The family Haleciidae (Cnidaria: Hydrozoa) from the Strait of Gibraltar and nearby areas

The amily Haleciidae (Cnida ia: Hyd ozoa) om he S ai o Gib al a and nea by a eas M.D. Medel, F.G. Ga cía & W. Ve oo Medel, M.D., F.G. Ga cía & W. Ve oo . The amily Haleciidae (Cnida ia: Hyd ozoa) om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (3), 11.xii.1998: 29-50, igs 1-7.—ISSN 0024-0672. M.D. Medel & F.G. Ga cía, Labo a o io de Biología Ma ina (Zoología), Depa amen o de Fisiología y Biología animal, Facul ad de Biología, Uni e sidad de Se illa, España, Spain. W. Ve oo , Na ional Museum o Na u al His o y, Leiden, The Ne he lands. Key wo ds: Cnida ia; Hyd ozoa; Lep olida; Haleciidae; S ai o Gib al a ; geog aphical dis ibu ion. A aunis ic s udy o he amily Haleciidae om he S ai o Gib al a and nea by a eas has been made; se en species a e desc ibed, igu ed and discussed. In o ma ion abou he zoogeog aphical dis- ibu ion, and condi ions o subs a es a e con ibu ed. A key o dis inguish he species o his amily om he seas o he Ibe ian Peninsula is p esen ed, and he Ibe ian Haleciidae a e compa ed wi h hose om he Medi e anean and he B i ish Isles. In oduc ion The S ai o Gib al a is an impo an zoogeog aphical a ea, because i is a ba ie o he es ablishmen o many species in he A lan ic o in he Medi e anean, bu i is also he way o communica ion o species wi h a wide capaci y o dispe sion. The shallow wa e hyd ozoan auna om he S ai o Gib al a has been unknown o a long ime. Besides, many haleciids ha e a shallow wa e dis ibu ion. In he p esen pape , se en species o Haleciidae ound in his a ea a e desc ibed and discussed, and a key o iden i y he species o he amily Haleciidae om he Ibe ian Peninsula is p o ided. The Ibe ian Haleciidae auna is mainly cosmopoli an and sub opical- A lan ic; he aunis ic composi ion is simila o ha o he B i ish Isles, Mo occo, and he Medi e anean. Th ee species ha a e conside ed as Medi e anean endemics, ha e no been ound a he Ibe ian Peninsula. Two species conside ed A lan ic in o i- gin we e ound in he S ai o Gib al a , o which one was also ound also by Ramil & Ve oo (1992) in he Albo an Sea. Lis o he locali ies and species collec ed S a ion/locali y coo dina es dep h (in m) Coas al s a ions -C1 El Po il 37˚12.40'N,07˚07.50'W 0-1 Ophiodissa mi abilis (Hincks, 1868). -C2 Caños de Meca 36˚11.00'N,06˚01.00'W 0 Halecium liou illei Billa d, 1934. -C3 Isla de Ta i a 36˚48.00'N,05˚36.00'W 0-45 Halecium delica ulum Cough ey, 1876; Halecium sibogae ma ocanum Billa d, 1934. -C4 Pun a Ca ne o 36˚04.36'N,05˚25.36'W 0-25 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus, 1758). -C5 La Ballene a-Ge a es 36˚04.54'N,05˚25.36'W 3-15 Halecium delica ulum Cough ey, 1876. -C6 San Ga cía 36˚06.35'N,05˚25.36'W 5-28 ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 29 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)30 Halecium delica ulum Cough ey, 1876. -C7 Té mica 36˚10.50'N,05˚25.00'W 9-30 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus, 1758); Halecium enellum Hincks, 1861. -C8 C.E.P.S.A. 36˚10.60'N,05˚25.80'W 12-25 Halecium delica ulum Cough ey, 1876; Halecium enellum Hincks, 1861. -C9 San Felipe 36˚09.45'N,05˚21.48'W 2-9 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus, 1758). -C10 Pun a Eu opa 36˚06.00'N,05˚21.00'W 5-18 Halecium halecinum (Linneus, 1758); Halecium enellum Hincks, 1861. -C11 Pun a del Desna igado 35˚53.50'N,05˚16.20'W 0-18 Halecium delica ulum Cough ey, 1876. -C12 Pun a Almina 35˚54.00'N,05˚16.20'W 20-25 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus, 1758); Halecium liou illei Billa d, 1934; Halecium enellum Hincks, 1861. -C13 Pun a Saudiño 35˚54.35'N,05˚17.95'W 38 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus 1758); Halecium liou illei Billa d, 1934; Halecium pusillum (M. Sa s, 1857); Halecium enellum Hincks, 1861. -C14 Aguja del Campo 35˚54.20'N,05˚19.65'W 20 Halecium delica ulum Cough ey, 1876; Halecium halecinum (Linnaeus, 1758); Halecium liou illei Billa d, 1934; Halecium pusillum (M. Sa s, 1857). -C15 Benzú 35˚55.10'N,05˚23.25'W 25 Halecium halecinum (Linnaus, 1758); Halecium enellum Hincks, 1861. Campaign FAUNA I -F23 Pláce de las Bó edas 36˚24.05'N,05˚00.99'W 30-32 Halecium halecinum (Linnaeus, 1758). -F55 T a alga 36˚05.91'N,06˚08.40'W 38-42 Halecium halecinum (Linnaeus, 1758). -F66 o Cádiz 36˚30.90'N.06˚22.10'W 25-28 Halecium halecinum (Linnaeus, 1758). Campaign AR.SA.’92 -A2 Cos a de Doñana 36˚48.50'N,06˚37.10'W 21 Halecium halecinum (Linnaeus, 1758). -A3 Chipiona 36˚43.60'N,06˚34.20'W 23 Halecium halecinum (Linnaeus, 1758). -A6 o Chiclana 36˚19.70'N,06˚29.90'W 84 Halecium halecinum (Linnaues, 1758). -A8 o Cádiz 36˚21.50'N,06˚35.90'W 112 Halecium halecinum (Linnaeus, 1758). -A10 o Doñana 36˚54.10'N,06˚47.96'W 41 Halecium halecinum (Linnaeus, 1758). -A16 o Doñana 36˚56.40'N,06˚50.30'W 40 Halecium halecinum (Linnaeus, 1758). -A17 o Doñana 36˚37.02'N,06˚49.30'W 147 Halecium halecinum (Linnaeus, 1758). ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 30 31Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) Taxonomic esul s In ao de Haleciida Hincks, 1868 Supe amily Halecioidea Hincks, 1868 Family Haleciidae Hincks, 1868 Genus Halecium Oken, 1815 ( ig. 1) Halecium delica ulum Cough ey, 1876 Halecium delica ulum Cough ey, 1876: 299; Ve oo , 1959: 27-30, igs 4-5 (synonymy). Halecium g acile; Mo z-Kossowska, 1911: 335, igs 7-8, pl. 18 ig. 2. Halecium medi e aneum; S echow, 1919: 34. Halecium lexile; Leloup, 1940: 6. Halecium enellum; Ga cía Co ales e al., 1978: 9, igs 1-2. Ma e ial examined.— C3: (17.2.1991), 3-4 m, wi h gono hecae (RMNH- Coel. 27842); (7.6.1992), 3-13 m, wi h gono hecae. C4: (25.7.1991), 3 m, wi h gono hecae. C5: (26.7.1990), 3 m, wi h gono hecae. C6: (29.10.1994), 3-9 m, wi hou gono hecae. C7: (28.8.1991), 2-3 m, wi h gono hecae; (14.7.1993), 3-13 m, wi h gono hecae. C8: (16.7.1991), 18 m, wi hou gono hecae. C9: (23.11. 1991), 2-5 m, wi h gono hecae. C11: (8.5.1986), 0- 18 m, wi h gono hecae. C12: (13.5.1986), 35 m, wi h gono hecae. C13: (6.5.1986), 38 m, wi hou gono hecae. C14: (5.5. 1986), wi hou gono hecae. Desc ip ion.— Hyd ocauli monosiphonic o polysiphonic, g owing in dense g oups, i egula ly b anched in all di ec ions ( ig. 1a, b). Axis and b anches di ided in o in e nodes sepa a ed by oblique nodes, al e na ely di ec ed le and igh . Each in e node wi h a hyd opho e, placed on one side a he dis al end, and al e na ely di ec ed o igh and le ( ig. 1c). P ima y hyd o heca shallow, g adually widening owa ds he ape u e, wi h a basal diaph agm, and wi h a ow o desmocy es on he wall o a achmen o he hyd an h ( ig. 1e). Rim e e ed, pe pendicula o leng h axis o hyd opho e; his wi h a hickening o pe isa c (pseudodiaph agm) a he adcauline side. Se e al hyd o hecae may o igina e om he diaph agm o a p e ious hyd o heca, wi h mo e o less de eloped hyd opho es ( ig. 1d). Male gono hecae o oid, elonga ed and la e ally comp essed ( ig. 1g). Female gono hecae almos sphe ical and sligh ly la e ally comp essed ( ig. 1 ). Measu emen s (in mm).— Hyd ocalus, maximum heigh 25 In e nodes, leng h 0.40 - 0.64 diame e 0.09 - 0.13 Hyd o heca, leng h 0.03 - 0.05 diame e 0.13 - 0.19 Gono heca, leng h 0.5 - 1 diame e 0.275 - 0.425 Subs a e.— Halecium delica ulum is e y abundan in he a ea s udied. I has been ound g owing on seaweeds, b yozoans, ascidians, ba nacles, and on Syn hecium ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 31 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)32 Fig. 1. Halecium delica ulum Cough ey, 1876, om Isla de Ta i a (C3). a, b, shape o colony; c, pa o hyd ocladium; d, eno a ed hyd o heca; e, p ima y hyd o heca; , emale gono heca; g, male gono he- ca. Scales: a, b, 1 cm; c, d, , g, 0.5 mm; e, 0.1 mm. a b g c d e ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 32 33Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) e ansii (Ellis & Solande , 1786). Roca Ma inez (1986) ound i on Eudend ium acemo- sum (Gmelin, 1791); Ramil and Ve oo (1992) on Se ula ella gayi (Lamou oux, 1821), and Wa son (1994) on sponges. Dis ibu ion.— H. delica ulum is conside ed a ci cumglobal species, cha ac e is ic o opical and sub opical wa e s (Rees & Ve oo , 1987), pene a ing Paci ic wa e s as a no h as Japan and Canada, and going in he A lan ic as a no h as Mo occo; sou hwa ds i s dis ibu ions ex ends in o he An a c ic. I is common in he li o al zone o he F ench coas s and in he Medi e anean (Ramil & Ve oo , 1992). I is also ci ed om he A ic (Leloup, 1940). Following Boe o & Bouillon (1993), we con- side his species o be cosmopoli an. The dep h eco ds a e be ween 2 m (p esen wo k) and 2035 m (Ramil & Ve oo , 1992). Halecium halecinum (Linnaeus, 1758) ( ig. 2) Se ula ia halecina Linnaeus, 1758: 809. Halecium halecinum; Oken, 1815: 91; Ve oo , 1972: 25-26, ig. 3a. Thoa halecina; Johns on, 1838: 119-120, pl. 6 igs 1-4. Ma e ial examined.— C4: (25.7.1991), 3 m, wi hou gono hecae. C7: (5.7.1991), 5 m, wi hou gono he- cae; (18.8.1991), 6 m, wi hou gono hecae; (27.1.1992), 8 m, wi h emale gono hecae. C9: (23.11.1991), 5 m, wi hou gono hecae. C10: (14.7.1993), 10-18 m, wi hou gono hecae. C12: (13.5.1986), 20-25 m, wi h male gono hecae. C13: (6.5.1986), wi hou gono hecae. C14: (5.5.1986), 20 m, wi hou gono hecae. C15: (10.5.1986), 25 m, wi hou gono hecae. F23: (13.7.1989), 30-32 m, wi hou gono hecae. F51: (19.7.1989), 27-28 m, wi hou gono hecae. F55: (20.7.1989), 38-42 m, wi hou gono hecae. F66: (27.7.1989), 25-28 m, wi hou gono hecae. A2: (10.10.1992), 21 m, wi hou gono hecae. A3: (10.10.1992), 23 m, wi h emale gono hecae (RMNH-Coel. 27841, slide 4024). A6: (11.10.1992), 84 m, male and emale gono hecae. A8: (11.10.1992), 112 m, wi hou gono hecae. A10: (13.10.1992), 41 m, wi hou gono hecae. A16: (20.10. 