A REVIEW OF GERARDIIDAE (ANTHOZOA: ZOANTHARIA ... - redmic

QAG
FUNDACION
A REVIEW OF GERARDIIDAE (ANTHOZOA: ZOANTHARIA)
FROM THE MACARONESIAN ISLANDS AND THE
MEDITERRANEAN SEA WITH THE DESCRIPTION
OF A NEW SPECIES
O. Ocaña* & A. Brito**
* Instituto de Estudios Ceutíes (IECICECEL-CSIC), Paseo del Revellín n° 30. Apdo. 953, 51001.
Ceuta, orth Africa, Spain.
E-mail: lebruni@telefonica.net ieceulies I @retemail.es
**Departamento de Biología Animal (Ciencias Marinas), Universidad de La Laguna.
CI Astrofísica Francisco Sánchez sIn, 38206 La Laguna, Tenerife. Islas Canarias. Spain.
E-mail: abrito@ull.es
We describe the new species of the zoanthid Gerardia macaronesica exclusively
known from Macaronesian islands (Azores, Madeira, Canarias y Cabo Verde). Moreover,
we offer diagnostic data in order to distinguish properly G. macaronesica from G. savaglia
which is the other known species of the family Gerardiidae. The ecological requirements
and the nature of the association between G. macaronesica and the antipatharians are discussed.
A complete study of the cnidom measurements of both species is offered; in addition,
the cnidom images of G. savaglia are also showed and some biogeographycal considerations
are taking in account.
Key words: Zoanthids, Gerardiidae, Macaronesian islands and Mediten'anean, new
species.
Se describe una nueva especie de zoantídeo, Gerardia macaronesica, conocida
exclusivamente de las islas de Azores, Madeira, Canarias y Cabo Verde, y se dan las características
de diagnóstico más importantes para distinguirla de la otra especie conocida de
la familia, Gerardia savaglia. Además, se discute sobre las relaciones con otras posibles
especies de Gerardiidae que se reparten por diferentes zonas geográficas. También se aportan
datos ecológicos sobre ambas especies y se contempla el parasitismo como el tipo de
asociación más plausible entre G. macaronesica y los antipatarios que la hospedan. Por primera
vez se realiza un completo estudio de los nematocistos de ambas especies y la iconografía
de los nematocistos de G. savaglia. Algunas consideraciones biogeográficas son
expuestas.
Palabras clave: Zoantídeos, Gerardiidae, islas macaronésicas y Mediterráneo, nueva
especie.

Recent studies of Gerardiidae have been carried out by several auth .'
Canary Islands and others Macaranesian archipelagos. These authors recorde~rs I~
speCles, Gerardia savaglia (=Savalia savaglia~ from the Canaries, Madeira an a SIn
Verde (see BRITO, 1983 [1] Y 1985 [2]; OCANA el al., 1995 [25]). Unfortu d ea
f G 1 "· M' '" nately
presence o . savag La In acaraneslan waters was a mlsldentlfication motiva '
l k f d
'd 'f' . d' . . ted by .1._
ac o goo I entl IcatlOn escnptors m the Ilterature (especialIy concern" "ti:
savaglia)
Gerardia
and the similarities showed by both, G. savaglia and the present ne~g to
macaronesica. As we pointed out previously (OCAÑA el al., 1995 [25?e c1
many taxonomical problems of zoanthids are increased due to the absence of a . 1~4
d
.. f h . A' . . ppropn
escnptlOns o t ese specles. new specles, G. macaroneslca, IS described; this
ISWI "d esprea d' In thM' e acaroneslan IS . 1and s, from Azores to Cabo Verde being abs specl f
' enl ro
the Mediterranean.
G. savaglia is a well known species (although not welI described) through !be
Mediterranean Sea, which was recorded also from Madeira by JOHNSON (1899) [1
named as Gerardia lamarckii. The presence of a protein skeleton was the reason arguedby
Johnson to inelude it into the Antipatharian. CARLGREN (1895) [4] merged the speci
into the Parazoanthidae family. This peculiar anatomical item (skeleton axis), unknown UI
any other group of soft hexacorals, plays an important role in the biology of these speci
and also supports the Gerardiidae family to be separated from the others (see OCAÑA etal
1995) [25]. After this taxonomical papel', new material and information has been aehieved
on both Gerardiidae, the Mediterranean and the Macaronesian species. There are enougb
morphological similarities (colour, growth and size) between both species (G. savaglia and
G. macaronesica) to think that there is only a single one. However, the autoeeology and
some other characters
both species.
fram the anatomy and the cnidom make a elear differenee between
The cnidom is one of the main characters in order to distinguish the speeies of
anemones sensu lalo (see SCHMIDT, 1972 [30] & 1974 [31]; OCAÑA, 1994 [24]) and w
demonstrate in this papel' that it is also important for taxonomical purposes in zoanthids
Obviously, zoanthids needs more descriptive information in order to define properly !he
species and the other taxonomical categories. Sensu SCHMIDT (1974 [31]: 544) Gerardl
is merged into the recent Zoantharia in which are ineluded all the zoanthids with speeialbmastigophores
(spirulae) in the column wall: Parazoanlhus Haddon & Schackleton, 1891
Epizoanlhus Gray, 1867, Zoanlhus Lamarck, 1801 and Gerardia Lacaze-Duthiers, 1864
However, there is not descriptive information to assure that this particular cnidom eharacter
is present in alI the species above mentioned. One of the major prablems on zoanthids t~
onomy is the absence of complete deseriptions in order to recognize properly ~hespecl~
Despite some authors offer enough anatomical information in the specialized blbhograpth
(structure and placement of the sphincters and disposition of the mesenteries, among ~
ers), the cnidom characteristics were low considered and usualIy, not well valuatedRT
CARLGREN, 1937 [5] y 1938 [6]; PAX, 1952 [26]). A more recent paper (HERBE'
dOI1l
1972 [13]) takes in account some cnidom categories, however, stilI it is needed more en! di
information, images and measurements, together with the other characters, in arder to
tinguish accurately the zoanthids species. ri¡,ed
G. macaronesica is the second species which belongs to Gerardiidae and des~dYin
worldwide. We offer a complete description of this species and also a comparatlve 51
. . .h G macaronesica fram G. savaglia. In addition, some cnidom data of
disunguIS .
to.· described) fram the Indopacific and New Zealand are documented.
'les (no . .
r pe\; . d' (T to HÁUSSERMANN (2003) [12] the genus Savalta Nardo, 1844 has taxccor
.
In"
. over Gerardia Lacaze-DuthJers,
.
18
6
4. However, due to the wlde
.
use of
. -al pflofllY . f 1864 '11 d'd ...
1Tl1\; d' I in the recent hterature, rom ti now, we eCI e to maIntam 11.
S
nU
Gerar IC
ge "
'1lnensfrom the Can aries studied in the present work was collected by scuba
The spec . ..
sampling trips during ~h.eBen~os PraJect (Universidad de.La.Laguna, .1980.-1985) and
I IDg d'l'ons and scient¡f1c praJects, headed by local mstltutlOns (Universidad de La
oth r expe I I . .
. Museo de Ciencias Naturales de Santa Cruz de Tenenfe), between 1985 and 1994.
guna,. ale organizations which organize colIeting expeditions, as Farmamar Expedition
OtherpflV .,.
1990), also ha eontributed to co~lect mor~ speclmens of Gerardlldae along th~ .Canary
Ilands. The material from Madelra Archlpelago belongs to the Museu MunicIpal do
unchal(MMF) colIeetions. The material fram Cape Verde has been collected by scuba div-
IDgduring several expeditions supported by different ins~ituti?ns fram the Canaries (Museo
d Ciencias Naturales de Santa Cruz de Tenenfe; UniversIdad de Las Palmas de Gran
anaria; Gobierno de Canarias). The sample fram Azores was sent by Universidade dos
ores, Departamento de Oceanografia e Pescas (DOP).
The material of G. savaglia was colIected in two different western Mediterranean
I alities: Ceuta (Gibraltar Strait) and Banyuls (France). Other consulted specimens fram
th Indopaeifie and New Zealand has been facilitated by the Copenhagen Zoological
u eum (ZMUC) and the Auckland Museum (AK) in order to make comparison
pecies.
with our
The holotype of Gerardia macaronesica is deposited in the Museo de Ciencias
aturales de Santa Cruz de Tenerife (TFMC). The paratypes are deposited in the Museo de
HI toria Natural de Madrid (MNCN), Museu Municipal do Funchal, Madeira (MMF) and
ationaal Natuurhistorisch Museum, Leiden (RMNH). Some colonies and most of the histol~giealslides
are also deposited in the colIection of the Departamento de Biología Animal,
Diversidadde La Laguna, Tenerife (DZ ZG). Part of the material examined and also the
pe~anenl enidom slides are deposited temporalIy in the private collection of O. Ocaña and
11 wlll be senl to the apprapriate institutions, once the studies has been coneluded.
Part of the material (specialIy the material from Canary Islands) were anaesthetized
~
nh
re
menlhol
. .' .
el'
ys
t
a
1
s an
d'
preserved In 8% formaldehyde. As some colon les
"
come from dlfntlmslIlullOnsand
prajects, most of them are fixed directly in alcohol not relaxing pre-
10USy ¡he 1 T '
di ' . p? yps. he general morphology and anatomy were studied by means of a stereo
sectmg mlcro Th
aJ'al h Scope. e anatomy and histology were studied following the Ramón y
mel od for topo h' ..
I J) S· grap ICstamIng. Mallory triple stain was also used (see GABE, 1968
. peclmens of l .
edit. . co onles fram Canary Islands and Cape Verde and also from the
crranean w .
and exa' ere sectlOned for histological purposes (more than 500 slides in a whole)
Int rfere mmed and studied with a light micrascope equipped with Nomarski differential
nee Contrast .
nn gel lh Opllc system. Permanent slides of cnidom were prepared using glyc-
, e same tech . -
The el '. ~Ique prepared for meiofauna (see OCANA, 1994 [24]).
I-IM1DT(1;;slflcatlOn and terminology of nematocysts used is essentially after
2) [30], as adapted by den HARTOG el al. (1993) [11]. The surveys of the

