The scleractinian (stone coral) cold watercoral Lophelia pertusa (L., 1758) is wellknown from the north-east Atlantic. In theFaroe area it was observed for the first timeat the eastern part of the Wyville ThomsonRidge at 969 m depth by the expedition ofHMS ‘Lightning’ in 1868 (Wyville Thomson1873) (Figure 1). L. pertusa builds largebanks, structures that can extend to sever-al hundred meters in diameter and riseseveral tens of meters above the sea floor.In contrast to tropical corals L. pertusa islacking symbiotic algae.

The geographical distribution of Lophelia pertusa within the areaLophelia pertusa is widely distributed andhas been recorded from most seas(Jungersen 1917, Zibrowius 1980, Cairns1994, Copley et al. 1996, Mortensen 1999,Hovland & Mortensen 1999). In the north-east Atlantic it is often to be found on theupper slope of off shore banks and near thecontinental shelf break at depth of general-

ly 200 – 400 m, in temperature of 4 – 8°Cand at a salinity of 33.5-35.2 S (Wilson1979a, Frederiksen et al. 1992).

In 1979 Wilson published a map of the oc-currence of Lophelia pertusa within thenorth-east Atlantic (Wilson 1979a). Hegathered the information from publishedrecords and also included sites known bylocal fishermen. The corals make up a riskof getting the fishing nets caught andripped, and the fishermen have for yearsmarked coral areas on their maps, whenev-er they came across any. Some of thebanks were destroyed in the attempt toturn the areas into good fishing grounds.As the knowledge about the Lophelia banksand their inhabitants has grown, public in-terest has increased. In some countries at-tempts have been made to protect Lopheliabanks, as for example in Norway, wheretrawling was recently regulated and insome areas even stopped.

During the BIOFAR program (1987-1991) aspecial effort was made to further registerthe Lophelia banks in the Faroe area and totest some of the stations within the Faroearea listed by Wilson (1979a) and pointedout by Faroese fishermen (Nørrevang et al.1994, Frederiksen et al. 1992). A map of ar-eas where Lophelia pertusa occurred wascompiled from all existing knowledge.(Figure 2).

In the Faroes Lophelia pertusa is the mostabundant coral. It occurs in single coloniesas well as banks of considerable dimen-sions. Lophelia was always found deeperthan 200 m by Frederiksen et al. (1992),with the largest occurrence on the shelf andupper slope of the banks at 250 – 450 m.The 200 m upper limit is general for the off-shore distribution in the North Atlantic (Zi-browius 1980) and probably relates to max-imum current speeds generated by extremewave conditions met during long periods(Frederiksen et al. 1992). In some Norwe-gian fjords Lophelia has been recorded asshallow as 39 m depth (Rapp & Sneli 1999).

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Lophelia pertusa and other cold water corals in the Faroe area

Figure 1. Lophelia pertusa (from Wyville Thomson 1873).

Growth and morphologyThe Lophelia colony is assumed to developfrom a planula, which settles on the sur-face of rocks, stones, pebbles or old coralbranches. As the colony grows and be-comes larger the network of intertwinedbranches forms attractive microhabitatsand shelter for other invertebrates. Withage the living tissue withdraws from thelowest part of the colony and only the out-er approximately 1 m of the coral banksupports living polyps (Wilson 1979b). Thedegradation of coral banks is carried out bya few very abundant species that actuallybore in the coral. They are mainly boringsponges such as species of Cliona and Alec-tona, but also boring polychaetes are im-portant excavators (Frederiksen et al.1992). When the inner and lower deadparts are attacked, the basal support of thecolony is weakened which can lead to aphysical collapse of the colony. However,even after the collapse, the living outerbranches of the colony will continue togrow (Wilson 1979b).

