Bijdragen tot de Dierkunde , 56 (2): 232-246 — 1986

Mediterranean Octocorallia: descriptionof Clavularia

carpediem n. sp. and synonymy of Clavularia crassa and

C. ochracea on etho-ecological grounds

by

Steven Weinberg

Institute of Taxonomic Zoology, University of Amsterdam, The Netherlands')

Abstract since then. As a result, the Mediterranean

octocoral fauna is quite well known, especially from the A new stoloniferan octocoral Mediterranean the circalittoral species that have recently been (Morocco) is described under the name Clavularia carpediem and critically compared with C. marioni Von Koch, 1891 revised (Weinberg, 1976, 1977, 1978). Only

ofthe and C. arctica (Sars, 1861). A thorough redescription three stoloniferan species have been described latter species is given. in the Cornularia as being very common area: The recent discovery that the common rock-dwelling (Pallas, 1766), Clavularia ochracea Von hitherto referred C. cornucopiae colonies belonging to a species to as 1878 and ochracea Von 1878 incubate their the outside Koch, Sarcodictyon roseum (Philippi, Koch, eggs on

2 of the rather unusual fact polyps, a among Octocorallia, 1842) ).

ofthis with C. leads to the species crassa (Milne synonymy Clavularia crassa (Milne Edwards, 1848), a far considered Edwards, 1848), so a rare species living live species believed to exclusively as an epi- the Mediterranean exclusively as an epibiont on sea-grass the marine biont on angiosperm Posidonia . oceanica is much while the rarer, two remaining

Résumé species appear to be very rare: Clavularia

( “Anthelia”) inermis, described by Berenguier Sous le nom Clavularia carpediem un nouvel Octocoralliaire but of which the type-specimen has Stolonifère méditerranéen est décrit des côtes marocaines, (1954), been have been et une comparaison critique est donnée de la nouvelle unfortunately lost, may

C. Von et C. espèce avec marioni Koch, 1891 arctica (Sars, rediscovered near Benidorm, Spain (Williams,

1861). Cette dernière est redécrite en détail. The other Clavularia pers. comm.). species, La découverte récente du fait les Stolonifères que com- marioni, originally described from Naples by substrats le muns des durs, connus jusqu’à présent sous Von Koch (1891), and of which the nom de C. ochracea Von Koch, 1878 incubent leurs oeufs type-

lost as was à l’extérieur des polypes, un phénomèneplutôt exception- specimen was well, subsequently nel chez les Octocoralliaires, conduit à la synonymie de found off Monaco, off Casablanca and around

C. l’on cette espèce avec crassa (Milne Edwards, 1848) que the Azores, in depths ranging from 845-2165 m considérait vivant jusqu’àprésent comme une espèce rare, (Thomson, 1927; Tixier-Durivault & d'Hondt, exclusivement en épibionte sur la Posidonie (Posidonia

In a recent of the oceanica). 1975). redescription species I from (Weinberg, 1978) studied a specimen

Naples (Zoologisch Museum Amsterdam coll.

the known INTRODUCTION no. Coel. 7831), presently only

specimen from the type-locality.

The study of Octocorallia (Coelenterata, In July and September 1982, while diving

in the MediterraneanSea of Morocco ) originated along the Mediterranean coast near in and has been pre-Linnaean times, going on Cabo Negro, an area extremely rich in

') Present address: European School, Boulevard Konrad Manuel (1981) brought the congenerity of Sarcodictyon

Adenauer, L-1115 Luxembourg. Forbes, 1849 and Rolandia De Lacaze-Duthiers, 1900 to

2 Still Rolandia in and established former ) described as rosea (Philippi, 1842) my my attention, correctly the name

Stolonifera the valid paper on Mediterranean (Weinberg, 1978). as only one by priority. BIJDRAGEN TOT DE DIERKUNDE, 56 (2) - 1986 233

