North-eastern Atlantic and Mediterranean species of Comulariidae Dana, 1846 (anthozoa: Stoloniferal) with the description of a new genus

No h-eas e n A lan ic and Medi e anean species o Co nula iidae Dana, 1846 (An hozoa: S oloni e a) wi h he desc ip ion o a new genus P.J. López-González, O. Ocaña, J.C. Ga cía-Gómez & J. Núñez P.J. López-González; O. Ocaña; J.C. Ga cía-Gómez & J. Núnez. No h-eas e n A lan ic and Medi e a- nean species o Co nula iidae Dana, 1846 (An hozoa: S oloni e a) wi h he desc ip ion o a new genus. Zool. Med. Leiden 69 (20), 29.xii.1995: 261-272, igs 1-21.— ISSN 0024-0672. P.J. López-González, Depa amen o de Biología Animal, Vege al y Ecología, Facul ad de Ciencias Expe imen ales, Uni e sidad de Jaen, Pa aje "Las Lagunillas" s/n, 23071 Jaen, Spain. O. Ocaña & J. Núñez, Depa amen o de Zoología, Facul ad de Biología, Uni e sidad de La Laguna, Tene i e (Cana y Islands), Spain. J.C. Ga cía-Gómez, Labo a o io de Biología Ma ina, Depa amen o de Fisiologia y Biología Animal, Facul ad de Biología, Uni e sidad de Se illa, Apdo. 1095, 41080 Se illa, Spain. Key wo ds: An hozoa; S oloni e a; Co nula iidae; Ce e a gen. no . The No h-eas e n A lan ic and Medi e anean species o he amily Co nula iidae a e e-examined and axonomically e ised. This amily is mainly cha ac e ized by he absence o scle i es. Up o now, only one genus, Co nula ia Lama ck, 1816, has been assigned o he amily. S udies o he in e nal and ex e nal ana omy o he Eu opean species o Co nula iidae, ca ied ou o he i s ime on a compa - a i e basis, leads us o p opose a new genus, Ce e a. New combina ions o he species included in he amily a e p oposed. The main cha ac e s o Ce e a gen. no . a e he p esence o a ine chi inous pe i- de m, a ious communica ion canals in he s olons and he absence o a addi ional chi inous laye ha co e s he pe ide m. Co nula ia a lan ica Johnson, 1861, is desc ibed o he i s ime subsequen o he o iginal desc ip ion, and is he e designa ed as he ype species o he new genus. The axonom- ic signi icance o he ana omical and his ological cha ac e s in he Co nula iidae a e discussed and he sys ema ic posi ion o he species p e iously a ibu ed o he genus Co nula ia is eassessed. In oduc ion Se e al e isions o he s oloni e ous oc oco als ha e been ca ied ou o e he las cen u y (e.g. Hickson, 1894, Goha , 1940, Weinbe g, 1978, and Baye , 1981). Only one genus, Co nula ia Lama ck, 1816, has been included in he amily Co - nula iidae Dana, 1846. I has been desc ibed as lacking scle i es, and ha ing sepa a e polyps uni ed by s olons. In he ype species, Co nula ia co nucopiae (Pallas, 1766), he s olons ha e only one canal, a hin pe ide m and a hick ex e nal chi inous cu icle co e ing he an hos eles and s olons (Benke and Hündgen, 1984). Howe e , se e al au ho s (Ligh , 1915; U inomi, 1950, 1955; Suzuki, 1971) also included species in he genus ha ing s olons wi h mo e han one canal, and he diagnosis o he genus and amily we e he e o e adap ed o include hese species. In Eu ope he amily Co nula iidae is ep esen ed by wo species, Co nula ia co - nucopiae and Ce e a a lan ica (Johnson, 1861) comb. no . Co nula ia co nucopiae is widely dis ibu ed in he wes e n Medi e anean and is also known om he A lan- ic coas s o sou h-wes e n Eu ope. Ce e a a lan ica was desc ibed om Madei a by Johnson (1861) as Co nula ia a lan ica, and had no been epo ed since i s o iginal desc ip ion. This species is edesc ibed he e and assigned o a new genus, oge he wi h o he , Japanese species. 