cnidorn are surnrnarized in tables, in which the means and ranges of length and .
nematocysts are included. A large collection of underwater irnages has been usedW1dth
to show the color and shape variations, some aspects of its autoecology and als In o
variety of environrnent where the new species may occurs. o the h¡
Familia Gerardiidae Roche & Tixier-Durivault, 1951(=Savaliidae Nardo, 1844?)
Género Gerardia Lacaze-Duthiers, 1864 (=Savalia Nardo, 1844)
Savaglia larnarckí, JOHNSON, 1899 [15]: 814-815, short description, the species is iden
fied with Gerardia savaglia (=Savalia savaglia) and merged into the Antipatharian Wt
doubts; Madeira and the Mediterranean Sea.
Gerardia savaglia, BRITO, 1983 [1]:89-91, figs. 1-3, autoecology and distribution; folio
ing JOHNSON (1899) [15], the species is identified as G. savaglia; the Mediterranean,
north Morocco, Canary Islands and Madeira. BRITO, 1985 [2]:178-183, fig. 35, lam. 3
description, habitat and some ecological notes; colour images; Canary Islands, Madeira,
Mediterranean and north Morocco. PÉREZ y MORENO, 1990 [27]: 81, a repetition
some Brito's data, one colour image. OCAÑA et al., 1995 [25]:153-164, figs. 2-4, colour
images, complete study of the anatomy, histology and cnidom, ecological notes; Can~
Madeira and Cape Verde islands. WIRTZ, 1995 [32]: 52, exclusively a color image,
Holotype: (TFMCBMCN/000188), colony (13 cm high and 13,5 cm wide) witb
more than 1000 polyps growing on Tanacetípathes cavernicola Opresko (200la) branch
almost all the axis of T cavernícola covered by the zoanthid, some calcareous sea weedsand
barnacles were noticed on the branches too; O. Ocaña coll., 14-08-2002, Punta Preta.
Tarrafal, (Santiago, Cape Verde Islands), 20 m, scuba diving, 15°1TN 23°46'0. paraty~
(MNCN 2.04/1079), branch of the same colony (with 100-150 polyps and 13 cm high) w' H
some areas of the antipatharian not covered by the zoanthid; same data as holotype; (RM !he
Coe!. 32362), branch with 150-300 polyps and 23 cm high, with some areas of re
. . . l' fE' tOléquata we
antlpathanan not covered by the zoanthld, severa speclmens o UnLce . 0.200
noticed living on the colony; same data as holotype; (MMF35616), branch wlth 10
polyps and 10 cm high, calcareous debris noticed; same data as holotype,
Other materials
'O 24 [7.19 O
Canary Islands: Tenerife: (DZ ZG-I) Las Caletillas, 28°21'N 16°21 , -. - thell
, An/lpa
75 m, by entangled in the fishermen nets and traps, small colony growmg on 1981.100
wo/lastoní, G. Dionis leg.; (DZ ZG-2) Punta de Güímar, 28°l8'N 16°20'0, 14-03-
f
· h men nets and traps, small colony growing on Antipathella wollas-
. thc IS er
taJlgl~d~n c ; (TFMC CN/0151) Playa Santiago, 28°15'N 16°50'0, 1~~03-19.89, 25 m
. D1on,.sI g'd" small colony growing on Antipathella wo.llastolU In vertIcal wall,
'uba IVlllg,
1 ¡ d h) se ZG-3) Punta Teno, 28°20'N 16°55'0, 15-09-1990,40 m collected by
ana leg.; (OZ 1 nies growing on rock platforms and also on small specimens of
. . , sl11allco o - 3) O O - I O h
ba dI\1l1g.· . . colour images (see Ocana et al., 1995:16 , . cana ego t er
I / / lI'o//a.\/OIlI, . PI S J
palIe (/ d but not collected, on different areas of the Island: aya an uan
'ere observe , . . I 25
Imens \\ l owing on a dead axis of A. wollastoní placed In a vertIcal wal ,
. t 's) co ony gr
Glgan c.. Z ZG-4) Tazacorte, 28°21'N 16°21'0, 3-11-1990, 30 m collected by scuba
, - D!llrna:
~
(O
. s of a big colon y growmg
..
on vertlca
l
wa
II
,co
l'
or Images,
O
.
O
cana
-
, I11CspecIlnen.
Ing. ~il;m;: (OZ ZG-5) Punta d~ los Frailes, 27°3~'N 1.8°00'0, 3-?~-1982" 23 m by
EL l growing on Antlpathella wollastolU, Tallarte Expedltlon leg., (DZ ZGmall
co ony . . .
ge, s . F 'Ies 27038'N 18°00'0,07-1985,35-45 m, by scuba dlvlng, small portlon
Puntade los ral ., ..
lony groWI
'ng on Antipathella wollastoní at the cave gate, A. Bnto leg, Gran Canana:
a o 7 P rto de Sardina 28°08'N 15°42'0, 5-10-1983, 12 m, by scuba diving, small
DZ ZG· ) ue, .
. g on Antipathella wollastoní other colonies were observed growlng on walls,
lonygrow1n ' , 0' 0'
Brito leg.; (DZ ZG-8) Punta de la Sal (El ~abron).' 27 52 N 15 22 O, 6-0?-1990, 30 m,
cuba diving, portion of a big colony growmg o~ ItS own skeleton, F. .Esplno leg, Oth~r
cimens were observed, but not collected, on dlfferent areas of the Island: El Cabron
nnaga). 10-20 big colonies growing on its own skeleton placed on a rocky platform wlth
d (color images), 30-40 m, Lanzarote: (DZ ZG-9) Puerto del Carmen, 29°03'N ]3°32'0,
1 1O-191U,40 m, portion from a big colon y growing on A. wollastoni and placed at the
nuuncave gate, color image (see Brito, 1983 y 1985) T. Cruz leg.; (DZ ZG-IO) Puerto del
armen. 29°03'N 13°32'0,22-06-1990, 55 rn collected by scuba diving, portion from a
lony growing on A. wollastoní placed in a crevice in a vertical wall with L. pruvotí, O.
aña leg. Many other specirnens were observed but not collected around the Montaña
lara islet. Fuerteventura: (DZ ZG-ll) Frente a Casas de Jacomar, 28°16'N ]3°53'0,21-06-
1990.35 m collected by scuba diving, portion from a big colony growing on A. wollastoní
and placed in the middle of a small tunnel, O, Ocaña leg.
Madeira: (MMF25715) Cais do Porto Novo (south Madeira coast), N 32° 39' 629"
16°48' 390",13-04-1994,31 m collected by scuba diving, portion from a colony grow-
IOg on A. \\'ol/astoni on its own skeleton, P. Wirtz leg.; (MMF27120) Antigo Cais do Porto
ovo (south Madeira coast), N 32° 39' 779" W 16° 48' 224",
b scuba diving, P, Wirtz leg.
19-07-1997,47 m, collected
lores: (DOP) Formigas Islets, 37° 16' 08"N 24° 46' 48"W, 25-08-2002, 25 m colted
hy scuba diving, several portion of colonies growing on A. wollastoni, not well prerved
material, F. Tempera leg.
12 19 Cape Verde: Santiago: (DZ ZG-12) Caleta do Sao Martinho, 14°54N 23°34'0, 31-
.,. K7, 17 m, collected by scuba diving portion of a big colony growing on
lanU('('1 I '
'/Hitles cavernícola, crevice in a vertical wall with many specimens of Euníce
rquuta living o th I . . . ..
Palmas'l '. . n e co ony, A. Bnto leg.; (C-16), (Matenal from UllIversldad de Las
rowin ,ile prOJect to Cape Verde), no data, scuba diving collected, portion of a colony
g On r ca . l' .
v ral . .' verntco a, F. espmo leg, Other colollles were observed, but not collected:
r (u\,('r"tes/ of the Tarrafal Bay, between 20 and 35 m, in shadow habitats, associated with
nlCO a O O - I
and Cav" " '. cana eg; King fish, 15°16'N 23°46'0, 25-40 m, vertical walls, tunnels
"s, assoclat d . h
47'0, 25.35 ~ Wlt T cavernicola, O. Ocaña leg,; Faro de Tarrafal, 15°1TN
m, blg caves and tunnels with vertical walls, associated with T caverníco-