1992), 40 m, wi h emale gono hecae. A17: (21.10.1992), 147 m, wi h emale gono hecae. Desc ip ion.— Hyd ocauli e ec , s ongly polysiphonic o e g ea e pa o hei leng h, b anched, pe isa c b own o yellowish in younges pa s. Axis and hyd ocla- dia in he same plane, b anches i egula ly al e na e ( ig. 2a). Monosiphonic pa s o colony di ided in o in e nodes sepa a ed by oblique nodes poin ing al e na ely le and igh . P ima y hyd o hecae sessile, disposed la e ally on an apophysis a ex eme end o each in e node and in al e na e posi ion, poin ing le and igh ( ig. 2b). Hyd o hecae shallow, wi h almos s aigh walls; im no e e ed, adcauline sides pa ly adna e o close o in e node. Se e al hyd o hecae p o ided wi h one o mo e hyd opho es o a ied leng hs, o igina ing om diaph agm o p e ious hyd o heca ( ig. 2b). Male gono hecae o oid, elonga ed ( ig. 2d). Female gono hecae g adually widen- ing owa ds la ened dis al end, p o ided wi h wo la e ally disposed hyd o hecae ( ig. 2c). Measu emen s (in mm).— Maximum heigh o he colony 180 Hyd ocladial in e nodes, leng h 0.33 - 0.64 diame e 0.13 - 0.19 ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 33 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)34 Fig. 2. Halecium halecinum (Linnaeus, 1758), a-c, om Chipiona (A3); d, om Pun a Almina (C1). a, shape o colony; b, pa o hyd ocladium; c, emale gono heca; d, male gono heca. Scales: a, 2 cm; b-d, 0.5 mm. a d b c ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 34 35Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) Hyd o hecae, leng h 0.02 - 0.05 diame e a im 0.13 - 0.18 Male gono hecae, leng h 0.70 - 1.05 diame e 0.29 - 0.375 Female gono hecae, leng h 0.75 - 1.00 diame e 0.425 - 0.550 Subs a e.— The ma e ial s udied was g owing on s ones, ba nacles and ubes o polychae es. I has been also eco ded om shells (Co nelius, 1975), seaweeds and o he hyd oids (Boe o & F esi, 1986). Discussion.— Halecium halecinum and H. beanii (Johns on, 1838) a e simila spe- cies. They mainly di e in mo phology o he emale gono hecae (Milla d, 1975; Roca Ma inez, 1986; Ál a ez, 1993). Howe e , adul colonies o H. halecinum a e egula ly pinna e while hose o H. beanii a e no , making a bushy appea ence (Ál a ez, 1993; Ve oo , 1946). In he p esen ma e ial all colonies, wi h o wi hou emale gono he- cae, ha e he same pinna e aspec and consequen ly ha e been conside ed o ep e- sen H. halecinum. Gili e al. (1989) poin ed ou ha , in H. beanii, he hyd opho es a e dis inc ly sepa a ed om he wall o he axis, in con adis inc ion o he condi ion obse ed in H. halecinum. Dis ibu ion.— Cosmopoli an (see Ál a ez, 1993). Dep h eco ds a e om be ween 0.5 m (Boe o & F esi, 1986) and 310 m (Naumo , 1960); occasionally ound much deepe (Medel & Ve oo , in p ep.). Halecium liou illei Billa d, 1934 ( ig. 3) Halecium Liou illei Billa d, 1934: 227, ig. 1; Leloup, 1937: 94, 116, ig. 2; Ramil Blanco & Pulpei o, 1991: 63-68, igs 1-3. Ma e ial examined.— C2: (11.2.1990), 0 m, wi h emale gono hecae. C12: (4.5.1986), (14.5.1986), 20-25 m, wi hou gono hecae. C13: (6.5.1986), 38 m, wi hou gono hecae (RMNH-Coel. 27844). C14: (5.5.1986), 20 m, wi hou gono hecae. Desc ip ion.— Hyd ocauli e ec , b anched o unb anched, some annula ions a basal pa o axis may be p esen ( ig. 3a). In e nodes wi h hick pe isa c, sepa a ed by oblique nodes, poin ing al e na ely igh and le , no in s aigh line bu gi ing axis a zig-zag appea ance ( ig. 3b). P ima y hyd o hecae disposed la e ally a he end o each in e node, al e na ely di ec ed le and igh , sessile, deep, wi h dis inc dia- ph agm, and g adually widening om base onwa d; im wide, occasionally e e ed bu no ecu ed. Adcauline side o hyd o heca longe han abcauline side, se o om in e node. Ring o desmocy es p esen inside hyd o heca. Se e al hyd o hecae may o igina e om diaph agm o p e ious hyd o heca, bu seconda ies usually wi h longe , annulla ed hyd opho e ( ig. 3c). Female gono hecae only ha e so a been obse ed, o igina ing om inside o hyd o hecae. They ha e a na ow annula ed pedicel and g adually widen owa ds hei end. Supe io pa is enla ged on one side, while on o he side wo closely adna e hyd o hecae a e p esen ( ig. 3d). ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 35 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)36 Measu emen s (in mm).