262 López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) A compa a i e s udy o he ana omy o he wo Eu opean species e ealed use ul cha ac e s o he classi ica ion o he amily Co nula iidae. Ma e ial and Me hods Colonies we e collec ed om ha d subs a a by hand in he in e idal egion, and by sno keling o SCUBA di ing in he uppe in ali o al a ea. They we e anaes he - ised in a sa u a ed solu ion o sea wa e and men hol, and p ese ed in 6-8% o malin. A s e eomic oscope was used o ex e nal ana omical s udies. His ological sec- ions, 8 u n hick, o dissec ed polyps and s olons we e made using he classic me h- od o opog aphical s aining o Cajal (Gabe, 1968). Specimens we e deposi ed in he Na ionaal Na uu his o isch Museum, o me ly Rijksmuseum an Na uu lijke His o ie (RMNH) o Leiden, he Museo Nacional de Ciências Na u ales o Mad id (MNCN), Spain, and he Labo a o io de Biologia Ma i- na (LMB) o he Uni e si y o Se ille, Spain. Desc ip i e pa O de S oloni e a Hickson, 1883 Family Co nula iidae Dana, 1846 Diagnosis.— Colonies small, wi hou scle i es. Polyps no joined in a common coenenchyma ous mass, bu connec ed by s olons wi h one o se e al in e nal canals. An hos eles and s olons co e ed by a pe ide m, and some imes a hick addi ional cu icle. Type genus: Co nula ia Lama ck, 1816. Co nula ia Lama ck, 1816 Diagnosis.— Co nula iidae wi h polyps connec ed by s olons ha a e weakly a ached o subs a um, a iable in leng h, ci cula in c oss-sec ion, and wi h one in e nal ca i y. S olons and an hos eles co e ed by a pe ide m and by a hick chi in- ous en elope, o ming a kind o heca; an hocodiae e ac able in o he heca. Type species: Tubula ia co nucopiae Pallas, 1766, by mono ypy. Co nula ia co nucopiae (Pallas, 1766) ( igs 1-9) Tubula ia co nucopiae Pallas, 1766: 80, ig. 37. Co nula ia ugosa Lama ck, 1816:112. Co nula ia co nucopiae; Schweigge , 1820: 425; Hickson, 1894: 334. No Co nula ia co nucopiae; Manuel, 1979: 393,1981: 34. Fo u he e e ences, see Weinbe g, 1978:141. Ma e ial.— RMNH Coel. 23136 (San a Ma ía del Ma , Cadiz, sou he n Spain, 36°11'N - 6°2'W, in e i- López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) 263 dal, 31.Í.1990, 1 agmen ); RMNH Coei. 23137 (El Cha o, Cadiz, sou he n Spain, 36°31'N - 6°16'W, in e idal, 9.Ü.1990, 1 colony); RMNH Coel. 23138 (Canos de Meca, Cadiz, sou he n Spain 36°11'N - 6°2'W, in e idal, ll.ii.1990, 1 colony); RMNH Coel. 23139 (Ta i a, Cadiz, sou he n Spain, 36o0'48"N - 5°36'W, lim, 9.VÜ.1990, 1 agmen ); LBM (Pan alán de San Felipe, La Linea, Cadiz, sou he n Spain, 36°9'N - 5022'W, 6 m, 26. ii.1990, 2 colonies); LBM (Pun a Almina, Ceu a, Spanish ci y in No h A ica, 35°54'N - 5°16'W, 25 m, 13.V.1988, 2 colonies). Mo phology ( igs 1-2).— Colonies small; polyps connec ed by s olons o a iable leng h. S olons ci cula in c oss-sec ion, abou 0.2 mm in diame e ( ange 0.12-0.25 mm), co e ed by a hick chi inous cu icle ha does no s ongly adhe e o he sub- s a um. An hos ele also co e ed by a hick en elope. In e ac ed polyps, he uppe pa o he hick chi inous cu icle appea s o be sepa a ed om he polyp ( ig. 2). In expanded polyps, he limi o he pe ide m is isible on he polyp body as a bo le- neck (also epo ed by Benke and Hündgen, 1984) (see whi e a ows in ig. 1). Ten a- cles wi h 14-18 pinnules on each side. Basal junc ions o polyps and s olons na ow and o simila diame e as he s ol- ons. Polyps anslucen ly whi e and he chi inous en elope beige o b own. Dis ance be ween polyps a iable (1.7-7 mm in he ma e ial examined). Heigh o expanded polyps 6.5-10 mm. Ana omy ( igs 3-4).