la, strong current environment, O. Ocaña leg. Sal: we observed colonies in Santa M .
16°35'N 22°54'0, 20-25 m, on vertical waJls and crevices, associated with Tarta
high sedimentation, O. Ocaña leg.; Palmeira Bay, 16°45'N 22°59'0
gate caves, associated to antipatharian, O. Ocaña lego
25 m ver·t·cavlernlc
, , Ica w 1I
a
Morphology: Several ways of growth and external appearances can occu' h
. r In t e
ent new specles. In the case that the colony creates its own skeleton, a basal plate is P
ated where the polyps are asexuaJly produced (Fig. 5e); after this the colon y Contin gen
erating many branches in a single plane or in several of them. Sometimes the spe ~esg
.. . c~
ge~erate a poor developed ~Ibbon hke skele~on, ha:dly notlced without an aCCurateob
vatlon of the colony. GroWIng up on an antlpathanan skeleton is the third way of
showed by this new species, the characteristic spines of those are some identificatio~T0har.
acters in order to distinguish the antipatharian skeleton where the G. rnacaronesica c
occurs. The colonies may reach one meter high.
In preserved material, the polyps measurements are variable between 0,2-1 cm hi
and 0,2-0,5 wide; due to the retraction stage the column can be cylindrical to flat mamm¡.
form; the texture should be roughly by the 90ncentration of sand in the ectoderm of
polyp walJ. In aJive and expanded conditions the sizes increase considerably in two OTthree
centimeters at least. Colonies with small polyps were frequently noticed in the material froII
Cape Verde. The tentac1es present medium size in expanded conditions; they are pointed.
42 in number, arranged in two, more or less, entacmeic cyc1es; we noticed 21 marginal t
The polyps are overhanging from coenenchyme, but extremely embedded polyp
noticed. Wide coenenchyme in the well developed areas with many polyps, fine and I
developed in the extreme areas of the colony.
The species present several colours, from yellow to orange in the tentacles, the
coenenchyme and the column (Fig. 5). As a consequence of the sediment accumulation and
other conditions, not well c1eared, the column can be brownish or whitish, or even pinki
in the column; the brownish column are typical from the colo nies growing on antipathan
ans, although not exclusively. Fixed specimens turned into red colour frequently allhou
orange or brownish are also possible.
Microanatomy and histology: The mesenteries present macrocnemic dispositio
although one specimen showed an intermediate arrangement; there are 28 mesenleries iothe
pharynx region and 42 in the upper column region. Most of the mesenteries are conoected
with a short, low folded pharynx; a single siphonoglyph, larger than wider, is noticed (FI
2d).
The sphincter is well developed and mostly endodermal, aJthough a final portioOo
.
It ISconsldered
. .
mesogleal (see OCANA
- . 5 b) The meseoler
el al, 1995) [25] (see Flgs. a y . an
ial musc1es are poorly developed, we failed in detecting retractor musc1es and weak ~ure
l
etobasilar musc1e only in some mesenteries. The tentacles present also a weak mus cu a
and it is arranged in short process. 1 pre5
The ectoderm present to 80 11mthick, being in the column poorly develo ped
i :iSu
ents glandular ceJls and b-mastigophores; there is afine cuticle that covers the who e '0 the
. 'd tegones, l
The tentacular ectoderm IS well developed and presents several cm om ca e cilia are
pharynx we noticed a few nematocysts, absence of glandular cel1s and larg
observed in the siphonoglyph.
. can reach to 120 Ilffi thick; it presents afine structure and more develm
'sog 1 oea . . .
')1le (;. I than in other anatorrucal areas. We notlced low cel1ular denslty, some
h' "0 umn
110 1 (; ~ d I content have been observed (Fig. 2c); sand incrustations are numerous
. h glan u ar
\\11 art of the column. The mesogloea is hardly noticed in some areas of the
. lhe outer p . .
aaUy10 h harynx wider in the slphonoglyph than In any other area of the pharynx.
nleries aod ld e . p can r~ach to 80 Ilffi thick, it is formed by absorption cells in the col-
Th' codo elm .
e l' dular cel1s among those; commonly, not folded and It presents a low
d some g an .
aO· Mt landular cells are observed along the mesentenes; a very low developlopmeol.
dOd s rgmis noticed in some mesenterical areas. There are filaments unilobuf
lhe en o e
010 '1 h I led (Figs 2 e and f) in the top of the mesenterial filaments. In the speciaod
In o u a· . .
"d we observed gonads development In all the macrocnemes (FIgs. 2 g and h).
o ludle, 'd ( . d l
oidom: A complete survey of t~~ C~l om mea~urements, categones an p acef
h 1alerial from different 10calJtJes IS presented In the table 1. We notlced one catnOolen
. .
of spirocysls and several cate~ones. of the other ~ematocysts eqUlpment: t~ree catery
of elliplical, some of them oVOld,splrulae (b-~astlgophores); one category ISthe sp.eal
pirulae only present in some groups of zoant~ld.s, as the presentogenus. T~ree pemcIlmasligophores)
categones: one category of ellJptlcal to some oVOldpemcIlII A; one cat-
~ of elliplical big penicilli E and other of rather small elliptical to ovoid penicilli E.
lb re are al50 lWOcategories of real, but not very conspicuous, homotrichs of elliptical
pe. a lhird liny category of nematocyst can be assume as a homotrich but it need confir-
1100.
Remarks.- The absence and the presence (or also the range of abundance) of some
adomcalegories found in the studied material of G. rnacaronesica, along the distribution
•should be explained as a part of the specific range of variation displayed by this taxon.
rtheless, lhe different origins of the studied material and, specially, the differences of
onservalion method used, should be also taking in account. Some tiny categories (as the
mall homolrichs fram the wall) may be easily overlooked by the observer; moreover, the
rang of abundance of these capsules categories, in the polyps from different colonies, does
nol have lo be underestimated. Other categories which are not very conspicuous, like the
medlUmsize homotrichs from the filament or pharynx, can be easily overJooked due to its
n morphology and placement. We offer (see Fig. 1) an ideal nematocysts equipment for
macarollesica, although, for identification matters we recommend to use the most common
categories, always present in all the specimens from different populations studied by
u ( e lable 1and Fig. 1).
bit 1: G('/'{¡rdia lila. . .
lJlary I lands COIoneSlca. Survey of the cmdom of five colonies from different localilies. A: El Hierro,
Igas hlcls ¡DZZG-6). B: Santiago, Cabo Verde (DZZG-12). C: Holotype (TFMCBMCN/OOO 188). D:
. lores (DOP n/n). E: Madeira (MMF25715)