— Maximum heigh o colonies 34 In e nodes o hyd ocaulus, leng h 0.38 - 0.90 diame e 0.13 - 0.24 In e nodes o hyd ocladium, leng h 0.53 - 0.70 diame e 0.12 - 0.17 Hyd o heca, leng h 0.03 - 0.08 diame e a he diaph agm 0.12 - 0.14 diame e a he im 0.19 - 0.22 Gono heca, leng h 0.95 - 1.00 diame e 0.50 - 0.53 Fig. 3. Halecium liou illei Billa d, 1934, om Caños de Meca (C2). a, shape o colony; b, pa o hyd o- cladium; c, p ima y and seconda y hyd o hecae; d, emale gono heca. Scales: a, 1 cm; b, d, 0.5 mm; c, 0.1 mm. d b c a ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 36 37Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) Subs a e.— The colonies we e ound g owing on ocks, associa ed wi h Tubula ia ce a ogyne C. Pé èz, 1920, and on B yozoa. The species has been ci ed also om sandy bo oms, g owing on Eudend ium spec., polychae e ubes, shells and ascidians (Ramil & Iglesias, 1988); also on Go gonacea (Al una, 1994b). Dis ibu ion.— A lan ic coas s o Mo occo (Billa d, 1934; Leloup, 1937) and Ibe - ian Peninsula (Ramil & Iglesias, 1988; Al una, 1994b). The dep h eco ds a e be ween 0 and 60 m (Al una, 1994b). Va iabili y.— Colonies ound a 0 m dep h we e monosiphonic and sca cely b anched, bu hose om deepe wa e we e polysiphonic a hei base and had well de eloped b anches. Discussion.— Al una (1994b) desc ibed he gono hecae o H. liou illei o he i s ime om Eu opean wa e s. They a e also p esen in ou ma e ial bu show sligh di - e ences wi h hose dec ibed by Al una (1994b) and Leloup (1937). Acco ding o he desc ip ions o hese au ho s he gono hecae a e la a hei dis al end, whe e hey show a single hyd o heca, while in he p esen ma e ial hey a e dis ally enla ged on one side and ha e wo hyd o hecae. New e ile ma e ial would be needed o esol e his con o e sy. Halecium pusillum (M. Sa s, 1857) ( ig. 4) Eudend ium pusillum M. Sa s, 1857: 154, pl. 1 igs 14-16. Halecium pusillum; Mo z-Kossowska, 1911: 347, igs 13c, 15-16. Halecium schneide i Bonne ie, 1898: 3, 10, pl. 1 igs 10-10b. Halecium annula um S echow, 1919: 33. Halecium co uga issimum; Pa i i, 1970: 25-26, ig. 24 (no Halecium co uga issimum T ebilcock, 1928. Ma e ial.— C13: (6.5.1986), 38 m, wi hou gono hecae. C14: (5.5.1986), 20 m, wi hou gono hecae. Desc ip ion.— Hyd ocauli e ec , wi h hin pe isa c and annula ed in e nodes. Hyd o hecae si ua ed a end o each in e node; b anches o igina ing la e ally, below hyd o hecae, cu ed a hei base ( ig. 4a). Hyd o hecae shallow, diaph agm dis inc , walls s aigh a basal hal and cu ed ou wa d a dis al pa ( ig. 4c); desmocy es p esen . Apical s olons a end o axis a e equen ly p esen ( ig. 4b). No gono hecae occu on colonies examined. Measu emen s (in mm).— Maximum heigh o he colony 1.2 Hyd o hecae, leng h 0.03 diame e 0.12 - 0.17 Diame e o hyd opho es 0.07 - 0.1 Subs a e.— The ma e ial s udied was ound g owing on seaweeds and on Euden- d ium spec. Ramil (1988) ound i on ocky and sandy bo oms wi h shells, on B yo- zoa, shells, ascidians, and on Obelia biden a a Cla k, 1875; Al una (1994a) on ba nacles, and Boe o & F esi (1986) on sea-g ass. Dis ibu ion.— Halecium pusillum is widely dis ibu ed in he Medi e anean ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 37 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)44 Fig. 7. Hyd odend on mi abile (Hincks, 1866), om El Po il (C1). a, colonies a ached o subs a e; b, shape o indi idual colony; c, s olon de eloping om hyd o heca; d, in e nodes wi h hyd o- and nema oheca; e, nema o heca. Scales: a, 1 cm; b, 5 mm; c, e, 0.5 mm; d, 0.2 mm. a b c e d ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 44 45Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) Discussion.— Se e al au ho s (Ve oo , 1959; Co nelius, 1979b; Medel & López González, 1996) ha e placed his species in he genus Ophiodissa S echow, 1919, because o he p esence o nema o hecae, ese ing he genus Hyd odend on Hincks, 1874, o he unique species wi hou nema o hecae, H. go gonoide (G.O. Sa s, 1874). Howe e , Rees & Ve oo (1987) poin ed ou ha Bonne ie (1898) epo ed nema- o hecae in he ype ma e ial o ha species. Acco ding o Co nelius (1995), he de el- opmen o nema o hecae in his g oup o species has ecei ed mo e axonomical a en ion ha i eally dese es and he consequen ly conside s he wo gene a synon- ymous. Howe e , he does no ejec he u u e e-es ablishmen o he genus Ophiodis- sa (S echow, 1919), as i s ype species is qui e di e en . We accep he synonymy o he wo gene a, because p esence o absence o nema o hecae is no a eliable cha ac- e . Fo he es ablishmen o gene a in he amily Haleciidae o he cha ac e s and p e e ably a combina ion o cha ac e s should be used. He e, we ollow Co nelius (1975) in conside ing Ophiodissa cacini o mis (Ri chie, 1907) an in alid species as i is p incipally based on he heigh o he colony. Al hough o he au ho s la e on a ibu ed addi ional speci ic cha ac e s, all o hose we e also obse ed by Co nelius (1975) in H. mi abile. This au ho poin ed ou he comple e simila i y in cha ac e es o he ype ma e ial o bo h species. Zoogeog aphical ema ks on he Haleciids om he Ibe ian Peninsula The amily Haleciidae o he Ibe ian Peninsula is ep esen ed by 16 species (see Medel & López González, 1996). O hese species 5 a e cosmopoli an [Halecium beanii, H. delica ulum, H. halecinum, H. sessile No man, 1867; and H. enellum], 4 a e opical- A lan ic [Halecium lankes e i (Bou ne, 1890), H. nanum Alde , 1859, H. pusillum and Mi ocomium ci a um Haeckel, 1879], 3 a e bo eal [Halecium lab osum Alde , 1859, H. mu ica um (Ellis & Solande , 1786) and Hyd an hea ma ga ica (Hincks, 1863)], 2 a e es ic ed o he A lan ic a ea nea o he Ibe ian Peninsula (Halecium liou illei and H. sibogae ma ocanum), 1 is A lan o-Medi e anean (H. pe osum S echow, 1919), and 1 is ci cum opical (Hyd odend on mi abile). Thus one- hi d o he haleciid auna o he Ibe ian Peninsula has a wide zoogeog aphical dis ibu ion (cosmopoli an and ci cum- opical) while wo- hi ds a e es ic ed o he A lan ic and he Medi e anean. Compa ison o he lis o he Ibe ian Haleciidae wi h ha published by Boe o & Bouillon (1993) o he Medi e anean (17 species), shows ha , wi h he excep ion o Halecium banyulense Mo z-Kossowska, 1911, H. conicum S echow, 1919, and Hyd an- hea aloysii (Zoja, 1893), he emaining Medi e anean species a e also ci ed om he Ibe ian coas s. The h ee lis ed abo e a e he only species conside ed endemic o he Medi e anean, bu hei absence om he Medi e anean coas o he Ibe ian Penin- sula is ema kable. On he o he hand, H. liou illei, conside ed an A lan ic species, has been ound in he S ai o Gib al a (Medel & López González, 1996; p esen wo k), so i s p esence in he Medi e anean could be expec ed. The same applies o H. sibogae ma ocanum, conside ed o be an A lan ic species, bu i s p esence in he S ai o Gib al a (Ramil & Ve oo , 1992; p esen wo k) and in he Albo an Sea (Ramil & Ve oo , 1992), show i o ha e an A lan o-Medi e anean dis ibu ion. Finally, he B i ish auna o Haleciidae (12 species a e enume a ed by Co nelius, 1995) also shows a high deg ee o simila i y wi h he Ibe ian auna. Only wo species ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 45 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)46 [Halecium undula um Billa d, 1922 and Hyd odend on go gonoide (G.O. Sa s, 1874)], bo h conside ed Bo eal species, a e no ound a he Ibe ian Peninsula. The se en spe- cies lis ed by Pa i i (1970) o he A lan ic coas s o Mo occo, ha e also been eco ded om he Ibe ian Peninsula. Key o he species o Haleciidae om he Ibe ian Peninsula 1 Hyd an h p o ided wi h in e en acula web ............................................................. 2 - Hyd an h wi hou in e en acula web. Fixed spo osacs .......................................... 3 2 Colonies eleasing ee medusae ................................................ Genus Mi ocomium One species: Mi ocomium medusi e um - Colonies eleasing ee medusoids ................................................ Genus Hyd an hea One species: Hyd an hea ma ga ica 3 Nema o hecae p esen ................................................................. Genus Hyd odend on One species: Hyd odend on mi abile - Nema ho ecae absen ........................................................................ Genus Halecium 4 4 Hyd o hecae ecu ed a he im ................................................................................... 5 - Hyd o hecae no ecu ed a he im .......................................................................... 10 5 Comma-shaped pe isa c old p esen a he base o hyd opho es .............................. .................................................................................. Halecium sibogae a . ma ocanum - Hyd opho es wi hou his cha ac e ............................................................................. 6 6 In e nodes smoo h ........................................................................................................... 7 - In e nodes wi h annula ions .......................................................................................... 9 7 Nodes oblique ................................................................................................................... 