensitivity assessment. Under all three scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C, increasing the temperature at which UK Lopelia pertusa and Madrepora oculata colonies are found to between 6-11°C, which is not outside their current temperature distribution. Whilst an increase of 1°C is not expected to cause catastrophic effects, laboratory experiments suggest an increase in temperature may lead to some stress on Lophelia pertusa, potentially leading to changes in feeding, and respiration rates. It must be taken into consideration that any negative impacts observed in laboratory experiments were observed when temperatures were increased suddenly by 2°C or more, and experiments run over relatively short timescales (< 6 months). Under future projections for all scenarios, the temperature increase in deep waters is expected to rise by only 1°C, with this increase occurring over the next 50-80 years, giving NE Atlantic populations of Lophelia pertusa the chance to adapt, as it has to warmer water temperatures in the Gulf of Mexico and the Mediterranean. Madrepora oculata appears to respond positively to an increase in temperature, and therefore is not expected to be sensitive to a 1°C increase in temperature. For this reason, the resistance has been assessed as ‘High’, whilst resilience has been assessed ‘High’. This leads to an overall sensitivity score of ‘Not sensitive’ at this level benchmark.

High High Not sensitive
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

Deep waters off the continental shelf (200 – 2,500 m) are expected to see a lower temperature rise (≈ 1oC) than shallow water habitats by the end of this century, regardless of scenario (FAO (Fisheries and Aquaculture Organisation), 2019).  Lophelia pertusa is typically found in areas where temperatures range from 4 to 12°C (Davies et al., 2008, Lunden et al., 2014),  with UK reefs and colonies generally in water temperatures of 6-10°C (Frederiksen et al., 1992, Duineveld et al., 2007, Roberts et al., 2009). The upper lethal limit for this species is thought to be 15°C (Brooke et al., 2013). Madrepora oculata is found in temperatures varying from approximately 5°C in the NE Atlantic (Wild et al., 2008) to 13.9°C in the Mediterranean (Freiwald et al., 2009).

Whilst temperatures are thought to generally be very stable in Lophelia pertusa environments (Dullo et al., 2008), there are occasions when substantial increases in temperature can be observed. For example on the Tisler Reef, in Norway, in 2006 and 2008 the temperature rose by approximately 4°C in 24 hours, spiked at 12°C and remained above 10°C for approximately 30 days (Guihen et al., 2012). No Lophelia pertusa mortality was observed during this time, although mass mortality of the deepwater sponge, Geodia baretti, occurred (Guihen et al., 2012). Furthermore, over coral mounds off the coast of North Carolina, US, incursion of the Gulf stream  can lead to temperatures fluctuating greatly, reaching 15°C on a regular basis, from a mean bottom water temperature of 9.9°C (Brooke et al., 2013). ). In the Gulf of Mexico, Lophelia pertusa occurred in areas where temperatures were 8.5-10.6°C (Davies et al., 2010). Temperature variations of 0.8°C were observed over 5-11 hr periods, associated with internal waves. Furthermore, high-frequency temperature variability over 20-30-minute periods was also recorded at one of coral sites (476m depth), where a temperature rise of 0.5°C occurred, followed by a slower temperature decline. These fluctuations exhibit 5-11 hour cycles. These observations suggest that this species can tolerate increases in temperature.

Results of the effect of short-term (≤6 months) experimental increases in temperature on Lophelia pertusa have been somewhat mixed, with some finding this species sensitive to increased temperatures (Dodds et al., 2007, Lunden et al., 2014), and others showing positive effects of increases in temperature (Büscher et al., 2017). Buscher et al. (2017) found an increase in growth rates in Norwegian populations in response to a six month increase in temperature from 8-12°C. In the study by Lunden et al. (2013) a 2°C increase in temperature (from 8-10°C) saw a 10% decrease in survivorship, although some genotypes were more sensitive than others to the increase in temperature. Naumann et al. (2014) examined the respiration rate and calcification rates of Lophelia pertusa and Madrepora oculata collected from the Mediterranean at 12, 9 and 6°C after acclimation for one month. Lophelia pertusa was found to acclimate to lower temperatures (9 and 6°C) and maintained a constant respiration rate although calcification rates were reduced by 58% at 12°C. On the other hand, Madrepora oculata respiration decreased at the lower temperatures. Madrepora oculata appears to exhibit greater growth at temperatures towards the upper end of its temperature range (Naumann et al., 2014), which may be why this species is more abundant in the Mediterranean (Arnaud-Haond et al., 2017).