— Re ac ed polyps elonga ed ( ig. 3). S olons ci cula in Fig. 1. Co nula ia co nucopiae, gene al o m. Aqua ium pho og aph showing expanded polyps and he limi o he pe ide m (indica ed by whi e a ows). Scale ba : 2 mm. 264 López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) Fig. 2. Co nula ia co nucopiae, gene al o m. Habi us o expanded and e ac ed polyps and s olons. AC, an hocodia; AT, an hos ele; PH, pha ynx; RP e ac ed polyp; ST, s olon; TE, en acle. Scale ba : 2 mm. c oss-sec ion, wi h only one in e nal canal o 120-150 u n in diame e ( ig. 4). Ex e nal laye , wi h concen ic g oo es, 8-13 u n hick ( ig. 5). Adhe ence o o eign pa icles o he ex e io is minimal. Mesogloea educed o less han 1 u n in hickness. Gene al aspec and de elopmen o ec ode m and endode m simila . Muscula p ocesses we e no obse ed. Co e o he an hos ele a ying in hickness om i s base (5 u n) o he dis al edge o he heca, whe e i becomes educed (1 u n) ( ig. 6). Ec ode m, mesoglea and endode m o an hos ele and s olon simila in appea - ance. In he an hocodia he hickness o hese h ee laye s is abou 10 u n, in he an hos ele and s olon abou 2 u n. In he an hocodia he ec ode m and endode m can be hicke han he mesogloea wi h ec ode mal and endode mal muscle p ocesses on mesogloeal a cs. In he en acles, he hickness o he mesogloea is a he uni o m (4- 5 u n) and he longi udinal muscula u e is be e de eloped han he ci cula muscu- la u e ( ig. 7). Pha ynx in c oss-sec ion i egula in ou line. I s mesoglea less han 1 u n hick, i s endode m 20-40 um hick. The single well-de ined siphonoglyph appea s midway, in he lowe pa o he pha ynx. Mesen e ies sho , wi h kidney-shaped ilamen s and weakly de eloped muscu- López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) 265 Figs 3-9. Co nula ia co nucopiae, ana omical and his ological de ails. Fig. 3. Longi udinal sec ion o a polyp and i s s olon. Fig. 4. C oss-sec ion o s olon. Fig. 5. De ail o wall o s olon. Fig. 6. Longi udinal sec ion o a polyp, showing he di e ence in hickness o he mesogloea in an hos ele (bo om) and an hocodia ( op). Fig. 7. C oss-sec ion o a en acle. Fig. 8. C oss-sec ion o a mesen e y and e ac o muscles. Fig. 9. T ans e se sec ion o a mesen e ic ilamen . CC, chi inous cu icle; LM, longi udinal muscula u e; ME, mesogloea; PO, polyp; RM, e ac o mus- cula u e; ST, s olon. Scale ba s: in ig. 3, 800 u n; in ig. 4,100 u n; in ig. 5,9 pm; in ig. 6, 20 u n; in ig. 7, 35 u n; in ig. 8,20 u n and in ig. 9,40 u n. 266 López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) la u e ( ig. 9). Mo e dis ally, he longi udinal muscula u e o he mesen e ies is di - use ( ig. 8). Habi a .— Co nula ia co nucopiae occu s in he in e idal and uppe in ali o al a eas, mainly on sponges, b yozoans, ascidians, hyd ozoans and algae. Dis ibu ion.