3 I found stoloniferan which consist of Octocorallia ), a Coel. 8328, 8329, 8330). The colonies

reminded of C. marioni the size of its of 35-150 at me by groups ca. (mean 90) polyps rising

I collected 1-4 2 from polyps. some colonies. Closer study intervals of mm (mean mm) a

of these C. network. revealed that although resembling stolonal The colonies appear as tight

bear clusters marioni, the specimens more resemblance of polyps (plate I, fig. A), even on

much wider to the boreal species C. arctica. The differences substrata that would allow for spac-

stolons between the Moroccan species and C. arctica are ing of the individual polyps. The are

that the former deserves of broad such, however, a tax- 2.5-20 mm wide, often consisting a

onomical of its In this the and flat membrane from which the arise. status own. paper polyps

The consist of 5.5-9 new species Clavularia carpediem will be described polyps (fig. 1) a mm high

and compared with C. marioni and C. arctica. anthostele which is completely stiffened by eight

The Furthermore, entirely new etho-ecological rows of densely packed sclerites. proximal

of which is observations on the species formerly described part the anthocodia, completely

few C. ochracea lead the of this retractile into the bears a as to synonymy taxon anthostele, only

with C. sclerites. On the distal of the anthostele crassa. part a

and with about halfa dozen crown points occur,

sclerites between each of MATERIAL AND METHODS intermediary pair

The consists of 5 9 points. crown to parallel

Living specimens were observed and photographed under of slender and rows long, sclerites, the spindle- water by meansof SCUBA divingprior to collecting. The sclerites of the shaped points are prolonged on were kept overnight in a solution of 0.5% the external side of the tentacles MgS04 in seawater in order to achieve anesthesia before by sparsely

fixation in 10% formalin. Subsequently, the colonies were distributed flat irregular tentacular sclerites

transferred tojars containing 70% ethanol. All specimens (%• 2). in the collections of the Museum of are kept Zoologisch

the of Amsterdam University (ZMA). These tentacular irregular plates (fig. 3) bear of Clavularia obtained from the Specimens arctica were few only a tubercles. Zoological Museum of the University of Bergen, Norway 49-167 107 Length: [Am (mean [Am). (ZMB: 457, 578, 579, 12162), the Zoological Museum of Width: 16-45 30 the University ofOslo, Norway (ZMO: B847, B1208) and |im (mean [im).

the Zoological Museum of the University ofCopenhagen, Average length/width ratio: 3.6.

Denmark numbers dates of (collection are registration;

The sclerites of the crown and points ZMK: 20101884 (3 x ), 05101889 (2 x), 15051890, (fig. 4)

undated with the consist of slender with rounded 04191896, 26111908, 22121953, and two very spindles

mention: "Tjalfe"). thorns.

Preparations of sclerites were obtained by dissolution of Length: 239-429 [Am (mean 295 [Am) the soft tissues in cold and concentrated sodium Width: 26-37 31 [im (mean [Am). hypochlorite (Javel). ratio: 9.5. The follow the Average length/width morphological descriptions nomen-

clature of Bayer et al. (1983). The anthocodial sclerites (fig. 5) consist of

relatively smooth spindles with rounded thorns.

Clavularia carpediem n. sp. Length: 135-300 [Am (mean 208 [Am).

Width: 26-41 32 [xm (mean [Am).

Description Average length/width ratio: 6.5.

Five colonies have been studied, the holotype The sclerites of the anthostele (fig. 6) consist of

(ZMA Coel. 8327) and four paratypes (ZMA two different forms: plump spindles densely

3 ) The following species were encountered: Clavularia Groot & Weinberg, 1982), Eunicella singularis (Esper,

crassa (Milne Edwards, 1848), Sarcodictyon roseum (Philippi, 1794), E. verrucosa (Pallas, 1766), Lophogorgia ceratophyta

1842), Maasella edwardsi (De Lacaze-Duthiers, 1888), (Linnaeus, 1758), L. viminalis (Pallas, 1766) and