8 - Nodes ans e se .............................................................................. Halecium enellum 8 P ima y hyd o heca sepa a ed om he apophysis by a node, gono hecae wi h ows o spines ............................................................................... Halecium mu ica um - Wi hou hese cha ac e s. Hyd o hecae wi h an abcauline hickening ....................... ........................................................................................................ Halecium delica ulum 9 Monosiphonic colonies wi h annula ed in e nodes; gono hecae co uga ed and wi hou e minal ape u e ............................................................... Halecium pusillum - Polysiphonic colonies wi h i egula ly undula ed in e nodes; emale gono hecae no co uga ed and wi h e minal ape u e ................................ Halecium lab osum 10 In e nodes o a ied leng h and wi h undula ions ................................................... 11 - In e nodes o equal leng h ............................................................................................ 12 11 Nodes dis inc ; emale gono hecae no co uga ed, no wi h hyd o heca opening dis ally ............................................................................................. Halecium lankes e i ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 46 47Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) - No dis inc nodes, se e al undula ions p esen ; emale gono hecae co uga ed, wi h wo hyd o hecae opening dis ally............................................. Halecium nanum 12 Hyd o hecae sessile ....................................................................................................... 13 - Hyd o hecae wi h a sho hyd opho e ........................................ Halecium pe osum 13 Hyd o hecae shallow ..................................................................................................... 14 - Hyd o hecae deep, in e nodes wi h a hick pe isa c ................. Halecium liou illei 14 Hyd o hecal eno a ions sessile, wi hou hyd opho e .................. Halecium sessile - Hyd o hecae eno a ions p o ided wi h dis inc hyd opho e ............................... 15 15 Colonies pinna e, s ems e ec , mo e o less pa allel; emale gono hecae wi h pai o hyd o hecae opening dis ally .................................................. Halecium halecinum - Colonies bushy, no egula ly pinna e, b anches lexuosus; emale gono hecae kidney-shaped; hyd o hecae opening in cen e o conca i y ......... Halecium beanii Re e ences Alde , J., 1859. Desc ip ions o h ee new species o se ula ian zoophy es.— Ann. Mag. na . His . (3)3(17): 353-356, pls 12-14. Al una P ados, A., 1994a. Es udio aunís ico, ecológico y biogeog á ico de los cnida ios ben ónicos de la cos a Vasca: 1-769, igs, annex I wi h pls 1-19, annex II-VII.— Ph. D. hesis. Uni e sidad de Na a a, Pamplona Al una (P ados), A., 1994b. No as sob e los cnida ios ben ónicos de la cos a Vasca I.-Mi ocomium ci - a um Haeckel, 1879 y Halecium liou illei Billa d, 1934 (Cnida ia: Hyd ozoa).— Kobie, 21: 43-54, igs 1-4. Ál a ez Claudio, C., 1993. Hid ozoos ben ónicos y ca álogo de an ozoos de la pla a o ma y alud con- inen ales de la cos a cen al de As u ias: 1-458, igs 1-75, pls 1-33, abs 1-3.— Ph. D. Thesis. Uni- e sidad de O iedo. Billa d, A., 1922. No e su deux espèces d’hyd oïdes du li o al d’Os ende (Obelia spinulosa Bale, Hale- cium undula um sp. no .).— Annls Soc. . zool. malacol. Belg., 52(1921): 135-139. Billa d, A., 1929. No e su un gen e nou eau e quelques espèces nou elles d’Halecidae.— Bull. Soc. zool. F ., 54: 305-307, ig. 1. Billa d, A., 1934. No e su quelques hyd oïdes du Ma oc.— Bull. Soc. zool. F ., 59: 227-231, 468. Boe o, F., 1981. Sys ema ics and ecology o he hyd oid popula ion o wo Posidonia oceanica mead- ows.— Pubbl. S az. zool. Napoli I, Ma . Ecol., 2(3): 181-197, igs 1-13, abs 1-4. Boe o, F. & J. Bouillon, 1993. Zoogeog aphy and li e cycle pa e ns o Medi e anean hyd omedusae (Cnida ia).— Biol. J. Linn. Soc., 48(3): 239-266, igs 1-5, abs 1-2, appendix. Boe o, F. & E. F esi, 1986. Zona ion and e olu ion o a ocky bo om hyd oid communi y.— Pubbl. S az. zool. Napoli I, Ma . Ecol., 7(2): 123-150, igs 1-10. Bonne ie, K., 1898. Neue no wegische Hyd oiden.— Be gens Mus. Aa b., l898(5): 1-16, pls 1-2. Bouillon, J., 1985. Essai de classi ica ion des Hyd opolypes - Hyd oméduses (Hyd ozoa-Cnida ia).— Indo-Malayan Zool., 2(1): 29-243, abs 1-32. Bou ne, G.C., 1890. No es on he hyd oids o Plymou h.— J. ma . biol. Ass. U.K., n. se . 1(4): 391-398, pl. 26. Cla k, S.F., 1875. Desc ip ions o new and a e species o hyd oids om he New England coas s.— T ans. Conn. Acad. A s Sci., 3: 58-66, pls 9-10. Co nelius, P.F.S., 1975. A e ision o he species o La oeidae and Haleciidae (Coelen e a a: Hyd oida) eco ded om B i ain and nea by seas.