A survey of the genera Sarcodictyon Forbes, 1847, and rolandia Lacaze-Duthiers, 1900, (anthozoa: octocorallia) in the North-eastern Atlantic and the Mediterranean

A su ey o he gene a Sa codic yon Fo bes, 1847, and Rolandia Lacaze-Du hie s, 1900, (An hozoa: Oc oco allia) in he No h-eas e n A lan ic and he Medi e anean O. Ocana, P .J. Lopez-Gonzalez, J. Nunez & J.C. Ga cia-Gomez Ocana, 0., P.J. LOpez-Gonzalez, J. NU iez & J.C. Ga cia-Gomez. A su ey o he gene a Sa codic yon Fo bes, 1847, and Rolandia Lacaze-Du hie s, 1900, (An hozoa: Oc oco allia) in he No h-eas e n A lan ic and he Medi e anean. Zool. Med. Leiden 73 (28), 6.iii.2000: 413-426, igs 1-5.- ISSN 0024-0672. O.Ocana, Ins i u o de Es udios Ceu es (CECEL-CSIC), Paseo del Re ellin, 30, Apdo. 51001 Ceu a (No h-A ica), Spain. J. NU lez, Depa amen o de Biologia Animal, Facul ad de Biologia, Uni e sidad de La Laguna, Tene - i e (Cana y Islands), Spain. J.C. Ga cia-Gomez & P.J. LOpez-Gonzalez, Labo a o io de Biologia Ma ina, Depa amen o de Fisi- alogia y Biologia Animal, Facul ad de Biologia, Uni e sidad de Se illa, Apdo. 1095, 41080 Se illa, Spain. Key wo ds: S oloni e a; Sa codiclyon; Rolandia; No h-eas e n A lan ic; Medi e anean. Sa codiclyon ca ena um Fo bes, 1847, is a common species in he A lan ic and he Medi e anean. How- e e , o en i has been mis aken o Rolandia co alloides Lacaze-Du hie s, 1900. On he basis o B i ish and Medi e anean ma e ial we s udied he main cha ac e s o dis inguishing S. ca ena um. We con- side Rolandia co alloides Lacaze-Du hie s, 1900, o be a alid species and we assign o i all he ma e i- al named Rolandia osea by S. Weinbe g (1978). Fu he mo e, a his ological s udy con i med ha R. co a/loides ypically o ms g oups o polyps embedded in a common coenenchyme. The e o e, he pos- sibili y o ans e ing R. co a/loides om S oloni e a o Alcyonacea is discussed. E ago gia osea Philip- pi, 1842, is he e conside ed o be a nomen dubium. In oduc ion The genus Sa codic yon Fo bes (in Johns on), 1847, is based on B i ish ma e ial o Sa codic yon ca ena um, desc ibed in he same pape . Fo bes & Goodsi (1851) desc ibed a second species, Sa codic yon agglome a a, e y simila o he ype species. They dis- inguished he species by ex e nal cha ac e s only"... a och aceous yellow colou and in he g ouping o polyps in h ees and i es ...". Subsequen ly, Sa s (1856) desc ibed Rhizoxenia ili o mis, which p obably ep esen s ano he species o Sa codic yon (see discussion o S. ca ena um). Gosse (1858) collec ed and s udied se e al colonies om he sou h coas o England and iden i ied hem as S. ca ena um. These B i ish pape s culmina ed in he syn he ic wo k o He dman (1885), ou s anding o i s me hodolo- gy used o he s udy o his genus. Ten yea s la e , He dman (1895) synonymized s. agglome a a wi h S. ca ena um. Meanwhile Philippi (1842) desc ibed E ago a osea om he Medi e anean. This species p obably belongs o Sa codic yon o Rolandia, bu his emains unce ain, due o he inadequa e desc ip ion gi en by his au ho ; i is e en possible ha i belongs o Scle an helia S ude , 1878 (see Ca pine, 1964). A e Philippi, species wi h some esemblance o E. osea we e gene ally e e ed o his species. Howe e , he