Paralcyonium spinulosum (Delle Chiaje, 1822), Alcyonium Paramuricea clavata (Risso, 1826).

acaule Marion, 1878, A. coralloides (Pallas, 1766) (see also 234 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

1. Two of Clavularia the other retracted. Sclerites occur Fig. polyps carpediem n. sp., one completely expanded, partly

in six distinct = = = = = stolon. body regions: te tentacles, po points, cr crown, ac anthocodia, as anthostele, st = - 235 BIJDRAGEN TOT DE DIERKUNDE, 56 (2) 1986

2. Detail of tentacle of Clavularia with sclerites of flat Fig. a carpediem n. sp., consisting irregular plates.

3. Tentacular of Clavularia of Fig. sclerites carpediem n. sp., consisting irregular plates

covered with tubercles, and slender spindles The stolonal sclerites (fig. 7) consist of the same

with rounded thorns. categories as encountered in the anthostele,

the surface of the slender Plump spindles: although processes

149-378 286 are more and sometimes Length: fjim (mean [Jim). spindles pronounced

Width: 53-71 (im (mean 61 fxm). more truncated and wart-like.

Average length/width ratio: 4.7.

Slender spindles: Plump spindles:

94-294 169 212-290 Length: [xm (mean (Jim). Length: (im.

Width: 51-55 Width: 24-37 |im (mean 31 fim). fim.

Average length/width ratio: 5.5. Average length/width ratio: 5.4. 236 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

4. Sclerites of and of Clavularia slender with rounded thorns. Fig. crown points carpediem n. sp.: spindles

Slender Cabo fair of sedi- spindles: around Negro carry a amount

90-261 192 the avoid Length: [im (mean [im). ment, species clearly seems to

Width: 26-45 34 substrata where silt accumulates. It should be (im (mean p).

Average length/width ratio: 5.7. noted that typical colonies of C. ochracea ( = C.

see next section) were found in the same in limited crassa, Cross-shaped sclerites occur numbers habitats. throughout the colony. So far, in spite of intensive prospecting, the

was not encountered in either Distribution and ecology new species

shallower or deeper habitats near the type- The been species has encountered so far in locality. shallow water (15-22 m depth) north of Cabo

Negro, near M'diq, Morocco. The colonies Etymology the side of rocks covered grow on upper by

calcareous encrusting algae or on the bare The specific name carpediem should be treated as

of stumps of gorgonian axes. Although the waters an "arbitrary combination letters" in BIJDRAGEN TOT DE DIERKUNDE, 56 (2) - 1986 237

5. Anthocodial sclerites of Clavularia with, rounded thorns. Fig. carpediem n. sp.: spindles

accordance with article ll.b.iii of the Interna- and Koren & Danielssen (1883) were completely tional Code of Zoological Nomenclature (3rd revised by Broch (1912). ed., 1985). Classicists, however, will recognize In the specimen of C. marionifrom Bocca Pic- here 70 the a contraction of "Carpe diem", a Latin cola, Naples, m depth, type-locality

the Coel. the distance between the proverb meaning "pluck day" (i.e. "enjoy (ZMA 7831), up

which life"), referring to the holiday during the to 14 mm tall polyps is 3-8 mm. The polypean author discovered the sclerites 40-210 124 new species. are [Am long (mean [Am) and

the stolonal sclerites measure 240-350 [Am

(mean 295 [Am). Both are rather smooth COMPARISON OF CLAVULARIA CARPEDIEM low thorns. For spindles with a more detailed WITH C. MARIONI VON KOCH, 1891 AND of this C. ARCTICA (SARS, 1861) description sample, see Weinberg (1978).

Because of the close resemblance of Clavularia In the original description (Von Koch, 1891:

marioni and C. and XXV carpediem with C. arctica, a fig. 11 pi. figs. 6-9) the length of the

of the main characteris- sclerites is 40-240 thorough comparison polypean some [Am (average tics of the three species was carried out. The approximately 150 [Am), while those of the results summarized in table I. are stolon measure 100-300 [Am (average approx-

imately 280 [Am).