— Bull. B . Mus. na . His ., Zool. 28(8): 373-426, igs 1-14, abs 1-12. ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 47 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)48 Co nelius. P.F.S., 1995. No h-Wes Eu opean Theca e Hyd oids and hei Medusae. Pa 1. In oduc- ion, Laodiceidae o Haleciidae. In: R.S.K. Ba nes & J.H. C o he s, eds, Synopses o he B i ish Fauna (New Se ies), 50: i- ii, 1-347, igs 1-73. Cough ey, M., 1876. C i ical no es on he New Zealand Hyd oida, subo de Thecapho a.— Ann. Mag. na . His ., (4)17: 22-32, pl. 3. Ellis, J. & D. Solande , 1786. The na u al his o y o many cu ious and uncommon zoophy es, collec ed om a ious pa s o he globe by he la e John Ellis, Esq. F.R.S. Soc., Reg. Upsal. Soc., e c.: i-xii, 1-206, pls. 1-63.— London, Benjamin Whi e and Pe e Elmsly. Ga cía Co ales, P., A. Agui e Inchau be & D. González Mo a, 1978. Con ibución al conocimien o de los hid ozoos de las cos as españolas. Pa e I : Halécidos, campanulá idos y plumulá idos.— Bol. Ins . esp. Oceanog ., 4(253) : 5-73, igs 1-32. Gibbons, M.J. & J.S. Ryland, 1989. In e idal and shallow wa e hyd oids om Fiji. I. A heca a o Se - ula iidae.— Mems Qd Mus., 27(2): 377-432, igs 1-41. Gili, J.-M., 1986. Es udio sis ema ico y aunis ico de los Cnida ios de la cos a ca alana: 1-565, igs, pls.— Ph. D. Thesis, Uni e si y o Ba celona. Gili, J.-M., J. Mu illo & J.D. Ros, 1989. The dis ibu ion pa e n o ben hic Cnida ians in he Wes e n Medi e anean.— Scien ia Ma ., 53(1): 19-35, igs 1-5. Gili, J.-M., W. Ve oo & F. Pagés, 1989. Hyd oids om he Wes A ican coas : Guinea Bissau, Namibia and Sou h A ica.— Scien ia Ma ina, 53(1): 67-112, igs 1-33. Gmelin, J.F., 1791. Linnaeus, C., Sys ema na u ae. Thi een h edi ion, edi ed by J.F. Gmelin. Vol. 1, pa 6 (Ve mes): 3021-3910.— Lipsiae, G. E. Bee . Haeckel, E., 1879. Das Sys em de Medusen. E s e Theil eine Monog aphie de Medusen [C aspedo- ae].— Denksch . med. na u w. Ges. Jena, 1: 1- 360; A las 20 pls. Hincks, Th., 1861. A ca alogue o he zoophy es o Sou h De on and Sou h Co nwall.— Ann. Mag. na . His ., (3)8: 152-161; 251-262; 290-297; pls 6-8. Hincks, T., 1863. On some new B i ish hyd oids.— Ann. Mag na . His ., (3)11: 45-47. Hincks, T., 1866. On Ophiodes, a new genus o Hyd oida.— Ann. Mag. na . His ., (3)18: 421-423, pl. 14. Hincks, T., 1868. A his o y o he B i ish hyd oid zoophy es. Volume 1: i- lx iii + 1-338, on ispiece, igs 1-45; olume 2: pls 1-67.— London, John an Voo s . Hincks, T., 1874. No es on No wegian Hyd oida om deep wa e .— Ann. Mag. na . His ., (4)13: 125- 137. Johns on, G., 1838. A his o y o he B i ish zoophy es: i-xii, 1-341, igs 1-51, pls 1-44.— Liza s, Edin- bu gh. Lamou oux, J.V.F., 1816. His oi e des polypie s co alligènes lexibles ulgai emen nommés zoophy- es: i-xxxi , 1-560, pls 1-19.— Caen, Poisson. Lamou oux, J.V.F., 1821. Exposi ion mé hodique des gen es de l’o d e des polypie s, a ec leu desc ip ion e celle des p incipales espèces, e c.: i- iii, old-ou able, 1-115, pls 1-85.— Pa is, Agasse. Leloup, E., 1935. Hyd ai es calyp oblas iques des Indes Occiden ales. (Zoologische E gebnisse eine Reise nach Bonai e, Cu açao und A uba im Jah e 1930, No. 13).— Mém. Mus. . His . na . Belg., (2)2: 1-73, igs 1-32. Leloup, E., 1937. Hyd oidea, Siphonopho a, Ce ian ha ia. I.- Hyd opolypes. In: Résul a s scien i iques des c oisiè es du na i e-école belge “Me ca o ”, ol. 1 p . i.— Mém. Mus. His . na . Belg., (2)9: 91-121, igs 1-16. Leloup, E., 1940. Hyd opolypes p o enan des c oisiè es du P ince Albe Ie de Monaco.— Rés. Camp. scien . Albe Ie de Monaco, 104: 1-38, pl. 1. Linnaeus, C., 1758. Sys ema na u ae pe egna ia na u ae, secundum classes, o dines, gene a, species cum cha ac e ibus, di e en iis, synonymis, locis. Edi io decima, e o ma a: 1-823.— Holmiae (S ock holm), L. Sal ii. Medel (So e as), M.D. & P.J. López-González, 1996. Upda ed ca alogue o hyd ozoans o he Ibe ian Peninsula and Balea ic Islands, wi h ema ks on zoogeog aphy and a ini ies. In: S. Pi aino, F. Boe o, J. Bouillon, P.F.S. Co nelius & J.M. Gili, eds, Ad ances in hyd ozoan biology.— Scien ia Ma ina, 60(1): 183-209, igs 1-2, abs 1-3. ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 48 49Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998) Medel (So e as), M.D. & W. Ve oo , 1995. Plumua ian hyd oids (Cnida ia: Hyd ozoa) om he S ai o Gib al a and nea by a eas.— Zool. Ve h., Leiden 300: 1-72, igs 1-28. Medel (So e as), M.D. & W. Ve oo , 1998. A lan ic Thy oscyphidae and Se ula iidae (Hyd ozoa, Cnida ia) collec ed du ing he CANCAP and Mau i ania-II expedi ions o he Na ional Museum o Na u al His o y, Leiden, The Ne he lands.