Branches broken off from the living coral,which fall onto the sediment, may continuegrowth, or become substrate for new plan-ulae, thus providing substrate for a newcolony. Some of the broken branches fre-quently fall into the colony where they arecaught and trapped amongst the intricatebranching and in time become cemented tothe structure (Wilson 1979b). As thecolonies and branchlets in the ring sur-rounding the original colony grow, they inturn generate debris around them that willprovide further colonies. When the sur-rounding ring becomes larger, difficultiesin feeding and long term stagnation of wa-ter movements may cause the death of theinner portion of the large Lophelia colonies(Chamberlain & Graus 1975).

The limitation in size of the colonies is, asalso summarized by Wilson (1979b), deter-mined by such factors as: the mechanicalstrength of the living, growing branches ofcoral (Stetson et al. 1962), the dynamics ofwater flow through the colony in relation tofeeding (Chamberlain & Graus 1975), andthe rate of weakening of the dead portionsof the colonies by activities of boring

sponges and polychaetes etc. Consequent-ly, the banks reach an equilibrium of for-mation and degradation.

FeedingThe knowledge about Lophelia’s feedingstrategy is limited, but from aquarium ex-periments lasting for three years in Trond-hjem Biological Station, Norwegian Univer-sity of Science and Technology, it has beenfound highly variable (Rapp & Sneli 1999).They suggested zooplankton and “marinesnow” to be the main food sources. They al-so found that the well known mucus pro-duction, formerly regarded as a mecha-nism to cope with stress, sedimentationand “fouling” organisms, was important inenabling large prey and large numbers ofprey to be captured and ingested. In situstudies on a bank off Norway showed thefood to be planktonic crustaceans (Hein-rich et al. 1997)

Unpublished results from aquarium exper-iments show that when fed different typesof prey Crustacea, with a size up to at least35 mm together with Chaetognatha wasthe preferred food source. Ctenophora,medusa and pure gelatine pellets were re-jected, indicating that Lophelia use bothtactile and chemosensitive receptors in dis-crimination and selection of prey. High

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Figure 2. Distribution of Lophelia pertusa in Faroe waters. Aand B are localities where coral associated fauna wasinvestigated (see Jensen & Frederiksen 1992).

concentrations of prey induced high cap-ture rates (Rapp et al., submitted).

Growth rates and age of coral banksGrowth is slow. Possible growth rates of 4.1– 7.5 mm/year were reported in a morethan 100 year old investigation (Duncan1877). These rates were direct measure-ments of Lophelia colonies settled on cableslaid out and recovered after 6 years. Recentstudies suggest, based on oxygen and car-bon ratio measurements related to growthline patterns in the skeleton, an averagegrowth of 4.3 mm/year in shelf corals and7.0 mm/year in corals from the Trondheimfjord (Mortensen & Rapp 1998). Thesegrowth-rates give an estimated age of mini-mum 300 years for a colony of 1.5 mheight. 15 m large banks might then be atleast 2000 years old, but accounting forfragmentation and decomposition, thebanks are undoubtedly much older. Agedetermination by isotope techniques allowsby comparison with similar Norwegian in-vestigations (Hovland et al. 1998) to esti-mate the age of Lophelia in the Faroe areato be approximately 9000-10.000 years(Unpublished information, Israelson andTendal). Mikkelsen et al. (1982) found thatdata derived from stable isotope ratioanalysis of corals from the Trondheim fjordindicated a growth rate of 2.5 cm/year,

however, Mortensen & Rapp (1998) found itto be surprisingly high and suggested thereason to be bulk-sampling of more thanone growth layer.

Lophelia banks as a habitatIn the mature coppice, only the outer 1 – 2m is alive. Beneath this, dead coral andfragments are found in a compact core. Thebanks function as sediment traps by re-ducing the bottom current speed, and oftenfine mud, sand and particles are trappedwithin the branches of the banks (Stetsonet al. 1962).

Compared to the surrounding seabed, thenetwork of intertwined branches forms avariety of habitats and shelter for many dif-ferent organisms, like for example the freespace between the coral branches, thesmooth surface of the living Lophelia, thedetritus laden surface of dead Lophelia andthe cavities inside dead Lophelia made byboring sponges (Mortensen et al. 1995)(Figure 3). The dead coral debris andtrapped sediment supports a rich fauna ofsponges, anemones, bryozoans, calcareoustube building polychaetes, brachiopods, bi-valves, asteroids and echinoids, and alsoprovide shelter for several scavengers (Wil-son 1979b).