Sensitivity assessment. Under all three scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C, increasing the temperature at which UK Lopelia pertusa and Madrepora oculata colonies are found to between 6-11°C, which is not outside their current temperature distribution. Whilst an increase of 1°C is not expected to cause catastrophic effects, laboratory experiments suggest an increase in temperature may lead to some stress on Lophelia pertusa, potentially leading to changes in feeding, and respiration rates. It must be taken into consideration that any negative impacts observed in laboratory experiments were observed when temperatures were increased suddenly by 2°C or more, and experiments run over relatively short timescales (< 6 months). Under future projections for all scenarios, the temperature increase in deep waters is expected to rise by only 1°C, with this increase occurring over the next 50-80 years, giving NE Atlantic populations of Lophelia pertusa the chance to adapt, as it has to warmer water temperatures in the Gulf of Mexico and the Mediterranean. Madrepora oculata appears to respond positively to an increase in temperature, and therefore is not expected to be sensitive to a 1°C increase in temperature. For this reason, the resistance has been assessed as ‘High’, whilst resilience has been assessed ‘High’. This leads to an overall sensitivity score of ‘Not sensitive’ at this level benchmark.

High High Not sensitive
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

Deep waters off the continental shelf (200 – 2,500 m) are expected to see a lower temperature rise (≈ 1oC) than shallow water habitats by the end of this century, regardless of scenario (FAO (Fisheries and Aquaculture Organisation), 2019).  Lophelia pertusa is typically found in areas where temperatures range from 4 to 12°C (Davies et al., 2008, Lunden et al., 2014),  with UK reefs and colonies generally in water temperatures of 6-10°C (Frederiksen et al., 1992, Duineveld et al., 2007, Roberts et al., 2009). The upper lethal limit for this species is thought to be 15°C (Brooke et al., 2013). Madrepora oculata is found in temperatures varying from approximately 5°C in the NE Atlantic (Wild et al., 2008) to 13.9°C in the Mediterranean (Freiwald et al., 2009).

Whilst temperatures are thought to generally be very stable in Lophelia pertusa environments (Dullo et al., 2008), there are occasions when substantial increases in temperature can be observed. For example on the Tisler Reef, in Norway, in 2006 and 2008 the temperature rose by approximately 4°C in 24 hours, spiked at 12°C and remained above 10°C for approximately 30 days (Guihen et al., 2012). No Lophelia pertusa mortality was observed during this time, although mass mortality of the deepwater sponge, Geodia baretti, occurred (Guihen et al., 2012). Furthermore, over coral mounds off the coast of North Carolina, US, incursion of the Gulf stream  can lead to temperatures fluctuating greatly, reaching 15°C on a regular basis, from a mean bottom water temperature of 9.9°C (Brooke et al., 2013). ). In the Gulf of Mexico, Lophelia pertusa occurred in areas where temperatures were 8.5-10.6°C (Davies et al., 2010). Temperature variations of 0.8°C were observed over 5-11 hr periods, associated with internal waves. Furthermore, high-frequency temperature variability over 20-30-minute periods was also recorded at one of coral sites (476m depth), where a temperature rise of 0.5°C occurred, followed by a slower temperature decline. These fluctuations exhibit 5-11 hour cycles. These observations suggest that this species can tolerate increases in temperature.

Results of the effect of short-term (≤6 months) experimental increases in temperature on Lophelia pertusa have been somewhat mixed, with some finding this species sensitive to increased temperatures (Dodds et al., 2007, Lunden et al., 2014), and others showing positive effects of increases in temperature (Büscher et al., 2017). Buscher et al. (2017) found an increase in growth rates in Norwegian populations in response to a six month increase in temperature from 8-12°C. In the study by Lunden et al. (2013) a 2°C increase in temperature (from 8-10°C) saw a 10% decrease in survivorship, although some genotypes were more sensitive than others to the increase in temperature. Naumann et al. (2014) examined the respiration rate and calcification rates of Lophelia pertusa and Madrepora oculata collected from the Mediterranean at 12, 9 and 6°C after acclimation for one month. Lophelia pertusa was found to acclimate to lower temperatures (9 and 6°C) and maintained a constant respiration rate although calcification rates were reduced by 58% at 12°C. On the other hand, Madrepora oculata respiration decreased at the lower temperatures. Madrepora oculata appears to exhibit greater growth at temperatures towards the upper end of its temperature range (Naumann et al., 2014), which may be why this species is more abundant in the Mediterranean (Arnaud-Haond et al., 2017).