si ua ion became mo e complica ed when Lacaze-Du hie s (1900) desc ibed ano he simila species om he Medi e anean, Rolandia co alloides. Thomson (1927) eco ded s. ca e- na um om Monaco, wi hou desc ibing he scle i es. La e on, Be enguie (1954) 414 Ocana e a1. A su ey o he gene a Sa codic yon and Rolandia. Zo01. Med. Leiden 73 (2000) ci ed S. ca ena um o he Medi e anean and claimed ha S. ca ena um and R. co al- loides we e he same species. Pax & MUlle (1962) dis inguished wo Ad ia ic species Zoan ha osea, Philippi, 1842, (possibly S. ca ena um) and Zoan ha co ii B och, 1935, (= R. co alloides), using scle i e cha ac e s. Weinbe g (1978a) conside ed E. osea and R. co alloides o be he same species, Rolandia osea. In ha pape Weinbe g assumed ha he scle i es in he genus Sa codic yon a e insepa ably used. This e o was c ea ed by Hickson's (1921: 367) misin e p e a ion o He dman's desc ip ion, epea ed by Oeich- mann (1936), Goha (1940) and Baye (1956). Baye (1981) pu an end o his con usion and illus a ed some scle i es o a specimen o S. ca ena um om B i ish wa e s (Isle o Man). In he same pape he e e ed Weinbe g's Rolandia osea o Sa codic yon. Finally, Manuel (1981) synonymized S. ca ena um and R. osea as Sa codic yon oseum. We s udied ma e ial om he A lan ic as well as he Medi e anean and ecognize wo species: Sa codic yon ca ena um Fo bes, 1847, and Rolandia co alloides Lacaze- Du hie s,1900. They a e desc ibed and depic ed below. Based on he scle i es and he ana omy we sugges o main ain R. co alloides in a genus di e en om Sa codic yon. Fo a compa ison o axonomic cha ac e s o Sa co- dic yon and Rolandia we e e o able 1. Ma e ial and Me hods The colonies s udied in he p esen wo k we e collec ed by SCUBA di ing du ing di e en sampling ips o ganized by he Na ional Museum o Na u al Sciences o Mad id along he sou he n Ibe ian Peninsula: "Fauna-I" in he Albo an Sea, "Fauna- II" in he Can ab ic Sea, "Fauna-ill" in he Balea ic wa e s, and "Fauna-IV" in Balea ic and Columb e es wa e s. Ma e ial o Rolandia co alloides om he Zoological Museum o Ams e dam (ZMA), p e iously s udied by Weinbe g, was also a ou disposal. Colonies we e anaes he ized wi h men hol c ys als and p ese ed in 70% alcohol; specimens used o his ological s udy we e ixed in 8% o maldehyde. The gene al mo phology and ana omy we e s udied by means o a s e eo dissec ing mic oscope. Ana omical and his ological de ails we e s udied ollowing he Cajal me hod o opog aphic s aining (see Gabe, 1968). Scle i es we e ob ained by dissol ing he so issues in concen a ed sodium hypochlo i e. They we e examined wi h a ligh mic o- scope and SEM. Fo e minology we used Baye e al. (1983). The colonies a e deposi - ed in he Na ionaal Na uu his o isch Museum, Leiden, The Ne he lands (RMNH), Museo Nacional de Ciencias Na u ales de Mad id (MNCN), and Labo a o io de Biologia Ma ina de la Uni e sidad de Se illa, Spain (LBM). Key o No h-eas e n A lan ic and Medi e anean species o Sa codic yon and Rolandia 1. Polyps embedded in a common coenenchyme; some imes s olons a e p esen . Scle i es mainly " used quad uple s" bu c osses and shu les a e also common Rolandia co alloides - Polyps g owing om na ow ibbon-like s olons, which a e some imes used o o m memb anous expansions. Scle i es wi hou " used quad uple s" 2 2. Scle i es include s ella e pla es Sa codic yon ca ena um Ocana e a1. A su ey o he gene a Sa codic yon and Ro/andia. Z001. Med. Leiden 73 (2000) 415 No s ella e pla es p esen bu six- adia es, b anched spindles, c osses & ods Sa codic yon cana iensis Family Cla ula iidae Hickson, 1894 Genus Sa codic yon Fo bes (in Johns on), 1847 Diagnosis.