-- Mean and range (in parentheses) Mean and range (in parentheses)
Nematocyst
Organ Origin
type of length and with (lf Nematocyst
N
of length and with of N Frequency
nematocyst capsules in /l
FreqUen~
orsall origin type
---nematocyst
capsules in /l
D Spirulae (c) 13,9 (13-18) x 4,9 (4-7,5) 13 Homotrichs ? (j)
8,9 x 1,1 1 Rare
ComInon A
E Spirulae (e) 17,4 (16-19) x 3,8 (3-4,5) 12
COInInon Spirulae (m)
19,2 (17,7-21) x 2,7 5 Rather common
A
E Spirulae (h) 17 x 2,5 ~rynl ---t- ¡.--
1 Rare
Soirulae (m)
18,5 (15,5 22,2) x 3 (2,2 3,3) 11 Common
B
E Spirulae (c) ¡.--
14,4 (13-16) x 5,3 (4,8-6) 18 ComInon
Spirulae (m) 16 (15-18) x 3 (2,5 3,8) 6 Rather-common
e
A 1---
Penicilli E (a) 24,2 (22,8-25,5) x 10 2
Uncomrnon
Soirulae (m) 16,9 (15-20,5) x 3 (2,5-3,5) 10 Common
O
A Penicilli E (b) 19,1 (18,9-19,4) x 7,1 (6,4-7,8) 2 Uncomrnon
Spirulae (m) 18,6 (15-21) x 3 (2,8-3) 7 Rather-common
E
B Penicilli E (a) 24,4 (23,3-26,6) x 10,4 (9,4-11,1) 5 Uncomrnon
Penicilli A (o) 14,4 x 4,4 1 Rare
A
t-
B PeniciLJi E (b) 13,6 (13,3-14,4) x 6,8 (6,1-7,2) 4 Uncommon
Penicilli A (o) 15,1 (13,3-16,7) x 3,7 (3,1-5,3) 5 Uncommon
B
C Penicilli E (a) 23 (21-25) x 9,3 (9-10) 4 Uncomrnon e Penicilli A (o) 15,1 (14-16) x 3,9 (3,5-4) 7 Rather-common
C Penicilli E (b) I1 (10-12) x 5,8 (5,5-6) 3 Uncornrnon E Penicilli A (o) 19 x 4,8 1 Uncommon
O Penicilli E (b) 15,8 (14-17,5) x 6,1 (5-8) 9 Uncornrnon A PeniciLJi E (q) 25 (22,2-27,8) x 10,5 (10-11,1) 2 Rare
E Penicilli E (b) 14,4 (11,5-17) x 5,3 (5-7,5) 13 Common A Penicilli E (p) 15,3 (13,3-16,7) x 6,8 (6,1-7,8) 15 Common
C Penicilli A (d) 14,5 (14-15) x 4,5 (4-5) 2 Rare B Penicilli E (q) 25,3 (31,1-27,8) x 9,8 (9,4-12,2) 13 Common
O Penicilli A (d) 16,5 x 4 1 Rare e PeniciLJi E (p) 13,4 (12-17) x 6,7 (5,5-8) 21 Very common
E Penicilli A (d) 19 x 4 1 Rare C Penicilli E (q) 24,3 (23-26) x 9,4 (8,5-10) 9 Rather-common
C Homotrichs (n) 9,5 (9-10) x 3,2 (2,8-3,5) 2 Rare O Penicilli E (o) 14,9 (13-17) x 5,6 (4-7) 18 Very common
D Homotrichs (f) 10,6 (10-11) x 3,3 (3-4) 6 Uncommon O Penicilli E (q) 26 x 8 I Rare
Tentacles A Spirocysts (i) (11,1-22,2) x (2,2-4,4) 10 Very common E Penicill i E (p) 13,5 (13-15) x 6,6 (5,5-7) 16 Common
B Spirocysts (i) (11,1-17,8) x (2,2-3,3) 10 Very common E Penicilli E (q) 26 (25-27) x 11,7 (11-12) 3 Rather-common
D Spirocysts (i) 16,7 (10-25) x 2,5 (2-3) 3 Very common B Homotrichs (n) 9,8 (7,8-12,2) x 1,3 (1,1-2,2) 11 Common
E Spirocysts (i) 20 (15-25) x 3 (2-4) 2 Very common C Homotrichs (n) 13,6 (12-15) x 2,7 (2,5-3) 5 Uncommon
A Spi rulae (g) 17,6 (14,4-20) x 3,9 (2,8-4,4) 20 Common O Homotrichs (n) 17,8 (17-19) x 3,5 (3-4) 6 Uncommon
B Spirulae (g) 16,6 (14,4-18,9) x 3,3 (2,2-4,4) 20 Common E Homotrichs ? (i) 9,5 (9-10)x2 2 Common
C Spirulae (g) 17,1 (16-19) x 3,9 (3,5-4,8) 26 Very common E Homotrichs (n) 14 x 2,8 1 Uncommon
C Spirulae (h) 16 (15-17) x 2,3 (2-2,5) 2 Uncommon FlIaments C Spirulae (r) 16,1 (15-17) x 3,5 (3-4) 7 Uncommon
D Soirulae (h) 16,5 (16-17) x 2,5 2 Rather-commCII A Penicilli A (v) 16,5 (14,4-18,9) x 4,9 (3,96,7) 22 Common
D Spirulae (g) 16,7 (15,5-18) x 3,9 (3,5-4,5) 8 Common B Penicilli A (v) 16,3 (14,4-18,9) x 4,9 (4,2-7,8) 21 Very common
E Spirulae (g) 17,5 (15-19) x 3,8 (3-4,5) 19 Very commo!!,.. C Penicilli A (v) 14,9 (13 16) x 4,2 (3,55) 15 Common
E Spirulae (h) 16,6 (12- 18) x 2,3 (1,8-3) 11 Rather-com~ O Penicilli A (v) 17,2 (15-20) x 4,3 (3,5-5) 12 Very common
A Penicilli A (k) 17,8 (14,4-20) x 4,7 (3,7 4,4) 3 Rare - E Penicilli A (v) 15,3 (14 19) x 4,5 (3,8-5) 12 Very common
B Penicilli A (k) 16,7 x 4,4 1 Rare - A Penicilli E (w) 24,2 (18,927,8) x 9,4 (6,7-10) 6 Uncommon
C Penicilli A (k) 14 (12 15) x 4 3 Uncomm~ A Penicilli E (u) 17,5 (13,3-18,9) x 7 (5,6-8,7) 30 Very common
D Penicilli A (k) 17 x 3,9 (3,8-4) 2 Uncomm~ B Penicilli E (w) 25 (24,4 25,5) x 11 (10,9 11, 1) 2 Rare
E Penicilli A (k) 17,8 (16-19) x 4,3 (3 5,5) 5 Uncom~ f..--B PeniciLJi E (u) 14,5 (12,2 16,7) x 5,9 (5,5-7,2) 20 Common
12 Comm~
C
A Penicilli E (1) 15 (13,3 16,7) x 6,7 (5,6-7,5)
Penicilli E (w)
Rare ___
21,9 (20 26) x 9,6 (8,5-10) 17 Common
I-----C
B Penicilli E (a) 23,9 (22,2-25,5) x 10 2
Penicilli E (u) 13,2 (11-15) x 5,8 (5 7) 17 Very common
1----0
B Penicilli E (1) 14,9 (13,3-16,7) x 6,5 (3,9-7,8) 25 Co~
co~11
Penicilli E (u) 15 (14-16) x 5,5 (5 6) r-----9
12 Very common
C Penicilli E (l) 13,3 (11-15) x 6,6 (6 7) 15 Com~
PeniciLJi E (w)
r------L
24,5 (24 25) x 11,8 (11,5 12) 2 Uncommon
D Penicilli E (1) 15 (12,5-17) x 5,9 (5-7) 25 Penicilli E (w) 25 x lO 1 Rare
16
Common
E
PeniciLJi E (u)
E Penicilli E (1) 13,1 (12-15) x 6,2 (5-7) 14,5 (13 18) x 6,4 (6 7,5) 16 Very common

