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