- Polyps g owing om na ow ibbon-like s olons, which a e some- imes used o o m memb anous expansions. Mesogloea o polyp wall wi h many lacunae o e icula . Re ac o muscle es ic ed o di use. S olons wi h 2 o 5 canals, some imes mo e. Scle i e ypes: six- adia es, s ella e pla es, c osses, b anched spindles and shu les. An hocodiae wi h minu e scle i es o none a all. Rema ks.- Baye (1981: 899) es ablished he sub amily Sa codic yinae, including di e se axa such as Cya hopodium Ve ill, 1868, and Tesse an helia, Baye , 1981, wi h used pla es and ope culum, and Sa codic yon and Scle an helia, wi h a comple ely di - e en mo phology. The cha ac e s used o dis inguish he sub amily a e based on ex e nal mo phology "Polyps sho , e ac ing almos lush in o s olons, o p oducing low, conical o cylind ical calices". Some species wi h la ge polyps, iz., Sa codic yon cana iensis (Ocana e al., 1992) and Scle an helia homsoni (Williams,i987) canno be i - ed in Sa codic yinae. The e o e, we decided no o conside his sub amily in ou p esen pape . The genus Scle an helia S ude , 1878, is e y simila o Sa codic yon. I only di e s by he p esence o s ella e pla es, disposed as a iled loo on s olon and calyx. Sa codic yon ca ena um Fo bes, 1847 ( igs 1-2) Sa codic yon ca ena a Fo bes (in Johns on), 1847: 179-180, pl. 33, igs 4-7, desc ip ion ex e nal mo phol- ogy and habi us, deep wa e , England (Loch Fine, Youghal, wes coas o Sco land); Gosse, 1858: 276-280, pl. 9, desc ip ion ex e nal mo phology, some da a o scle i es and many aqua ian obse - a ions, England (Ilsam), a in e idal ca e ns; He dman, 1885: 31-49, pis 1-3, desc ip ion ex e nal mo phology, ana omy, his ology and scle i es, England (Ta be , Loch Fyne, Sound o Sky, A os bay, Sound o Mull, C ouling Island, Heb ides), ci cali o al and occasionally in shallow wa e s; He dman, 1895: 163-168, pl. 8, desc ip on o he yellow a ie y, me ging Sa codic yon agglome a a wi h Sa codic yon ca ena um. Sa codic yon ca ena u n; Thomson, 1927: 8-9, pl. 2 ig. 17, p o pa e, Dogge Bank No h A lan ic; Mad- sen, 1944: 11, only eco ded, 55 m deep; Be gen, No way; Tixie -Du i aui & d'Hond , 1974: 1366, eco d and dis ibu ion, Azo es (Flo es). Cla ula ia ca ena a; Hickson, 1894: 332, pu in a di e en genus as a esul o a misin e p e a ion o He dman's pape . Sy npodiu n ca ena u n; Molande , 1915: 43, diagnosis and ema ks; No way, England, Sco land and I eland. Sa codic yon oseu n; Manuel, 1981: 36-38, ig. 8, desc ip ion ex e nal mo phology, scle i es, ci cali - o al, B i ish coas s, Sou h-wes Eu ope and he Medi e anean, pe haps no hwa d o Scandi- na ia? (misiden i ica ion). ?Rhizoxenia ili o mis Sa s, 1856: 65-67, Pl. X, igs. 13-17, desc ip ion ex e nal mo phology, some habi a da a, ci cali o al, colou o he colony g eyish-whi e, possibly a di e en species, could be a di - e en genus. ? E ago a osea Philippi, 1842: 36, ig. 2, b ie desc ip ion, Naples. No Rolandia osea; Weinbe g, 1978: 167-173 (= Rolandia co alloides). Ocana e al. A su ey o he gene a Sa codic yon and Rolandia. Zool. Med. Leiden 73 (2000) 416 Ma e ial.