Organ IOrigin I
Nematocyst
type
Mean and range (in parentheses)
of length and with of IN
nematocyst capsules in /l
Ecology
A Homolrichs (I)? 9,2 (7,8-10) x 1,7 (1,4-2) 3
B Homotrichs (I)? 9,6 (7,8-12,2) x 1,8 (1,1-2,8) 17
e Homolrichs (s) 12,3 (11-13) x 3,3 (2,5-4) 4
o Homotrichs (s) 15 x 2,8 1
o Homolrichs (I)? 8,3 (8-9) x 2 3
E Homolrichs (s) 13,4 (12-15) x 2,3 (2-2,5) 6
E Homolrichs (I)? 8,7 (8-10) x 2,1 (1,8-2,5) 6
Gerardia macaronesica is a typical species from circalittoral and deep infrali
bottoms that often occur on Antipathella wollastoni, forming a very interesting associatil
This association is very characteristic around the Macaronesian islands, except in
Verde, where the zoanthids occur mostly on the antipatharian Tanacetipathes cavernicola.
has been collected from 12 (only in some special habitats) to 100 meters deep, althoughv,
little is known about its abundance in deeper bottoms. G. macaronesica is able to show
eral ecological adaptation forms, but on infralitora! bottoms always is placed in dark
tats, as vertical walls, cave gates, ruffs and big crevices, and in other habitats with the Ii;
conditions diminished. Big colonies with their own skeleton have been noticed at some
cial environments, where they find the ecological conditions to do it (this on!y has
observed in scarce sites at Canary Islands). These environments seem to be exposed pl
with high water movement. This is a key factor in order to understand the ecological di
bution of this species. We find the most important concentration of G. macaronesica
ing on its own skeleton in flat rocky and moderate sandy bottoms (Fig. 8a), but also in VI
tical walls at the exposed islets (Fig. 8b), both habitats with high water movements. '
morphology of the colonies and branches orientation can be quite different from one col,
to another, being more related to water movements than any other factor. In the ex
islets in Canary Islands (Montaña Clara, norte de Lanzarote) G. macaronesica was obse.
together with the gorgonian Paramuricea grayii and a species of the genus Eunicel/a suD
be identified. In deep infralitoral and circalitora! bottoms, and with different environ ,
conditions, the zoanthid can also form rather neglected or not very conspicuou~ pOPula:..
growing on walls and crevices (also exclusively noticed at Canary Islands), wlth very
skeleton development (OCAÑA et al., 1995) [25], like fine axis hardly noticed froro
wall (Fig. 9b). In this habitat the polyp's density is lower than in the other, but the zoan
can cover a big piece of the substrate (Fig. 9a). . tatI
The species grow on antipatarian on many occasions; some parts of the anl1P~~
coenenchyme are damaged and destroyed by the zoanthid. It seems to be a case o it .
sltlsm, . . where the zoant h'd I IS . growmg . on th e antlpatanan .. coenenc hy, me covenng . skc
including alive parts inside the colenteron. The final result is to get the antipat~nanofSUI
ton as an axis where grow up. Still, have to be studied if there is some absorptJOnlauSi
stances that may bring some trophy benefits to the zoanthid, but it should be a ~nes"
option to take it in account. Having demonstrated the serious damages that G.maca
.. lharians, we think that the inquilinism (see MORTON, 1989) [21] can
lhe:anllp