C. marioni

Conclusion This species was described in detail by Von

Koch (1891) and Weinberg (1978), while the C. marioni has somewhat smaller and plumper

original descriptions of C. arctica by Sars (1861) polyps with shorter tentacles than C. carpediem 238 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

6. anthostele of Clavularia covered with tubercles. Bot- Fig. Sclerites of carpediem n. sp. Top row: plump spindles densely tom row: slender spindles with rounded thorns.

(but only preserved specimens of the former water species, while C. marioni is restricted to species have been observed!) and the polyps are deeper water.

each than is in more spaced from other the case

and C. carpediem. The crown points of C. marioni C. arctica contain less sclerites than C. carpediem. Its pol-

sclerites obtained ypean are smaller, while stolonal Specimens of this species were from sclerites in are longer. The plump spindles with the Zoological Museums Copenhagen (Den-

tubercles in and coarse that occur anthostele mark), in Oslo (Norway) and in Bergen (Nor-

of C. absent in C. marioni. ZMB stolon carpediem are way), including the specimens nrs. 578

Although C. carpediem occurs within the known and ZMB 579 collected by Koren & Danielssen, distributional range of C. marioni (Mediterra- being the syntypes of their C. arctica, which

there is who added the nean, Casablanca, Azores), no overlap were studiedby Broch, following in C. Dan.'s depth range, carpediem being a shallow- note (in Norvegian): "the type of Kor. & TOT 56 - 1986 239 BIJDRAGEN DE DIERKUNDE, (2)

7. Sclerites of stolon of Clavularia covered with tubercles and slender Fig. carpediem n. sp.: plump spindles densely

spindles with rounded thorns.

Table I Cl. described arctica, as n. sp. The species is,

described M. Sars of Cc Clavularia however, previously by as: Comparison of the main characteristics =

Cm = marioni Von 1891 and Rhizoxenia carpediem n. sp., C. Koch, arctica”.

Ca = C. arctica Data from Von Koch (Sars, 1861). (1891), Another specimen, ZMB 457, identified by Broch and Weinberg (1978) new observations; (1912), carries label Sars, a reading (in Norvegian): * = important differences between Cc and the other "Loc. Vardo for is possibly a writing error species. Vadso". Since the descriptions by Koren &

Cc Cm Ca Danielssen (1883) and Broch (1912) contain

data the I only few numerical on sclerites, add height anthostele (ram) 7.5-9 7.5 9-16

a new of the maximum height (mm) 16 14 22 description syntype (ZMB 578): maximum width polyp (mm) 2.5 3.0 3.0 Vadso, 50-60 fathoms tentacles 7.5 5.5 4.0 Locality: (90-110 m). max. length (mm)

the consists of a of number of pinnulae 8-13 8-10 10-11 Description: sample piece 2 wide stolon is distance between polyps stone, on which a mm found

1-4 3-8 3-8 (mm)* with about 6 polyps still attached to it, and

about detached sclerites anthostele: 20 polyps with a maximum maximum 720 width of and maximum of length ((xm)* 380 240 2.5 mm a height 11.5

170 125 425 average length ((xm)* mm (Koren & Danielssen, 1883, also mention 5.1 4.9 4.4 average shape (L/W)* 10-12 mm, but Broch, 1912, has described sclerites stolon: polyps up to 22 mm). maximum length (|xm)* 290 350 510 Some sclerites are brown, filled with a 190 295 290 average length ((im)' matrix. 5.5 6.8 4.5 fibrous, average shape (L/W)* organic Tentacular sclerites (fig. 8): flat, irregular observed 15-20 70-2165 35-300 depth range (m)* 103-243 165 plates; length: [Am (mean [Am), 240 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

Fig. 8. Sclerites of tentacles of Clavularia arctica (ZMB 578), consisting of irregular plates.