- Spain: (MNCN 2.04/722), Albo an Island, 15. ii.89, 35°55.95'N 3°01.56'W, wo colonies, 34-44 m, ci cali o al, Fauna-I c uise (s a. 33); (LBM/PI 99) C ina is, 36°10.10'N 5°23.70' W, ii.1990, h ee colonies, in ali o ai, 20-25 m, P.J. Lopez leg.; (RMNH Coel. 23984), Cadiz, La A una a, 36°10.54'N 5°20.00'W, iii.1988, one colony, ci cali o al 40 m, P.J. Lopez leg.; (MNCN 2.04/723) Can ab ic Sea, No he n Ibe ian Peninsula, 43°28.62'N 2°53.77'W, 21. i.91, one colony, 88 m, ci caIi - o ai, Fauna-II c uise (sm. 145-A); (MNCN 2.04/724) Ho migas Islands, 37°39.52'N 0038.92'W, 21. ii.96, one colony, 25 m, in ali o ai, Fauna-IV c uise (s a. 299B5); G ea B i ain: (MNCN 2.04/712), Po land Ha bou , Do se , ii.1977, one colony on limes one, R.L. Manuel leg.; (RMNH Coel. 24087), Millpo , Island o Cumb ae, Sco land, xi.1979, one colony on shell, R.L. Manuel leg.; (RMNH Coel. 24088) Isle o Man (1 mile no h o he Cal o Man), ii.1980, one colony, 30 m, a ached on shell, R.L. Manuel leg. Diagnosis ( o a de ailed desc ip ion, see He dman, 1885; 1895).- Colonies wi h 2 o mo e han 20 polyps, joined by ibbon-like s olons, some imes used o o m mem- b anous expansions. Calyx leng h 3 o 6 mm. An hocodiae whi e, an hos eles and s olons eddish o pale-yellow. Gene ally he an hocodiae ha e small scle i es: c oss- es, shu les and six- adia es, 24-52 pm long and 15-32 pm wide. S olons wi h six- adi- a es, shu les and c osses, 70-116 pm long and 30-110 pm wide. The s ella e pla es a e mos abundan in he an hos eles; hey a e 56-114 pm long and 32-120 pm wide. A dis inc pe ide m is p esen on he an hos eles and s olons. Mesogloea o he polyp walls wi h many lacunae, o ally o pa ially co e ed wi h g anula endode - mal ma e ial. Endode m o he polyps poo ly de eloped. Ec ode m o he en acles well de eloped. The scle i es a e loca ed be ween he ec ode m and he mesogloea. Mesen e ies wi h a s ong, es ic ed e ac o muscle in he middle. Pha ynx wi h a educed mesogloea wi hou lacunae; endode m well de eloped, cilia ed and glandu- la . S olons wi h 2-4 in e nal canals. Ecology.- The colonies s udied we e collec ed om in e idal o ci cali o al bo - oms be ween 0 and 100 me es, on s ones and shells, occasionally close o Alcyonium palma um Pallas, 1766, o associa ed wi h ascidians. Acco ding o Manuel (1981) com- pe i ion o space may de e mine he g ow h o m o his species. Dis ibu ion.- A lan ic and No h Sea: England, Sco land, Wales, Dogge bank; Medi e anean: Algeci as Bay, Albo an Island and Ho migas Islands (Alican e). Discussion.