G. macaronesica is exc1usively known from the Macaronesian archi I
deira, Canary Islands, Cape Verde and Azores). In Canary Islands is well kn pe a~os (
islands and some inlets. OWnIn a11
Gorgonia savaglia Bertoloni, 1819.
Gerardia lamarckii LACAZE-DUTHIERS, 1864 [16]: 169, genus description. LAC
DUTHIERS, 1865 [17]: 1-63, comparison with several speci~s of Antipatharians, no pl
LACAZE-DUTHIERS,
·
ay: d tae.
1882 [18]: 10-17, 1 planche, descnptlün of the parasitic Lauro
~
Gerardia savaglia, CARLGREN, 1895 [4]: 319-334, remarks the position into zoan
and placed the species into Parazoanthiídae. ROCHE ET TIXIER-DURIVAULT, 1951[
402-409, plate 1, external description, anatomy (mesenterial cyc1es), bioquimical chara
of the axis, merged into the Gerardiidae family. ROSSI, 1958 [29]: 1-7, figs. 1
Gerardiidae, some external characters, autoecology; Mediterranean (Egean Sea, Actri
Argelia, Genova Gulf). LAUBIER & THEODOR, 1967 [19]: 223-225, fig.l, Gerardii
some data of external anatomy and habitat, repetitíon of the Rossi's data, Banyuls -Sur-
HERBERTS, 1972 [13]: 72, 142 Y 144, cited only in the key, merged into Parazoanthii
Schmidt, 1974 [31]: 544, recent zoanthids. MANUEL, 1981 [20]: 78, merged
Parazoanthiidae. ZIBROWIUS, 1985 [33]: 349-352, on the associated species Balssia~
GILI et al., 1987 [9]: 13-17, short description, autoecology, comparative table, Catalo
coas!. CALVÍN, 1995 [3]: 307, short description and some data about the habitat, ca
image of a colony with possibly its own skeleton, the Mediterranean.
Savalia savaglia, HÁUSSERMANN, 2003 [12]: 505, 1 abb., short description, and col
picture, merged into the family Savaliidae Nardo, 1844.
Ceuta: (Priv. Col.) Ciclón de fuera, North Bay, 35°54'N 5°17'0, 10_05-1998.30
by scuba diving collecting, small piece of a big colony growing on its own skel elO ° tfi
Haussermann, 2003: 505 for color picture of this particular colony), rock and sand pla
near a vertical wall colonized by Paramuricea cLavata "forest", other small colon~;oS
observed growing on P cLavata, O. Ocaña leg., (Priv. Col.). Sarchal, South Bay, 11col
SOI7'O, 15-01-2004,30-35 m, by scuba diving collecting, some branches of a sm~ f(1l
growing on P cLavata, community of several gorgonians growing on a rocky plal ole~t,
specimen of the crustacean Balssia gasti living on the branches, copepodaoin ~ut~o~'E
Ocaña leg. France: (ZMUC, no collection number) Banyusl Sur Mer, 42 ?9 N p da
08-1968,40 m, by scuba diving collecting, small piece of a colony growlOg on .
F. Fabricius lego
. ferial ofller species examined
rtlllve ",a
Gt
,.
ran f(/
° I l'
"P' New Zealand: (AK 82153) Bay of Islands, Motuterakihi Te Pahi
., ., . .. '
174°05.8'E, 23 m, by scuba dlVlllg collectIng, smal1 colony possibly growd
i; lo~~nskeJeton, smooth rac~ dome. with san?-silt s.kirt, A. ~. Stephenson leg.
o 'fíc' (X-489 , ZMUC), zoanthld groWlllg on antlpathanan, medlUm size colony, no
p el.. (Z_9695/KAH-9901l59, ZMUC), zoanthid growing on its own skeleton or on
dala. :'Ig colony no more data; (TZ-61, ZMUC), zoanthid growing on its own skeleoacea.
" '
~ 00 gorgonacea, medium size colony collected by scuba diving.
"0 i.~(for a more accurate description see LACAZE-DUTHIERS, 1864 [16]; ROCHE
TI IER_DUR1VAULT,1951 [28]).
The species is known to occur grawing on the gorgonian Paramuricea cLavata or
o wilh ils own skeleton. When the species generates its own skeleton, as happen with G.
amnel'ic(/ above described, a basal plate is generated where the polyps are asexually
u ed unlillhe colony is generated with many branches in a single plane or in several of
m
In prescrved material, the size of the polyps varies between 0,1-0,6 cm high and 0,2m
wide. The column can be cylindrical to flat mammiform if the specimen is strangly
cled: ~exlur~roughly due to the co~centration of sand in the ectoderm of the polyp wall.
lum Slze pOll1tedtentac1es, to 27 III number, arranged in two, more or less, entacmeic
I : we nOliced between 13-18 marginal teeth (sometimes
ooticed propcrly).
those marginal teeth can not
Pol~ps are overhangi~g fram coenenchyme, but extremely embedded polyps are
d. W.ldccoenenchyme In the well developed
loped 111Ihe extreme areas of the colony.
areas with many polyps, fine and less
The species present several colours, fram yellow to orange in the tentacles
o nch~mc and column. We only detected pale yellow colonies (see Fig. 10). When fixed
fonnahne, Ihe colonies turn to red colour.
Internal "6 anatomy . and microanato my. . M acracnemlc. d' ISposltlün .. of the mesenteries
rer~: - -2 8m.esent~nes. Pharynx short and not specially folded a single siphonoglyph
an WIdcr ISnotlced (Fig. 4b).
The sphincler is e d d 1
pecimen' I . d n? ~rma and not well developed, it seems to be weak (at least in
11 coversshaelsfte )h
f andd¡ffIcult to be distinguished fram the rest of the endodermal mus-
Weakand
. ,
t
o
b
t e colum
n more or
l'
ess (FIg. 4a). Retractors and parietobasilar mus-
are arrang
no
d .
o served praperl
y.
Th
e
t
entac
I
es present also a weak musculature and
T e 111short process.
he ectoderm can reach 75 h' k .
111 the phary . f..IID t lC In the column and 100 f..Imthick in the tentab
nx we noltced a few b
o erved in th . h nematocysts, a sence of glandular cells and large cilia
k. present a re Slp onoglyph. The mesogloea of the column wall can reach to 125 f..Im
ar lile structure and di'
as. Wc notic d I more eve opment In the column than in other anatomib
rved: some sa
e
d
ow
.
cellular d
ensIty,
. b
ut
I
acunae wlth
.
glandular content are frequent-
Ode . n Incrustatio ..
rm. Thc meso l' ns were notlced In the outer part of the tissue c10se to the
Wld'
er In the siph
g oea IS hardly
no
f
Ice
d'
In some areas of the mesenteries and
'
the pharmeSogloeaof
the onloglYPhthan in any other area of the pharynx. In the inner face of
al ti co umn it has b .
Ssucand also big ho~ . een notlced a system of big lacunae like, with endootnchs
(Figs. 4d, e and f). These areas are common along the

column wall; the endoderm associated to them can reach to 120 J.lm thick. W,
observed unilobulated filaments. All the macrocnemes bear gonads in the s e Only
ied (Fig. 4c). pec1rnen
Cnidom: A complete survey of the cnidom (measurements catego'
. . . , ' nes a d
ment) of the matenal from the two Medlterranean 10calItIes studied is in Tabl n pl
Fig, 3), We noticed one category of spirocysts and several categories of the e
II
(see
. h . f 11' , oth cysts eqUlpment: t ree categones o e Iptlcal, so me of them ovoid
el'
.
ne
mastigophores); one category is the special spirulae only present in some g~Osplrulae
th'd I S as the present genus. T'wo pemcI '11' I ( p-mastlgophores) . categories: one cate ups of
tlca
. l
to some OVOI
'd'
pemcI
'11'A d . gorYofel
1 an one category of ellIptical big penicilli E es .
mon In
'h
t especia
' 1
areas o
f
t
h
e en
d
oderm of the wall (see description above),
peclally Co
Table n. Gerardia savaglia. Survey of the cnidom of Iwo colonies from different Mediterranean I .
Ceuta (North Africa) (Priv. Col.). B: Banyuls (France) (ZMUNI n/n) ocahu
Colurnn A
A
B
A
A
B
Tentacles A
B
A
A
B
B
A
B
A
Pharynx A
B
A
Filarnents A
Ecology
B
Nernatocyst
type
Spirulae (e)
S irulae (d)
S irulae (e)
Penieilli A (b)
Penieilli E (a)
Penieilli E (a)
S iroe sts (e)
S iroe sts (e)
S irulae (h)
S irulae ( )
S irulae (h)
S irulae (o)
Penieilli A (f)
Penieilli A (f)
Penieilli E
.)
Penieilli A (k)
Penieilli A (k)
Mean and range (in parentheses)
of length and with of N
nernatocyst capsules in 11
15,6 (13-20) x 12,4 (10-15)
16 x 2
13,1 (9-16) x 10,6 (9-12)
16,5 (13-20) x 4 (3-5)
18,5 (14-21) x 9,3 (8-11)
18,2 (16-20) x 9,1 (8,5-10)
20 (15-25) x 2,8 (2-3,5)
20 (15-25) x 2,8 (2-3,5)
18 (16-20) x 3 (2,5-4)
22 (20-27) x 4,8 (4-5,5)
17,8 (16-23) x 3,3 (2,8-4)
24,4 (22-27) x 4,5 (4-5)
19,2 (15-23) x 4,4 (4-5)
18 x 4
26 x 10
16,6 (13-20) x 3,2 (2,5-4)
14,2 (12-16) x 4,3 (3,5-5)
17 (15-19) x 4,8 (4-5,5)
l3,6 (12-15) x 4,5 (3,5-5)
14,2 (12-16) x 4,3 (3,5-5)
5 Uneommon
1 Rare
10 Uneommon
51
20
2 Rare
2
2
28
4 Uneommon
23
12
II Uncommon
1 Rare
1 Rare
37 Rather-co
11 Rather-com
2 Rare
14 Common 13 Comm on
G. savaglia is a typical species from circalittoral bottoms (see ZIBROWI.U~~:,9
, , d to PO/OII
[33]; GIL! et al., 1987 [9]; CALVIN, 1995 [3]) frequently found aSSOCIate . ¡"on WI
clavata, In the Strait of Gibraltar area the species commonly occurs in asSOC1aI
_ rsonal observation). Although other species of gorgonians has been
O
Ocana, pe .
'(l (. .' to the zoanthids (see ROCHE et TIXIER-DURIVAULT, 1951[28]),
. asS'lctatlOn "k 1 (F' 10)' h' b 1
10 ' ' olonies growing wlth ItS own s e eton Ig. , In t IS case a asa
1, 'cd sOme c . .) h l' . .
no Il . del' to erect the colony (see dIagnosIs; ot el' co omes growIng on ItS
'['Itcd In 01' ,
gent: ' ecorded in the literature (ROSSI, 1958 [29]). In the case the specles
I
n has been l' " "
10 / t l't is noticed some expanslOn of the zoanthlds, USIng ItS own skele-
P e al'a a, , .
Ing on , . (. h ppen al so with G. macaronesica) only generate ItS skeleton In so me
G rtw/w,¡s a "'1 h
a" . nian do not offer the appropnate support; a Slml al' p enomenon was
h
. thc gOlgo ,
ert: , '1 by ROCHE et TIXIER-DURIVAULT (1951 [28]: 405). We always fInd
d prcVIOUSY ., ,
, h'lbitats with high blOdlverslty patterns and constant water movement,
In some , ,
I .' t be toa strong' locally G, savaglia colomes have been always observed
h II can no '
g 'Ians "forest" 01' very close to those. The common occurrence of G. savaglia
the gorgon .
. P clavata it seems to be, accordmg to ROCHE et TIXIER-DURIVAULT
gorgol1lan ' "
I 1281l, a similar parasitic phenomenon also observed In G. macaroneSlca on
tharian species, .
pa G. s(Jl'lIg/ia is a planctonic captor, able to prey on several specles of copepods as we
d resl 01'these (partially digested) inside the coelenteron as a gut contento We find also
d apod Ba/ssia gasti in association with one of the colony (for more information about
ZIBROWIUS, 1985 [33]).
sexual reproduction has been noticed by budding in several polyps following a simpallems
find in G. macaronesica. We find gonads development in the specimens studn
May and August, but specimens analyzed on January do not present any gonads what
n islenl with the Mediterranean environmental conditions.
G. sal'ag/ia it seems to be a species rather known in the western Mediterranean
talonian coast, Baleares, Algerian coast, Naples and Geneva Gulfs and the Strait of
braltar) including the Adriatic Sea and apparently less common in the Levantin basin
re has hcen exclusively recorded from Aegean Sea,
Gerardia mac " .
rn the M, .aroneslca IS consldered here as a endemic element, exclusively known
acaroneslan t Th' ,
hlpelag(). . wa ers. e presence of thls taxon along the whole Macaroneslan
s and the c' .
thids is h oexlstence wlth the endemic antipatharians, where frequently the
Olhcr cOsted,.support the later hypothesis.
anog y Ocañ~d~~lc Macaronesian anthozoan, as the Actiniaria Actinia nigropunctata den
rnatic events' 03, has been recently described, although in those cases, the Pleistocene
. seems to pl '
nce of end'
emlc sp .
ay
.
an Important role and was used by some authors to justify
_
the
eCles In macaronesian waters (see den HARTOG & OCANA, 2003