During the BIOFAR project, a thorough in-vestigation of the fauna associated withLophelia pertusa banks was carriedthrough at two sites indicated in Figure 2.This is the only investigation on the faunaassociated with Lophelia within the Faroearea; however, investigations of Norwegiancoral banks (Burdon-Jones & Tambs-Ly-che 1960) and blocks from the Bay of Bis-cay (LeDanois 1948) suggested a rich asso-ciated fauna. In the Faroe area 11 blocksfrom site A and 14 blocks from site B of 0.2– 2.0 kg were examined. An echogram ofsite A showed the bank to be 10 m high and110 m wide. The area is exposed to tidalcurrents with an average speed of 50 cm/s.Site B has an average speed of 35 cm/s.(Jensen & Frederiksen 1992). At both sitesthe average temperature is 6 – 8°C with astandard deviation of 0.5 – 1.0°C (Wester-berg 1990). Clay and silt were found be-tween the coral branches at both sites.

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Figure 3. Blocks of live and dead Lophelia pertusa. The intricatebranching provides space and shelter for a variety ofspecies. (© section of photo by O.S. Tendal)

As expected a highly diverse and rich faunawas shown to be associated with the Lophe-lia banks. In the BIOFAR study 300 specieswere identified, of which 256 species werefound on the blocks examined and 42species were identified from loose coralrubble. A list of species occurring at mini-mum half of the 25 blocks is provided inTable 1.

The most species rich groups were Poly-chaeta (67 species), Bryozoa (45 species)and Porifera (29 morphological types) andin number of individuals, not counting thecolonial species, the dominant groups werePolychaeta, Bivalvia, Echinodermata andBrachiopoda (Jensen & Frederiksen 1992).

Jensen & Fredriksen (1992) found thatmost individuals occurred in the dead coralblocks from the inner part of the bank orcolony, and only 20 species, represented bymore than 1 individual, were found exclu-sively on the living part of the corals. Thecoral polyps are not interconnected, as ep-ithelium does not cover the entire surfaceof the skeleton and fauna associated withthe live coral blocks were found betweencalices rather than on or in the living tissueof Lophelia. Polychaeta and Gastropoda oc-curred in twice as many individuals ondead as on live coral. Crustacea, Sipuncu-la, Bivalvia, and Nematoda were found in 4- 8 times as many individuals on dead ason live coral. Ascidia, Anthozoa and Echin-odermata were found in more than 10times as many individuals on dead as onlive coral and Brachiopoda were found tobe more than 50 times as frequent on deadthan on live coral.

Of the 20 most abundant species only fourshowed a correlation between the numberof individuals and coral weight. For livecoral blocks these were the polychaete Eu-nice norvegica, the bivalve Modiolula phase-olina and nemertea sp. A., and on deadcoral only the polychaete family Para-onidae.

Many juveniles were found in the corals byJensen & Frederiksen (1992) and thecorals may provide a good nursery areawith protection from predators like it is

known from the holdfasts of the largebrown algae Laminaria in shallower water(Christie et al. 1994, Worsaae 1998).

The associated fauna is facultative as noneof the species occurring in the sampleswere exclusively found in Lophelia pertusabanks and obligatory associated to it. Manyof the species are common in the Faroesearea, and the corals may thus act as an im-portant habitat for recruitment of speciesto the more barren surrounding deep waterareas. This is contrary to Dons (1944) whofor a Norwegian bank mentioned about 50species to be obligate Lophelia associatedfauna and nearly always found on thestone-coral banks he investigated. How-

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Figure 4. Trawl catch from 375 m depth west of the Faroe Is-lands, BIOFAR Stn. 535 (Nørrevang et al. 1994). Thecharacteristic animals are, in addition to Lophelia,large branches of the octocoral Paragorgia, the bivalveAcesta, the brittle star Gorgonocephalus, and largesponges of the genus Geodia (© photo by A. Klitgaard).

ever, Burdon-Jones and Tambs-Lyche(1960) only found a limited overlap ofspecies when comparing Dons species listwith what they found on a coral bank in aNorwegian fjord near Bergen, and suggest-ed as Jensen & Frederiksen (1992) for theFaroes that no such obligate fauna exists.