Fig. 9. Sclerites of anthocodia of Clavularia arctica (ZMB 578): spindles with rounded thorns and some warts. 56 - 1986 241 BIJDRAGEN TOT DE DIERKUNDE, (2)

width: 29-60 45 Anthostele [im (mean [Am), average (fig. 11): large, mostly plump,

140-660 length/width ratio: 3.6. tuberculate spindles; length: [xm (mean

Crown and slender 423 width: 33-140 97 points (fig. 10): spindles (xm), p (mean [xm), with rounded 194-480 ratio: 4.4. small, thorns; length: average length/width

smooth [im (mean 351 [Am), width: 33-70 [j.m (mean 48 Stolon (fig. 12): rather spindles,

ratio: 7.4. small with [Jim), average length/width many ones, some plump spindles

Anthocodia with rounded 78-410 292 (fig. 9): spindles tubercles, length: |im (mean fxm), thorns and some tubercles; length: 175-330 [im width: 29-95 |im (mean 64 [xm), average

Broch mentions (mean 237 [Am), width: 25-58 [Am (mean 42 length/width ratio: 4.5. (1912)

ratio: 5.7. stolonal sclerites of 510 [Am), average length/width up to [xm.

Fig. 10. Sclerites of crown and points of Clavularia arctica (ZMB 578): slender spindles with small, rounded thorns. 242 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

Fig. 11. Sclerites of anthostele of Clavularia arctica (ZMB 578): plump, tuberculate spindles. - 1986 243 BIJDRAGEN TOT DE DIERKUNDE, 56 (2)

12. Sclerites of stolon of Clavularia arctica thorned tuberculate ones. Fig. (ZMB 578): many small, spindles, some larger, 244 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

Conclusion tions on their different modes of reproduction.

The rather in which C. peculiar way crassa in In C. arctica the polyps are larger than C. been incubates its eggs has recently observed by tentacles shorter carpediem, while the are (pre- Tixier-Durivault d'Hondt & (1975). However, served The the specimens!). polypean as well as reconfirmation of planulae emission by C. stolonal sclerites are much larger than the cor- ochracea is needed to establish the validity of two C. the responding ones in carpediem, and shapes distinct species with certainty." different from of the sclerites are definitely those of C. carpediem. C. arctica occurs deeper month of 1984 I made During the August some than C. carpediem, and mostly in boreal waters underwater observations in several stations (Madsen, 1944), although Thomson (1927) that well near Banyuls-sur-Mer (France), might mentions the species from deeper water around be conclusive to solve this taxonomical riddle. the Azores. No distributional overlap occurs. On vertical rock surfaces between 5 and 10 m Neither of the species have been encountered so depth there exist large populations of what I far around the British Isles or the Atlantic along the regarded so far, because of habitat, as coasts of France, Spain and Portugal. On the 4th of belonging to Clavularia ochracea.

observed for the first time col- August 1984, I a SYNONYMY OF CLAVULARIA OCHRACEA VON its under the ten- ony incubating orange eggs KOCH, 1878 AND C. CRASSA (MILNE EDWARDS,

tacles, on the outside of the I: 1848) polyps (see plate with fig. C, and compare Kowalevsky &

In revision of the Mediterranean I, 1 and la and my Marion, 1883, plate figs.

d'Hondt In Stolonifera (Weinberg, 1978), I paid particular & Tixier-Durivault, 1975, fig. 4). attention the Clavularia. At that time the the was to following days same phenomenon there lot in was still a of confusion, and six species observed on dozens of colonies, depths rang-

the 9th of names existed: marioni Von Koch, 1891, crassa ing from 5 to 10 m, and on August

(Milne Edwards, 1848), ochracea Von Koch, the first planulae detached themselves from the

mother The 1878, petricola Kowalevsky & Marion, 1883, polyps. same phenomenon was