[10]). We should consider this hypothesis: G. macaronesica was generated as a e
of the Pleistocene isolation suffered by Macaronesian waters fram Africa coast ~nseq
ference of these Pleistocene species, there is not consistent praves about the' ~t~
macaronesica during the Pleistocene periodo Attending to the current Macaroo ng . 1ll
graphical and ecological distributions of the zoanthid, it seems more plausibly t~eSI~
ed before the glacial period and could survive to those fluctuant climatologic ~t tl e
circalittoral or upper bathyal bottoms. The actual presence ofthe zoanthid in the ~ ang
and, especially, in the Azores support the latter hypothesis of having survivar
Pleistocene times. According to HOEK (1987) [14], the climatic perturbations e d
bring the.destruction of most of littoral species fram the Azores in opposition toC~:ld
accepted Idea (see MORTON et al., 1998 [22]); some new records of Actiniaria from
(Ocaña et al. in prep.) support the latter idea. The high ecological possibilities of G
aronesica are consistent with this assertion and invite to think about that particular id
The abundance of G. macaronesica in Cape Verde call to explore the POSsibili~
zoanthids as a Guinean element than could reach the Macaronesian archipelagos, th
the warm periods suffered during the Pleistocene. If we assume the last argument, it WIU
no rare that the species could even occur on other similar habitat from close related p
as some spots of the Mauritanian, Senegal coasts and also at Guinea Gulf, extending its
tributional range. Following the same argument, it can be also possible that G. savaglia
be a relict Guinean species that could support the cold periods inside the Mediterranean.
similar phenomenon is well known in some species of fishes such us the pomacen
Chromis chromis (Lusitanic-Mediterranean) and Chromis limbata (Macaronesic-Guin
(EDWARDS, 1986 [7]). G. macaronesica can occur apparently very little (at least in the
ical habitats available with scuba diving equipment) in the Azores than in the other i I
graup, this remarks the northern distributionallimit of this taxon. The occurrence of G.
aronesica in association with Antipathella wollastoni remarks this antipatharian to be
most common macaronesian host-substrate. Regarding this assertion there are not m
possibilities to find G. macaronesica further north than the Azores.
G. macaronesica is a new species of Gerardiidae and it constitutes the second s
of such family described worldwide, being G. savaglia the first species which belon
such family, which has been already described. The great shape sirnilaritYand other anaI
ical characters between both species of Gerardia studied in the present paper and ~he
of appropriate description of the G. savaglia in the literature gets us into the conf~slon,
.. . . M . . 1 t n (see OCANA el
we faIled III recogOlzlllg our acaroneslan maten a as a new axo . f
1995 [25]). Fortunately, the intuition of Dr. Zibrowius (com. in litt.) put us Lllalert o
affair as he always felt they were different taxa. Th s
G. macaronesica and G. savaglia present dlfferent
.
cnl
'd
om eqUlpm
. ents
.
e. "0
. . . h'l G savagl G. macaronesica presents a hlgh number of cnldom categones, meanw le· f eniciJli
ent less cnidom categories, being less armed in a general sense. The absence °f Ph
. . f h f ues o t e
(p-mastigop~ores and even holotr~chs by.other authors! III most o ~ e ISs different
ies
G. savaglia IS one of the malll cnldom dlfferences whlch makes thlS spec G sava
. . . d·ft s from ..
G. macaronesica (see Figs. 1 and 3). In addltlon, G. macaroneslca 1 er. hore
in having common elliptical capsules of special spirulae (special b-mastJg~p presen
SCHMIDT, 1974 [31]) in the body wall (see Fig. 1) meanwhile G. savag ¡a
of special spirulae (see Fig. 3). There are also some minor, but conaree
eutegory the cnidom morphological categories and others, this differences
difli re 'es umong f h 'd h h h' (
nc ., urements and placement o t e COI om t raug t e tlssues see
. h the meas
lId Wlt . t are specially concentrated in the tentacles, although they are also
11) Splrocys s . .
I aod . ,1 h tissues, being very common III the pharynx and the fllaments of
'hout al te.. . h . h
Ihroug . 'd"d e We find also some small, other tlllY, IllCOnSplcuoUS omotnc s
. )1 Gerar 11 a . . . d'ff
peeles 1 • eSI'ca not found in G. savaglia. Some shght cnldom I erences
. G macmon . . .
n 10. . T bleI) from different 10calItles has to be archlved as a part of the
. I mIes (see a . .
o co 1 f' h 'pecies thraugh the Macaronesian waters. Part of these cmdom dlfrunge
o t e s .. . Id
uoo 1 due to the fixed methodology applJed to the maten al whlch shou
hould be a so .
s . 'f" lt'es to study the material. Differences in the cnldom of the body wall
ome dIf ICU I .. .
d t the quantity of the sand Illcluded III the body wall of the speclmens.
Id be also ue o
. he oint of view of the morphology and histology, both species are very sim-
From. tI however P the species di'ff ers III . h aVlllg . d'ff I erent num ber of tentac Ies
as a conn
g oera . b . G . B h .
'lmong them being high the num er III . macaroneslca. ot specles pres-
I haraetcr, '. . . . . .
d~ocr d d mal sphincter
.
but It IS less developed III G. savagha,
"
meanwhIle III G. mac-
lea wc noticed it to be stranger and with some mesogloeal portlo~ (although stIlI have
oofirmed if this character can be considered constant) (see OCANA et al., 1995 [25];
I . 2a and b). The presence of a remarkable system of big lacunae-like along the col-
~Ihe body wall in G. savaglia, is the main micro anatomical difference between both
I . Thcse lacunae (see Figs. 4d and e) are absent from G. macaronesica and they pres-
I ndodermal content and also concentration
!hors) (see Fig. 4f).
of big homotrichs (holotrichs sensu other
G. macaronesica differs also fram other possibly Gerardiidae from more distant
on . The taxonomical relation of G. macaronesica and G. savaglia with those specimens
Iallvely idcntified as Gerardia sp. fram New Zealand (Ocaña & den Hartog, in prep.)
m lo be rather close; the absence of penicilli E (holotrichs from other authors) from the
am nts and the presence of a different morphological category of especial spirulae (b-
Ilgophores from other authors) in the body wall of the material from New Zealand make
ugh diffcrcnces with those Mediterranean and Macaronesian species. Unfortunately, the
r xamined material was not fixed in formaline and this complicates enormously the
nomieal work, especially in the cnidom concern. However, the different sizes categories
nd I~allthe specimens examined differ from what was found in G. savaglia and G. macne
lea. Thc material from Indopacific (ZMUC/X489) is growing on antipatharian too,
I h~weda characteristic feature in the mesogloea of the body wall not found in the other
thld> T~c other material studied from the Indopacific (ZMUCITZ61; ZMUC/Z9695)
po slbly In association with gorgonians skeleton (although we can not assure it) and
nt diffcrcnt e 'd . .
ni om measurements than what IS expected from our Medlterranean and
aroncsian m t . 1 Th .
I et d'h a ena . e posslble Gerardia sp. (ZMUC/Z9695) present small and rather
ne ¡e. (omotnehs in the body wall, also found in some material identified
(a scc Table 1 and Fig. 1).
as G. mac-
The authors . h
dlgos. all of th: IS
to thanks to Drs. Fernando Tempera, Filipe Porteiro and Frederico
m fram the Departament of Oceanography and Fisheries of Azores