The coral banks in the north-east Atlanticare built by Lophelia pertusa alone, where-as other cold water corals like Madreporaoculata (L., 1758) and Solenosmilia varia-bilis Duncan, 1873 are found scattered andsometimes use Lophelia as substrate Theassociated fauna consists mainly of sus-pension and particle feeders like the coralitself (Figure 4). Lophelia banks are foundin areas with considerable water movementand abundant suspended material (Jensen& Frederiksen 1992).

The study by Jensen & Fredriksen (1992)only included the closely associated fauna.The main part of the mobile species are notobtained with the type of gear used (trian-gle dredge) and free-living species could belost during haul up. Video investigationswith ROV (Remote Operated Vehicle) onNorwegian banks showed the presence ofseveral fish species at the banks. Especial-ly redfish and saithe dominated the fishfauna (Mortensen et al. 1995).

Large numbers of Eunice norvegicus (L.,1767) were found associated with thebanks. This large polychaete can grow upto 20 cm in length. It lives in parchment-like tubes in the living part of the coralwhere the coral strengthens the tubes byovergrowing and encapsulating them. An-other very common species was the brit-tlestar Ophiactis balli (Thompson, 1840)which lives in the dead skeletons of thepolyps or other cavities, with the body hid-den and protected and arms extended tofeed. Such cavities are also often occupiedby the bivalves Hiatella arctica (L., 1767) orAcar nodulosa (Müller, 1776) (Jensen &Frederiksen 1990).

The importance of food supply on the distribution of Lophelia pertusa banksLophelia pertusa is a suspension-feederand as such is dependent on food being

brought to it. Studies on the distribution ofcorals in Norwegian fjords suggest areas ofstrong current as a preferred habitat (Dons1944). This was not supported by Frederik-sen et al. (1992) for the distribution ofLophelia pertusa in the Faroes. They foundthe densest concentrations to occur closeto areas where the conditions allow inter-nal tidal waves to break at the sloping bot-tom. They argued that the food supply topassive suspension feeders like corals notnecessarily increases with increasing cur-rent speed but that the crucial factor is theparticle flux to the bottom mixed layer. Twodifferent types of possible increases in sed-imentation rates were discussed (Frederik-sen et al. 1992).

One is by internal waves generated by theadvection of stratified water across bottomcontours by the barometric tide. This gen-erates rays of energy, that enhance the ver-tical mixing and results in nutrient richbands parallel to the shelf break contour,which in turn enhances phytoplanktonproduction within the bands. The in-creased production would at least season-ally lead to a higher vertical detritus fluxand thereby higher food availability for thesuspension feeders at the bottom. However,the horizontal advection is much largerthan the downward flux of small particles,and these would be transported away fromthe production sites before reaching thebottom. Only faecal pellets and marinesnow have a sinking rate large enough toenable the surface production to reach thebottom mixed layer close to the productionsite. As the water depth increases the verti-cal detritus flux to the bottom becomesweaker and the spatial separation betweenproduction site and the area where the sur-plus is deposited increases. Another expla-nation could be that when the bottom slopeequals the rays, called the critical slope,the local increase in mixing intensity willlead to a thickening of the bottom mixedlayer and to a resuspension of organic par-ticles that may have been deposited, there-by increasing the availability of food (Fred-eriksen et al. 1992, Klitgaard & Tendal,this volume).

This could explain the known distribution

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Table 1. Invertebrates associated to Lophelia pertusa occurring at a minimum half of the 25blocks are shown. Number shows frequency of occurrence.

