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.