— The species has been epo ed om he wes e n Medi e anean (S iasny, 1941; Pax & Mülle , 1955, 1962; Weinbe g, 1978; Bibiloni & Co ne , 1980; Ocha an & Anadón, 1981; Gili, 1982; Roca, 1986) and he A lan ic coas s o F ance (Teissie , 1965), Po ugal (Saldanha, 1974), he Azo es (Tixie -Du i aul & d'Hond , 1974) and As ú ias (no he n Spain) (Ocha an & Anadon, 1981). Ce e a gen. no . Diagnosis.— Co nula iidae wi h polyps connec ed by la ened s olons ha a e s ongly a ached o he subs a um, a iable in leng h, i egula ly elonga e in c oss- sec ion, and wi h se e al in e nal canals. An hos eles and s olons co e ed by a weak pe ide m only. Type species: Co nula ia a lan ica Johnson, 1861, designa ed he ein. E ymology.— The name Ce e a has been adop ed in honou o M Joaquin Ce - e a, ecen ly deceased, who helped us in ob aining inancial suppo and wi h he o ganiza ion o ma ine biological expedi ions a ound he S ai o Gib al a . The gen- de o he name is eminine. Ce e a a lan ica (Johnson, 1861) comb. no . ( igs 10-21) Co nula ia a lan ica Johnson, 1861: 298. ?An helia ine mis Be enguie , 1954: 61. ?Co nula ia ine mis; Roca, 1986: 540. Ma e ial.— MNCN 02.04/0003 (Isla de Ta i a, Cadiz, sou he n Spain, 36°0'48'N - 5°36'W, 2 m, ii.1991, 1 colony); RMNH Coel. 23140 (San a Ma ía del Ma , Cadiz, sou he n Spain, 36°31'N - 6°17'W, in e i- dal, 30.ii.1990, some agmen s); RMNH Coel. 23141 (Isla de Ta i a, Cadiz sou he n Spain, 36°0'48MN - 5036'W, 2 m, 6.Ü.1991, 1 colony + 3 agmen s); RMNH Coel. 23142 (Ceu a, Spanish ci y in No h A i- ca, 35°53'N - 5°16'W, 3-5 m, 15. iii.1991, 3 agmen s); RMNH Coel. 23143 (idem, 9-12 m, 17. iii.1991,1 colony); RMNH Coel. 23144 (idem, 1 m, 20. iii.1991, 1 agmen ); RMNH Coel. 23158 (Candelá ia, Tene i e, Cana y Islands, 28o34*30"N - 16°20*W, 3 m, 4.Ü.1989, 2 colonies); RMNH Coel. 23159 (Pun a Hidalgo, Tene i e, Cana y Islands, 28°21'N - 16°22'W, 1 m, 9.iii.l990, 2 colonies); RMNH Coel. 23160 (Mon ana Roja, La Teji a, Tene i e, Cana y Islands, 28°30'N - 16°33'W, in e idal, unde s ones, 20.Í.1990, 2 colonies). Mo phology ( igs 10-11).— Colonies small; polyps up o 1 cm in heigh , connec - ed by la ened s olons o a iable leng h, and maximally 2-3 mm in wid h. Polyps anslucen ly o ange. An hos eles ( ep esen ed by he basalmos 2-3 mm o he polyps) and s olons co e ed by a hin b ownish pe ide m. Junc ions o polyps and s olons wide. Ten acles wi h 12-14 pinnules on each side. Ana omy ( igs 12-21).— Re ac ed polyps conical o sphe ical ( ig. 12). Ou line o s olons i egula ly elonga e in c oss-sec ion. Each s olon con aining 3-5 p incipal canals and o he s smalle in size ( ig. 17-18). Fo eign pa icles adhe ing o ex e nal Lopez-Gonzalez e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) 268 López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) Figs. 12-21. Ce e a a lan ica, ana omical and his ological de ails. Fig. 12. Longi udinal sec ion o a e ac ed polyp and i s s olon. Figs 13-14. De ails o wall o polyp, showing he mesogloeal lacunae and he weak chi inous cu icle. Fig. 15. T ans e se sec ion o an hocodia, showing he well-de eloped endode m. Fig. 16. De ail o ans e se sec ion o a en acle, displaying a hick mesogloea and he lon- gi udinal muscula u e. Fig. 17-18. C oss-sec ion o s olon, showing he mesogloeal canals. Fig. 19. T ans e se sec ion h ough a mesen e y. Fig. 20. Oocy e. Fig. 21. Packe o spe ma ogonia. CA, canal; CC, chi inous cu icle; EN, endode m; LM, longi udinal muscula u e; ME, mesogloea; ML, mesogleal lacunae; OV, oocy e; PO, polyp; RM, e ac o muscle; SP, packe