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
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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).
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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
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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
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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
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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
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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).
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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
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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
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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
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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
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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
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