— Zool. Ve h., Leiden 320: 1-85, igs 1-24. Medel (So e as), M.D. & W. Ve oo , in p epa a ion. Al an ic Haleciidae and Campanula iidae (Cni- da ia: Hyd ozoa) collec ed du ing he CANCAP and Mau i ania-II expedi ions o he Na ional Museum o Na u al His o y, Leiden, The Ne he lands.— Zool. Ve h., Leiden. Milla d, N.A.H., 1957. The Hyd ozoa o False Bay, Sou h A ica.— Ann. S. A . Mus., 43(4): 173-243, igs 1-15. Milla d, N.A.H., 1975. Monog aph on he Hyd oida o sou he n A ica.— Ann. S. A . Mus., 68: 1-513, colou pla e, igs 1-143. Mo z-Kossowska, S., 1911. Con ibu ion à la connaissance des hyd ai es de la Médi e anée occiden- ale. II.- Hyd ai es calyp oblas iques.— A chs Zool. exp. gén., (5)6(10) (= 46): 325-352, igs 1-16, pl. 18. No man, A.M., 1867. Repo o he Commi ee appoin ed o he pu pose o explo ing he coas s o he Heb ides by means o he d edge. P. II. On he C us acea, Echinode ma a, Polyzoa, Ac ino- zoa, and Hyd ozoa.— Rep. B . Ass. Ad m Sci., 36, (No ingham, 1866): 193-206. Nu ing, C.C., 1899. Hyd oida om Alaska and Puge Sound.— P oc. U. S. Na n Mus., 21: 741-753, pls 62-64. Nu ing, C.C., 1901b. [Le e o he edi o ].— Am. Na ., 35: 789. Oken, L., 1815. Okens Leh buch de Na u geschich e. D i e Theil. Zoologie, 1: 1-842, i-x iii, i-xx iii (sic).— Jena, Schmid. Pa i i, G., 1970. Ca alogue des cnidai es e c énai es des cô es A lan iques ma ocaines.— T a . Ins . scien . Ché i ien, Zool., 35: 1-149, igs 1-172. Peña Can e o, A.L., 1995. Hid ozoos ben ónicos de las Islas Cha a inas: aunís ica, ecología, biocenología y biogeog a ía: 1-589, pls 1-61, abs.— Ph. D. Thesis. Uni e si a de València Pé èz, C., 1920. No es su la aune ma ine du Boulonnais II. Tubula ia ce a ogyne n. sp.— Bull. Soc. zool. F ., 45: 171-177, igs 1-4. Philbe , M., 1935. Les hyd ai es de la égion malouine.— Bull. Ins . océanog . Monaco, 673: 1-36, igs 1-6. Ramil Blanco, F .J., 1988. Hid ozoos de Galicia: 1-525, pls 1-22.— Ph. D. Thesis. Uni e sidad de San- iago. Ramil Blanco, F. & A. Iglesias Diaz, 1988. La amilia Haleciidae (Cnida ia, Hyd ozoa) en las cos as de Galicia.— Thalassas, 6: 71-78, igs 1-8. Ramil Blanco, F. & E. Fe nández Pulpei o, 1991. Su la p ésence d’Halecium liou illei Billa d, 1934 (Cni- da ia, Hyd ozoa, Haleciidae) su les cô es eu opéennes.— Annls Ins . océanog ., n. se . 67(1): 63- 68, igs 1-3. Ramil (Blanco), F. & W. Ve oo , 1992. Repo on he Hyd oida collec ed by he “BALGIM” expedi- ion in and a ound he S ai o Gib al a .— Zool. Ve h., Leiden 277: 3-262, igs 1-68, abs 1-83. Rees, W.J. & W. Ve oo , 1987. Hyd oids om he John Mu ay Expedi ion o he Indian Ocean, wi h e iso y no es on Hyd odend on, Abie inella, C yp ola ia and Zygophylax (Cnida ia: Hyd ozoa).— Zool. Ve h., Leiden, 237: 1-209, igs 1-43, abs 1-37. Rees, W.J. & E. Whi e, 1966. New eco ds and auna lis o hyd oids om he Azo es.— Ann. Mag. na . His ., (13)9(100-102): 271-284. Ri chie, J., 1907. On collec ions o he Cape Ve de Islands ma ine auna, made by Cy il C ossland o S And ews Uni e si y, July o Sep embe 1904. The Hyd oids.— P oc. zool. Soc. Lond., l907: 488- 514, pls 23- 26. Roca Ma ínez, I., 1986. Es udio de los Cnida ios Ben ónicos de las aguas cos e as de Mallo ca: 1-32, igs 1-18.— Ph. D. Thesis. Uni e si a de les illes Balea s. Sa s, G.O., 1874. Bid ag il Kundskaben om No ges Hyd oide .— Fo h. VidenskSelsk. K is iania, 1873: 91-150, pls 2-6. ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 49 Medel e al. Haleciidae om he S ai o Gib al a and nea by a eas. Zool. Med. Leiden 72 (1998)50 Sa s, M., 1857. Bid ag il kundskaben om middelha e s Li o al-Fauna, Reisebemae kninge a I ali- en. I Classis: Polypi.— Ny Mag. Na u id., 9: 110-164, pls 1-2. S echow, E., 1919. Zu Kenn nis de Hyd oiden auna des Mi elmee es, Ame ikas und ande e Gebie e, nebs Angaben übe einige Ki chenpaue ’sche Typen on Plumula iden.— Zool. Jb., Sys . 42(1): 1-172, igs 1-56 (A-F2). T ebilcock, R.E. 1928. No es on New Zealand Hyd oida.— P oc. R. Soc. Vic ., n. se . 41(1): 1-31, pls 1- 7. Van P aë , M. 1979. Les ypes de polypes d’Hyd ai es conse és au Muséum Na ional d’His oi e Na- u elle de Pa is.— Bull. Mus. na n. His . na . Pa is, (4)1, sec ion A(4): 871-940, igs 1-113. Ve oo , W., 1946. Hyd ozoa (C 1) A. Hyd opolypen.— Fauna Nede l., 14: 1-336, igs 1-137. Ve oo , W., 1959. The Hyd oida o he opical wes coas o A ica.— A lan ide Repo . Scien . Res. Danish Exped. coas s op. W. A ., 1945-1946, 5: 211-325, igs 1-57. Ve oo , W., 1972. Hyd oids om he The a, Vema and Yelcho c uises o he Lamon -Dohe y geo- logical obse a o y.— Zool. Ve h., Leiden 120: 1-247, igs 1-83. Zoja, R., 1893. In o no ad un nuo o id oide.— Mi . zool. S a . Neapel, 10(4): 519-526, pl. 33. Recei ed:10.i .1998 Accep ed: 17.i .1998 Edi ed: J.C. den Ha og ZM 72-03 (Medel e al.) 05-01-2007 10:31 Page 50