NEMATODADeotostoma sp. 14

POLYCHAETAEusyllis blomstrandi Malmgren 1867 13Sphaerosyllis spp. 17Typosyllis armillaris (O.F. Müller, 1776) 19Paraonidae indt. 14Lanassa cf. venusta (Malm, 1874) 14Polycirrus cf. norvegicus Wollebæk, 1912 16Perkinsiana socialis (Langerhans, 1884) 14Placostegus tridentata (Fabricius, 1779) 19Protula tubularia (Montagu, 1903) 14

MOLLUSCA, Bivalvia* Modiolula phaseolina (Philippi, 1844) 19

Heteranomia squamula (smooth) (L., 1758) 17H. squamula (scaly) (L., 1758) 16Hiatella arctica (L., 1767) 20

CRUSTACEAGnathia sp. juvenile 19G. abyssorum G.O. Sars 1872 13G. dentata G.O. Sars, 1871 14

SIPUNCULAGolfingia minuta (Keeferstein, 1862) 19

BRYOZOACrisia eburnea (L., 1758) 18C. aculeata Hassall, 1841 13Idmidronea atlantica (Forbes in Johnston, 1847) 18Cabarea ellisii (Fleming, 1814) 13Porelloides laevis (Fleming, 1828) 13Sertella beaniana (King, 1846) 14

BRACHIOPODACrania anomala (O.F. Müller, 1776) 14Terebratulina retusa (L., 1758) 15Macandrevia cranium (O.F. Müller, 1776) 15

ECHINODERMATAOphiactis abyssicola M. Sars, 1861 17O. balli (Thompson, 1840) 14

* By wrong identification the species is given as M. modiolus in Jensen & Frederiksen (1992).

of Lophelia pertusa in the area of the uppershelf and in the vicinity of critical slope ar-eas in the Faroes. Such conditions wouldnot only be a benefit for growth of Lopheliabut also for other suspension feeders(Frederiksen et al. 1992), such as the largesponges as is indicated by their distribu-tion (Klitgaard & Tendal, this volume).

Others have suggested that the Lopheliabanks off Mid Norway are linked to areas ofhydrocarbon seeps (Hovland et al. 1998,Hovland & Thomsen 1997). The reason forthis kind of distribution is again suggestedto be an enhancement in food supply in re-lation to surrounding more barren areas.At hydrocarbon seeps the water becomesenriched with inorganic and organic car-bon components that might be utilized bymicro-organisms which again can be usedas food by filter feeders and so forth. If thechemical and micro-organic condition isstable over prolonged periods, and the sub-strate and current regime is favorable, par-ticle feeders including Lophelia find opti-mum conditions for settling and growth(Hovland et al. 1998).

Variation in morphologyVariation in morphology of Lophelia mightbe correlated with differences in the energylevels at the place of growth. From bottomcontours, it is assumed that slender, tallforms are from low energy environmentscompared to the compact forms that mightbe found in high-energy environments. Thedifference in tall and compact forms is alsoreflected in the internal structure of thecorals as the compact forms have a denserskeleton (Mikkelsen et al. 1982). Compactand extended growth forms may occur indifferent parts of the same colony indicat-ing differences in the rate of budding, prob-ably correlated with the availability of food.The spacing and orientation of the polypscertainly suggest that the growth attemptto maximize the exploitation of the foodsupply (Freiwald et al. 1997).

Reproduction and recolonizationWilson (1979b) did not find evidence ofsuccessful settlement of planulae from thesubmarine dive on the Rockall Bank andhe concluded that almost all growth in L.

pertusa was by asexual budding. Accord-ingly death of L. pertusa will leave the givenarea barren for a long time. Even when re-colonized, it will take many years for thenewly settled Lophelia planulas to grow in-to banks of considerable size, due to theslow growth rate, and for the rich associat-ed fauna to develop. Of course recoloniza-tion will only take place if the disturbancethat killed the former Lophelia banks hasceased.

Lophelia pertusa and most sessile suspen-sion feeders are effected and physically dis-turbed when the water becomes loadedwith suspended inorganic particles as aneffect of nearby trawling or by the dis-charge of cuttings from the oil industry. Wedo not know the long-term effect of drillingactivities on neighbouring Lophelia pertusabanks or other hard bottom communities.However, the effect from a rise in sedimentinflux could be a partial or total burial ofcoral colonies. The sediment covers thesurface and all the hollows of the colony,and in time the corals and their associatedfauna may be smothered and totally buriedif the sediment load continues.