Sensitivity assessment. Under all three scenarios (middle and high emission, and extreme scenarios), waters off the continental shelf are expected to increase marginally by approximately 1°C, increasing the temperature at which UK Lopelia pertusa and Madrepora oculata colonies are found to between 6-11°C, which is not outside their current temperature distribution. Whilst an increase of 1°C is not expected to cause catastrophic effects, laboratory experiments suggest an increase in temperature may lead to some stress on Lophelia pertusa, potentially leading to changes in feeding, and respiration rates. It must be taken into consideration that any negative impacts observed in laboratory experiments were observed when temperatures were increased suddenly by 2°C or more, and experiments run over relatively short timescales (< 6 months). Under future projections for all scenarios, the temperature increase in deep waters is expected to rise by only 1°C, with this increase occurring over the next 50-80 years, giving NE Atlantic populations of Lophelia pertusa the chance to adapt, as it has to warmer water temperatures in the Gulf of Mexico and the Mediterranean. Madrepora oculata appears to respond positively to an increase in temperature, and therefore is not expected to be sensitive to a 1°C increase in temperature. For this reason, the resistance has been assessed as ‘High’, whilst resilience has been assessed ‘High’. This leads to an overall sensitivity score of ‘Not sensitive’ at this level benchmark.

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.) Therefore, sensitivity to marine heatwaves is probably ‘Not relevant’ in this bathyal habitat.

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.) Therefore, sensitivity to marine heatwaves is probably ‘Not relevant’ in this bathyal habitat.

Medium Very Low Medium
Q: High
A: High
C: Medium
Q: High
A: High
C: High
Q: High
A: High
C: High

Due to their calcified nature, both deep and shallow water scleractinian corals are generally thought to be highly sensitive to ocean acidification (Hall-Spencer & Harvey, 2019). The deep-water species, Lophelia pertusa, is a plastic species which has a wide bathymetric range (30-3000 m), lives in several different temperate habitats, and can thrive in the Mingulay Reef complex, where tidal downwellings cause daily variations in pH equivalent to a 25 year jump in atmospheric CO2 (Hennige et al., 2015). This suggests that it may have some tolerance to changes in pH.

Results of the impact of short-term experimental acidification on Lophelia pertusa generally show negative effects such as a decrease in respiration rates (Hennige et al., 2014; 0.3 unit decrease), and calcification rates (Maier et al., 2009; both at 0.15 and 0.3 unit decrease), or no observable effects (Maier et al., 2013b). Georgian et al. (2016), found variability in response between populations. For populations in the Gulf of Mexico, decreasing pH led to a decrease in respiration, prey capture rates and net calcification, whilst populations in Norway responded to a decrease in pH by increasing respiration and prey capture rates, leading to increased calcification (Georgian et al., 2016). Longer term studies (≥ 6 months) show potential for acclimation, with results suggesting that Lophelia pertusa is able to maintain calcification at levels of ocean acidification expected for the end in both the mid- and high-emission scenario  (Form & Riebesell, 2012, Maier et al., 2013a, Movilla et al., 2014, Hennige et al., 2015). Experimental exposures have similarly shown that calcification rates (Maier et al., 2012, Maier et al., 2013b) and respiration rates (Maier et al., 2013a) are maintained at ocean acidification levels expected for the high emission scenario in Madrepora oculata.

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 400m under the high emission scenario (RCP 8.5) and 600m for the middle emission scenario (RCP 4.5) (Zheng & Long, 2014).

For both Lophelia pertusa (Form & Riebesell, 2012, Hennige et al., 2015, Büscher et al., 2017, Kurman et al., 2017), and Madrepora oculata (Maier et al., 2016), when the aragonite saturation state fell below 1, net calcification became close to zero, or fell below zero, as rates of dissolution overtook calcification. Experimental evidence suggests that some Lophelia genotypes may be more resilient than others to ocean acidification (Kurman et al., 2017). Whilst it is thought that most Lophelia pertusa populations occur above the aragonite saturation horizon, Lophelia pertusa is known to persist at aragonite saturation states < 1 along the Californian margin (Gomez et al., 2018), and under experimental conditions fragments managed to maintain slight net positive calcification under low pH and an aragonite saturation state of 0.76 (±0.10) during a 12 month duration (Hennige et al., 2015), suggesting some tolerance to future conditions. This said, this is the first observation of Lophelia pertusa below the ASH (Gomez et al., 2018), and Lophelia was not found at an aragonite saturation state < 1.25 in the Gulf of Mexico (Lunden et al., 2013). Lophelia pertusa has been shown to be able to upregulate the internal pH and therefore saturation state at the site of calcification (McCulloch et al., 2012), although upregulation is energetically costly (McCulloch et al., 2012). The persistence of Lophelia pertusa along the California margin may be due to extremely high productivity of the area (Gomez et al., 2018), whilst maintenance of net calcification under experimental conditions (Hennige et al., 2015) may be due to an abundant food source.