University, their interest, sending the material from the Azores, within the p .
rOJects
(EU LIFE- Nature B4-3200/98-509) and Marov (PDCTM/P/MARl15249/1999) .
tially funded the mission that allowed the collection of the species in the A~oreWhlch
thank to Ricardo Araújo, Director of the Museu Municipal do Funchal who sent S : We
material for our paper and provided us with collection numbers and other rele~rn
Special gratitude is due to Dr. Rogelio Herrera who pro vides us with some ima ant
logical interest. Our colleagues Gustavo Lorenzo, Fernando Espino and Dr. Peterg~.of
sent us some interesting images of biological value. To our administrative person Ilrtz
(Ceuta) who meet all my requirements and especially to Enrique Gómez Barceló. ~rOf
Núñez identified species of polychaeta associated with the antipatharians. Part of the'J
rial studied in this paper was collected during Bentos Project (1980-1985) and Farro
Expedition (1990). Some funding in order to study part of the Mediterranean material
provided by Fauna Ibérica Project (subproject of Anthozoa) from Museo Nacional
Ciencias Naturales of Madrid (MCNM). Histological sections of the Mediterranean ma
al were partly prepared by Mr. T.J.G.M. van Oyen from the RMNH. Dr. Hoecksema and
M. Slierings, both from the RMNH of Leiden and Miguel ViIlena from MCNM who sent
collection number and accepted our type material. The authors thank to Gustavo Dioni
interest in collecting samples from the fishermen of Gerardia macaronesica. We are
grateful to Dr. Juan José Bacallado, head of the Museo de Ciencias Naturales de Tene
for inviting and encouraging us to publish this paper in a Canarian Journal. Many than
Dr. Zibrowius who puts us in alert of the genus Gerardia and provided us with sugge ti
in behalf of our paper.
This paper is part of a current research on zoanthids, headed by the Canary 1 I
Government (Viceconsejería de Medioambiente).
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(Anthozoa: Zoantharia) en las Islas Canarias. Tethys, 11(1): 89-91.
[2] BRITO, A., 1985. Estudio taxonómico, eco lógico y biogeográjico de los.Ant~zoosdde
región litoral de las islas Canarias. Tesis doctoral (no publicada). Umversldad e
Laguna.
[3] CALVÍN, J. e., 1995. El Ecosislema Marino Mediterráneo: Guía de su Flora y Fa
Calvín editor. 797 pp.
. h Ql/;, K Vele
[4] CARLGREN, O., 1895. Ueber de Gattung Gerardw Lac.-Dut. 'jvers. .
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. . . pomacenln
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dI: la pata
la)
catalanas (MedIterraneo occidental). MISc. Zool.,
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•.~ C DEN O. OCAÑA & A. BRITO, 1993. Corallimorpharia collected dur-
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l. C'nidom of G
erardia macaronesica. The letters refer lo lhose in Table 1.

PHARYNX
Penicilli A
FILAMENTS
Homotrich? Penicilli E
Ur 2 H'
. "tology and '
of the ,phincter' mlcroanatomy of Gerardia macaronesica: a. sphincler: b. detail of lhe mesogloeal por-
'PennatogOnium' h ' e, mesogloeallacunae; d, siphonoglyph: e. lrilobulated fílament: f. unilobulated filamenl; g,
, ,ovule.

COLUMN
Penicilli E
•.• 4. ""tOlo gy and .
unac )\tcm {lf h mlcroanatomy of Gerardia savaglia: a, sphincler; b, pharynx and siphonoglyph; c, ovule;
tcm t c body wall; e, lacunae syslem ofthe body wall and detritus holes; f, hornotrichs in the lacu-

· from Gran e
Figure 5. Polyps morphology and coloration 01"Gerardia lIIac{//YJllesica: a. detall of a colony. f a colon
(El Cabrón): b. detail of a colony from Canaries: e. detail 01"a colony I"rom Cape Verde: d. detall ~ran Can
Cape Verde (Tarrafal): e. detail of the basal plate 01"a colony growing on its own skeleton from
Cabrón).
6. AutoeeOlogieal data f .
ape Verde (Tarrafal)' a .a. GerardlQ macaronesica growing on the antipatharian Tallacelipalhes cavernicoa
. ,Inltlal phase of the colonization: b, antipatharian colony almost covered by G. maca-

F
·
Igure
7 .
. Autoecologlcal data
. ..
of GerardlO macarolleslca grawIng
. h 11 llastani al Canari
on Anllpat e a ;va f groW1
Azores: a, a colony placed in a vertical wall (Azores); b, lhe same habitat, but noticeable is the way o
this antipatharian (Canaries).
I't 8. \utnccnlngical data of .
In a Ingle plane f GerardlO macaronesica growing
Unt ncar MOnta- ro
m
¡a rack platform with sand influence;
na C ara lslet.
on its own skelelon
b, a 101of colonies
al Canaries: a. a big colony
placed on a slope botlom of

ck pla 1forrn :"
Figure 9. AUloecological dala of Gerardia macaronesica growing on vertical walls and ro Y
zoanthids can cover a big piece of the substrate; b, detail.
I't lO. Hig colo .
urircu cla,' 1 ~,Yof Gerardla savaglia growing on its own skelelon in lhe circaliuoral communily
a a forest") from Ceuta.