Concluding remarksLophelia banks are widely distributed with-in the Faroese EEZ, and are particularlyfound close to the shelf break or on placeswith similar high-energy water movementconditions.

Lophelia banks get very old and the esti-mated age of Lophelia in the Faroe area isapproximately 10,000 years. Growth isslow, and if destroyed either by physicaldamage or by smothering, the banks areonly very slowly rebuilt, if at all.

Although no obligate associated Lopheliafauna exists, the banks are very rich in as-sociated species and in number of individ-uals, and may function as nursery and re-cruitment area for the more barren sur-roundings.

Primnoa resedaeformisThe octocoral Primnoa resedaeformis (Gun-nerus 1763), ‘sea corn’ or ‘rice coral’ has

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been found many times at the Faroes (Fig-ure 5). It was first recorded by Thomson(1906) in the Faroe-Shetland Channel, lat-er by Madsen (1944) from two localitieswest of the Faroes, by Grasshoff and Zi-browius (1983) from Bill Bailey Bank, andfinally during the BIOFAR project at 18 lo-calities all around the Faroes (Nørrevang etal. 1994, Tendal in prep.).

It has been taken at 90-1020 m depth, withmost records between 200 and 500 m. Thetwo localities shallower than 200 m are onthe western and eastern side of the Faroeseplateau, while the only two deeper than500 m, both at around 1000 m, are on thenorthern and southern flanks of the BillBailey Bank, respectively. All records, ex-cept the northernmost of the deepest andone in the Faroe-Shetland Channel arefrom areas dominated by North Atlanticwater, with temperatures of 6-8°C. The re-maining two are from water of 2-4°C (West-erberg 1990).

Most samples are fragments, but a numberof whole specimens, about 1 m high weretaken by the trawls. According to literaturethis is the normal maximum size for thespecies all over its North Atlantic distribu-tion area (Broch 1912, Carlgren 1945,Breeze et al. 1997). Specimens of that sizeare supposed to be of considerable age, atleast 500 years old (Strömgren 1970, Risket al. 1998). P. resedaeformis has been re-ported to be viviparous but nothing isknown of the reproduction frequency(Thomson 1906; the remark by Breeze et al.1997 that the development is pelagic seemsunconfirmed).

The species is to be considered very vulne-rable to physical damage.

Paragorgia arboreaThe octocoral Paragorgia arborea (Linné1758), ‘sea tree’ or ‘bubble gum coral’ isknown from several locations in the Faroes(Figure 6). It was first mentioned by Mad-sen (1944; the same specimen mentionedand shown on a photograph in Vedel Thån-ing 1958), and during the BIOFAR project 6other records turned up (Tendal 1992, Nør-revang et al. 1994, Tendal in prep.).

It has been taken at 260-649 m depth, with6 of the records shallower than 500 m. Allrecords are from the southern part of thearea, from North Atlantic water with a tem-perature of 6-9°C, and with calculated cur-rent velocities of 45-60 cm/sec (Westerberg1990).

The largest fragment found during the BIO-FAR project was a branch 12 cm thick,about 150 cm long and with a branchingpart 50 cm wide. Fishermen have toldabout colonies of about 2.5 m height. The

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Figure 5. Distribution of Primnoa resedaeformis in the Faroearea.

Figure 6. Distribution of Paragorgia arborea in the Faroe area.

species grows very slowly, and specimensof that size are at least 1500 years old (Ten-dal & Israelson, unpublished). The repro-duction pattern is unknown (the remark byBreeze et al. 1997 that the development ispelagic seems unconfirmed).

Tendal (1992) believed the species to be

more widespread, as the gear used duringthe BIOFAR project in many cases was toosmall. Of the 7 samples, 5 were obtained bytrawl.

The species must be considered very vul-nerable to physical damage.

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