Tixier-Durivault 1969 observed between 16th and inordinata & Lafargue, again July August and steveninoae d'Hondt & Tixier-Durivault, 5th, 1986.

established that the 1975. So it has been clearly now

I show was able to that several of these were common rock-dwelling Mediterranean colonies

and I maintainedthree of Clavularia behave in the as was synonyms, species only. can same way

Besides Clavularia marioni from I known from deeper water, already the colonies living on sea-

C. the the It still that described crassa as species living on grass. seems possible, however, rhizomes of the marine angiosperm Posidonia Kowalevsky & Marion's (1883) observations oceanica and its the outside of and on incubating eggs at were correct, that, perhaps depending the polyps, under the tentacles, and Clavularia environmental conditions, these clavularians ochracea the switch from of their as species living on (see Weinberg, can one way incubating

rock, the tunicate Microcosmus and to another. 1979): spp., eggs even on the crustacean Pisa nodipes, incubating Since the important ethological criterion of

octocoral internal external its eggs internally, in normal fashion, vs. embryological develop-

the free- has thus there be on mesenteries and expulsing ment disappeared, seems to

valid Clavularia into swimming planula larvae. no more reason to separate

I 1978: and that lives However, concluded (Weinberg, a rock-dwelling species one on

"I wish to stress fact that the I therefore conclude the 160): the separa- sea-grass. to synonymy tion of the C. and their the species crassa C. ochracea, of the two species, where according to

rule morphology being very similar, is based mainly priority Clavularia crassa (Milne Edwards,

& henceforth. upon Kowalewsky Marion's (1883) observa- 1848) should be used - 245/I BIJDRAGEN TOT DE DIERKUNDE, 56 (2) 1986

Plate I

Fig. A (top): Holotype of

Clavularia carpediem n. sp. (ZMA Coel. 8327), photo-

graphed in situ at 22 m

depth north of Cabo

Negro, Morocco. Note that

the polyps are densely

grouped, a typical feature

of the species (1.7 x).

Fig. B (left): Expanded

polyps of Clavularia crassa

rock surface growing on a

amidst sponges, Banyuls-

sur-Mer, ca. 10 m depth.

Fig. C (right): Colony of

Clavularia crassa in the same

habitat as above. The

orange eggs are being

incubated in a mucous

sheath at the outside of the

polyps, under the ten-

tacles. (All underwater pic-

tures by the author.) 246 S. WEINBERG - MEDITERRANEAN OCTOCORALLIA

A. A.-F. 1883. Documents ACKNOWLEDGEMENTS KOWALEVSKY, & MARION, pour

l'histoire embryogenique des Alcyonaires. Annls. The for loan of following persons are acknowledged Mus. Hist. nat. Marseille, (Zool.) 1 (4): 1-50, pis. material: Dr. Endre Willassen (Zoological Museum, I-V. University of Bergen, Norway), Dr. Marit Christiansen MADSEN, F. J., 1944. Octocorallia. Danish Ingolf Exped., (Zoological Museum of the University of Oslo, Norway), 5 (13): 1-65, pi. I. Dr. Kay Petersen (Zoological Museum, University of MANUEL, R. L., 1981. British Anthozoa. Synopses of the Copenhagen, Denmark) and Dr. Rob van Soest (Institute British Fauna (New Series), 18: 1-241 (Academic of Taxonomic Zoology, University of Amsterdam, The Press, London etc.). Netherlands). The magazine "Vita Marina" kindly con- MILNE EDWARDS, [H.], 1848. Les Zoophytes. In: G. C. tributed the separation negatives for plate I. CUVIER al. Le distribue et eds., regne d'apres

son organisation: [i-v], 1-160, 1-94; Atlas: [i-v], 1- REFERENCES 97, pis. 1-97 (Fortin, Masson et Cie., Paris).

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First draft received: 12 May 1986

Revised draft received: 10 July 1986