As the aragonite saturation horizon shoals, dissolution of exposed coral skeleton will occur, although live coral tissue will protect against dissolution. This may be compounded by the fact that under high CO2 conditions, Lophelia branches grew longer and thinner, and there is a noticeable change in biomineralisation processes, lead to branches becoming 20-30% weaker than those in control conditions (Hennige et al., 2015). Results from the field suggest that Lophelia pertusa branches found in aragonite undersaturated waters have a lower skeletal density than populations found in saturated waters (Gomez et al., 2018). Dissolution of exposed skeleton, and a weakening of the integrity of live coral branches will lead to a loss of structural complexity becoming more susceptible to mechanical damage and bioerosion (Hennige et al., 2015).

Sensitivity Assessment. Whilst there is evidence that some populations of Lophelia pertusa can survive at an aragonite saturation state < 1 (Gomez et al., 2018), this is the only record of this species being found in undersaturated waters, in an area of extreme productivity. Whilst this offers some hope for future cold-water coral colonies, evidence suggests that aragonite undersaturation is likely to lead to some mortality of this habitat, and even if net calcification can be maintained, it is likely that there will be some loss of colony integrity, making it more susceptible to mechanical damage or bioerosion. Under the middle emission scenario, the aragonite saturation horizon is expected to rise to approximately 800m, so is unlikely to affect the upper bathyal depth zone. Therefore, Lophelia pertusa and Madrepora oculata colonies are expected to maintain growth, without suffering dissolution. Under this scenario resistance has been assessed as ‘High’, whilst resilience has been assessed ‘High’. This leads to an overall sensitivity score of ‘Not sensitive’ at this level benchmark.

Under the high emission scenario, the aragonite saturation horizon is expected to rise to approximately 400m, meaning that 50% of the upper bathyal depth zone (200-600 m) is expected to become undersaturated in aragonite. Whilst some Lophelia pertusa and Madrepora oculata colonies may be able to maintain calcification, there will likely be some mortality and some dissolution of exposed skeletons. For this scenario resistance has been assessed as ‘Medium’. Cold water corals are slow growing, and the undersaturation of waters will continue into the future, reducing growth even further, therefore their resilience has been assessed as ‘Very low’. This gives a sensitivity assessment of ‘Medium’.

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

Due to their calcified nature, both deep and shallow water scleractinian corals are generally thought to be highly sensitive to ocean acidification (Hall-Spencer & Harvey, 2019). The deep-water species, Lophelia pertusa, is a plastic species which has a wide bathymetric range (30-3000 m), lives in several different temperate habitats, and can thrive in the Mingulay Reef complex, where tidal downwellings cause daily variations in pH equivalent to a 25 year jump in atmospheric CO2 (Hennige et al., 2015). This suggests that it may have some tolerance to changes in pH.

Results of the impact of short-term experimental acidification on Lophelia pertusa generally show negative effects such as a decrease in respiration rates (Hennige et al., 2014; 0.3 unit decrease), and calcification rates (Maier et al., 2009; both at 0.15 and 0.3 unit decrease), or no observable effects (Maier et al., 2013b). Georgian et al. (2016), found variability in response between populations. For populations in the Gulf of Mexico, decreasing pH led to a decrease in respiration, prey capture rates and net calcification, whilst populations in Norway responded to a decrease in pH by increasing respiration and prey capture rates, leading to increased calcification (Georgian et al., 2016). Longer term studies (≥ 6 months) show potential for acclimation, with results suggesting that Lophelia pertusa is able to maintain calcification at levels of ocean acidification expected for the end in both the mid- and high-emission scenario  (Form & Riebesell, 2012, Maier et al., 2013a, Movilla et al., 2014, Hennige et al., 2015). Experimental exposures have similarly shown that calcification rates (Maier et al., 2012, Maier et al., 2013b) and respiration rates (Maier et al., 2013a) are maintained at ocean acidification levels expected for the high emission scenario in Madrepora oculata.

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 400m under the high emission scenario (RCP 8.5) and 600m for the middle emission scenario (RCP 4.5) (Zheng & Long, 2014).