- Sa codic yon ca ena um is a well known species since He dman's desc ip ion (1885, 1895). The main cha ac e s o dis inguish his species a e he p es- ence o minu e scle i es in he an hocodiae (especially in he abo al ace o he en a- cles) and he es ic ed e ac o o he mesen e ies (He dman, 1885: pl. 2 ig 10; pl. 3 igs I, 3). Howe e , in he en acles o some colonies om he wes e n Medi e anean scle i es a e sca ce o absen al oge he . Mo eo e , al hough hese colonies also ha e mesen e ies wi h s ong e ac o s hese a e less es ic ed. As hese di e ences a e g adual, we conside hem in aspeci ic a ia ion. Howe e , u u e s udies migh e eal he Medi e anean popula ion o ep esen a di e en subspecies. Sa codic yon ca ena um di e s om S. cana iensis in ha ing s ella e pla es in he coenenchyme and small scle i es in he polyps. Also he mesogloea o he polyp wall and he e ac o muscles a e clea ly di e en (see Ocana e al., 1992). Rhizoxenia ili o mis (Sa s, 1856) om he No h Sea has some esemblance wi h Sa codic yon. Sa s dis inguished his species om S. ca ena um by di e ences in colou and shape: "No e espece No egienne se dis ingue de l'espece B i annique pa la Ocana e al. A su ey o he gene a Sa codic yon and Rolandia. Zool. Med. Leiden 73 (2000) 417 Fig. 1. Scle i es o Sa codic yon ca ena um.I-12 an hos ele; 13-25 s olon; 26-41 en acles. Scale ba : 150 pm. Oca a e al. A su ey o he gene a Sa codic yon and Rolandia. Zool. Med. Leiden 73 (2000) 418 Fig. 2. His ology o Sa codic yon ca ena um. A, polyp wall. B, c oss-sec ion o a en acle. C, E & F, c oss- sec ions o mesen e ies, e ac o muscles and ilamen . D, de ail o he pha ynx and siphonoglyph. G, c oss-sec ion o s olon, H, de ailso he canals in he s olon. Scale ba s: A = 75 pm; B = 13 pm ; C = 42 pm; D = 30 pm; E = 28 pm; F = 28 pm; G = 67 pm; H = 39 pm. couleu ..., pa les s olons, qui son beaucoup plus e oi s, n'ayan pas la moi ie de la la geu des cellules polypi e es; e pa la plus g ande longeu des s olons en e les cellules, ...". This axon could be ano he Sa codic yon species, bu could jus as well belong o Scle an helia o e en Cla ula ia, as some species o he la e genus can ha e a s iking ex e nal esemblance o Sa codic yon. This is, o ins ance, he case wi h Sa - codic yon densum and Sa codic yon cha co i, desc ibed om he Azo es by Tixie -Du i- aul & d'Hond (1974: 1367-1372; i~s 3-8). These las -named species should be 419 Ocana e al. A su ey o he gene a Sa codiclyon and Rolandia. Zool. Med. Leiden 73 (2000) placed in Cla ula ia (see Ocana e al., 1992: 423-424). G ieg (1887) desc ibed Rhizoxenia alba om he No h Sea, which also clea ly belongs o Cla ula ia (see G ieg, 1887: 15, pl. 1; Molande , 1918: 15-19, igs 5-6). As he ype ma e ial o E ago a osea Philippi, 1842, is almos ce ainly los and because o he inadequa e desc ip ion we conside E. osea as a nomen dubium. Genus Rolandia Lacaze-Du hie s, 1900 Diagnosis.- Polyps embedded in a common coenenchyme. Some imes ibbon- like s olons a e p esen , which can use in o memb anous expansions. The mesogloea o he polyp wall is hin and wi hou lacunae, whe eas i is hick and wi h many lacu- nae in he calyx. Re ac o muscles well de eloped and es ic ed. S olons wi h 9-12 canals. Scle i es sca e ed o e he su ace o he calyx. Scle i e ypes: c osses and " used quad uple s", shu les and six- adia es. Polyps wi hou scle i es. Rolandia co alloides Lacaze-Du hie s, 1900 ( igs 3-4) Rolandia co alloides Lacaze-Du hie s, 1900: 424-449, pis 15-16, desc ip ion ex e nal mo phology, scle- i es and his ology, nea