o spe ma ogonia; ST, s olon. Scale ba s: in ig. 12,1 mm; in igs 13-14, 25 u n; in ig. 15, 37 u n; in ig. 16,15 u n; in igs 17-18, 360 pm; in ig. 19, 50 pm; in ig. 20, 35 u n and in ig. 21, 65 u n. López-González e al. Species o Co nula iidae. Zool. Med. Leiden 69 (1995) 269 su ace o pe ide m. Ec ode m sca cely de eloped. Mesogloea wi h a complex sys em o lacunae, lined wi h hin (5 um) endode m. Pe ide m o an hos eles simila o ha o s olons. Ec ode m sca cely de eloped. Mesoglea well-de ined, eaching i s maximal hickness in he an hos ele. La ge lacu- nae illed wi h cells p esen in mesogloea o bo h an hos eles and s olons ( igs 13-14). Endode m o an hos eles weakly de eloped, eaching i s maximal de elopmen dis- ally in he an hocodia ( ig. 15). Ec ode m o en acles and pinnules well de eloped, 10-12 u n hick; longi udinal muscula u e s ong, suppo ed by mesogloeal p ocesses, up o 10 u n hick in he pinnules and axis o he en acles. Ci cula muscula u e p esen bu no well ma ked. Endode m well de eloped, 4-5 u n hick ( ig. 16). Mesen e ies wi h a hin mesoglea and weak muscula u e ( ig. 19). The colony agmen s om San a Ma ía del Ma (Cadiz) (RMNH Coel. 23140) collec ed in Feb ua y 1990, con ain oocy es up o 69 u n in diame e ( ig. 20). The col- ony om Ta i a (MNCN 02.04/0003) collec ed in Feb ua y 1991, con ains well de el- oped packe s o spe ma ogonia, up o 205 u n in diame e ( ig. 21). In he pha ynx, he mesoglea is educed in hickness o 5 u n. Ec ode m well- de eloped, abou 45 pm hick. Siphonoglyph dis inc , wi h mesogloeal lacunae a i s mo e p oximal le el; i s cilia mode a ely long. Habi a .— The specimens we e collec ed unde s ones, in ock issu es and ca es in in e idal and uppe in ali o al a eas. Dis ibu ion.— So a , Ce e a a lan ica had only been epo ed om Madei a (Johnson, 1861). The p esen eco ds ex end he a ea o dis ibu ion o Tene i e (Cana y Islands) and he S ai o Gib al a . Discussion Th ee species o be accommoda ed in Co nula iidae ha e been epo ed om Eu opean wa e s, including he Azo es, Cana y Islands and Madei a, iz., Co nula ia co nucopiae (Pallas, 1766), Ce e a a lan ica (Johnson, 1861) and "An helia" ine mis Be enguie , 1954. The las -named species is s ill insu icien ly desc ibed and a ca e ul ana omical s udy is in ac necessa y o es ablish whe he i is di e en om Ce e a a lan ica (see below). Co nula ia co nucopiae is a well known Eu opean species (see dis ibu ion) and needs no u he discussion by i sel . Ce e a a lan ica (Johnson, 1861) was o iginally desc ibed om Madei a as Co nu- la ia a lan ica. Johnson's b ie desc ip ion o his species eads as ollows: "Basal band na ow, [...], des i u e o spicula, bu in es ed by a hin epide mis con aining pa i- cles o sand; when e ac ed, o ming a leshy hemisphe ical bu on". The cha ac e s men ioned in his desc ip ion jus i y i s inclusion in Co nula iidae, bu do no ma ch Co nula ia co nucopiae. In Co nula ia co nucopiae, he hicke anslucen chi inous en elope o he an hos ele o ms a heca. In ully e ac ed polyps he dis al pa o his heca is emp y ( ig. 2), an aspec well illus a ed by Weinbe g (1978). This heca is absen in Ce e a. Ano he di e ence exis s in he junc ion o polyp and s olon. In Co nula ia co nucopiae his junc ion is na ow, while i is wide in Ce e a a lan ica. The las -named species had no been epo ed again since i s o iginal desc ip ion. Ou