For both Lophelia pertusa (Form & Riebesell, 2012, Hennige et al., 2015, Büscher et al., 2017, Kurman et al., 2017), and Madrepora oculata (Maier et al., 2016), when the aragonite saturation state fell below 1, net calcification became close to zero, or fell below zero, as rates of dissolution overtook calcification. Experimental evidence suggests that some Lophelia genotypes may be more resilient than others to ocean acidification (Kurman et al., 2017). Whilst it is thought that most Lophelia pertusa populations occur above the aragonite saturation horizon, Lophelia pertusa is known to persist at aragonite saturation states < 1 along the Californian margin (Gomez et al., 2018), and under experimental conditions fragments managed to maintain slight net positive calcification under low pH and an aragonite saturation state of 0.76 (±0.10) during a 12 month duration (Hennige et al., 2015), suggesting some tolerance to future conditions. This said, this is the first observation of Lophelia pertusa below the ASH (Gomez et al., 2018), and Lophelia was not found at an aragonite saturation state < 1.25 in the Gulf of Mexico (Lunden et al., 2013). Lophelia pertusa has been shown to be able to upregulate the internal pH and therefore saturation state at the site of calcification (McCulloch et al., 2012), although upregulation is energetically costly (McCulloch et al., 2012). The persistence of Lophelia pertusa along the California margin may be due to extremely high productivity of the area (Gomez et al., 2018), whilst maintenance of net calcification under experimental conditions (Hennige et al., 2015) may be due to an abundant food source.

As the aragonite saturation horizon shoals, dissolution of exposed coral skeleton will occur, although live coral tissue will protect against dissolution. This may be compounded by the fact that under high CO2 conditions, Lophelia branches grew longer and thinner, and there is a noticeable change in biomineralisation processes, lead to branches becoming 20-30% weaker than those in control conditions (Hennige et al., 2015). Results from the field suggest that Lophelia pertusa branches found in aragonite undersaturated waters have a lower skeletal density than populations found in saturated waters (Gomez et al., 2018). Dissolution of exposed skeleton, and a weakening of the integrity of live coral branches will lead to a loss of structural complexity becoming more susceptible to mechanical damage and bioerosion (Hennige et al., 2015).

Sensitivity Assessment. Whilst there is evidence that some populations of Lophelia pertusa can survive at an aragonite saturation state < 1 (Gomez et al., 2018), this is the only record of this species being found in undersaturated waters, in an area of extreme productivity. Whilst this offers some hope for future cold-water coral colonies, evidence suggests that aragonite undersaturation is likely to lead to some mortality of this habitat, and even if net calcification can be maintained, it is likely that there will be some loss of colony integrity, making it more susceptible to mechanical damage or bioerosion. Under the middle emission scenario, the aragonite saturation horizon is expected to rise to approximately 800m, so is unlikely to affect the upper bathyal depth zone. Therefore, Lophelia pertusa and Madrepora oculata colonies are expected to maintain growth, without suffering dissolution. Under this scenario resistance has been assessed as ‘High’, whilst resilience has been assessed ‘High’. This leads to an overall sensitivity score of ‘Not sensitive’ at this level benchmark.

Under the high emission scenario, the aragonite saturation horizon is expected to rise to approximately 400m, meaning that 50% of the upper bathyal depth zone (200-600 m) is expected to become undersaturated in aragonite. Whilst some Lophelia pertusa and Madrepora oculata colonies may be able to maintain calcification, there will likely be some mortality and some dissolution of exposed skeletons. For this scenario resistance has been assessed as ‘Medium’. Cold water corals are slow growing, and the undersaturation of waters will continue into the future, reducing growth even further, therefore their resilience has been assessed as ‘Very low’. This gives a sensitivity assessment of ‘Medium’.

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

At a depth of 200 – 600 m, these biotopes will not be affected by sea-level rise.  Therefore, sensitivity to this climate change pressure is probably ‘Not relevant’ in this bathyal habitat.

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

At a depth of 200 – 600 m, these biotopes will not be affected by sea-level rise.  Therefore, sensitivity to this climate change pressure is probably ‘Not relevant’ in this bathyal habitat.

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

At a depth of 200 – 600 m, these biotopes will not be affected by sea-level rise.  Therefore, sensitivity to this climate change pressure is probably ‘Not relevant’ in this bathyal habitat.

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 (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

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 relevant (NR)
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

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Citation

This review can be cited as:

Garrard, S.M., Perry, F. & Tyler-Walters, H., -unspecified-. Discrete [Lophelia pertusa] colonies on Atlantic upper bathyal rock and other hard substrata. 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 11-08-2020]. Available from: https://www.marlin.ac.uk/habitat/detail/1191

Last Updated: 01/01/1970