Alge ia coas ; Molande , 1929: 41-42, only ema ks; Rossi, 1965: 163 & 174, eco d and ecology in co alligenous communi y, Pun a di Mesco (I aly). Rolandia osea; Weinbe g, 1978a: 167-173, pis 16-18, desc ip ion o ex e nal mo phology, scle i es, dis- cussion and synonymy, eggs obse ed om May o Sep embe , Banyuls-su -Me , Ad ia ic sea, Gul o Naples, Gul o Ma seille, and nea Alge ia. Al hough he au ho included Thomson's eco d in his synonymy appa en ly he does no admi ha R. osea could be p esen in he A lan ic Ocean; Weinbe g, 1978b: 39, o o 4, eco d and jus some ecological da a; Gili, 1982: 114 & 116, igs 57a-c, 58d, b ie desc ip ion and ecology, Banyuls de la Ma enda, Illes Medes. Rhizoxenia osea; Ma ion, 1882: 985, eco d, on Poseidonia sp. hizomes, 15 m, Gul o Ma seille; Thom- son, 1927: Pl. 3, igs 2-4, 6, 12; b ie desc ip ion bu su icien o ecognizing he " used quad u- ple s", 845 m, Azo es. E ago a co ii B och, 1935: 5-8, ig. 2, desc ip ion ex e nal mo phology, scle i es show clea ly he " used-quad uple s", 15-20 m, Ad ia ic Sea (Spli ). Zoan ha co ii; Pax & Mulle , 1962: 263-264, ig. 140, b ie desc ip ion and some da a o scle i es which a e la ge han hose in Zoan ha osea, Ad ia ico (Spli ), 15-20 m. Sa codic yon ca ena um; Be enguie , 1954: 62-65, pis 5-71; desc ip ion, wo ecological g ow h o ms, R. co alloides and S. ca ena um synonymous, Gul o Ma seille. ? E ago a osea Philippi, 1842: 36, ig. 2, b ie desc ip ion, Naples. No Zoan ha halassan hos Lesson, 1826, e e ed o Cla ula ia, see Weinbe g, 1978a. No Rhizoxenia halassan hos; Eh enbe g, 1834: 55, only eco d and new combina ion. No Rhizoxenia osea; on Koch, 1891: 661-663, ig. 13, con usion wi h an Alcyonium species, Naples. Ma e ial.- Spain: (LNM/PI 101) C ina is, 36°10.10'N 5°23.70'W, ii.1990, h ee colonies, in ali o al 20-25 m, P.J. Lopez leg.; (RMNH Cael. 23985) La A una a, 36°10.54'N 5°20.00'W, iii.1988, wo colonies, ci cali o al 40 m, P.J. Lopez leg.; (RMNH Cael. 23986) Gib al a ha bou , 36°08.90'N 5°21.60'W, .1992, wo colonies, ci cali o al28 m, P.J. Lopez leg.; (MNCN 2.04/726) Columb e es Islands (Wes - e n Medi e anean), 39°53.56'N 0040.20'E, 16. ii.96, one colony, 25 m, in ali o al, Fauna-IV c uise (s n. 278B8); (MNCN 2.04/727) Columb e es Islands (Wes e n Medi e anean), 39°51.27'N 0040.55'E, 18. ii.96, wo colonies, 30 m, in ali o al, Fauna-IV c uise (s n. 285B22); (MNCN 2.04/728) Colum- b e es Islands (Wes e n Medi e anean), 30049.77'N 0032.10'E, 2O. ii.96, one colony, 10 m, in ali o al, Fauna-IV c uise (s n. 296B7); (MNCN 2.04/731) Ho migas Islands, 37°39.52'N 0038.92'W, 21. ii.96, ." Ocana e al. A su ey o he gene a Sa codic yon and Rolandia. Zool. Med. Leiden 73 (2000) 420 Fig. 3. Scle i es o Rolandia co alloides. 1-13 an hos ele; 14-25 s olon. Scale ba : 75 pm. Ocana e al. A su ey o he gene a Sa codic yon and Ro/andia. Zool. Med. Leiden 73 (2000) 421 Fig. 4. His ology o Ro/andia co al/oides. A, c oss-sec ion o ilamen s and e ac o muscles. B, c oss- sec ion o a en acle. C, wo polyps embedded wi h a connec ing (solenial) canal. D, de ail o he canal. E, solenium isola ed in he mesogloea, F, h ee polyps embedded in he mesogloea. Scale ba s: A, B & E = 50 pm; C &F = 200 pm.