Marine Ecology.

ISSN 0173-9565

REVIEW ARTICLE

Hydrozoa (Cnidaria) symbiotic with Porifera: a review

S. Puce1, B. Calcinai1, G. Bavestrello1, C. Cerrano2, C. Gravili3 & F. Boero3
1 Dipartimento di Scienze del Mare, Università Politecnica delle Marche, Via Brecce Bianche, Ancona, Italy
2 Dip.Te.Ris., Università di Genova, Corso Europa, Genova, Italy
3 Di.S.Te.B.A., Università di Lecce, Lecce, Italy

Keywords Abstract
Hydroids; hydrozoans; marine benthos;
sponges; symbiosis. Many hydroids are symbiotic with other organisms. Sponges, because of their
complex canal system and their filter-feeding activity inducing a continuous
Correspondence water flow, are used by numerous species as either exclusive or facultative sub-
S. Puce, Dipartimento di Scienze del Mare, strata. The associated hydroid fauna thriving on or inside sponges shows a
Università Politecnica delle Marche, Via Brecce
wide range of relationships with their hosts. Hydroids may be simply epibiotic
Bianche, I-16131 Ancona.
E-mail: s.puce@univpm.it
on sponges, their stolons running on the host surface. Alternatively, the stolons
may grow inside the sponge body, the polyps emerging from the sponge sur-
Accepted: 16 August 2005 face, having also the possibility of retraction inside the sponge tissue. Finally,
stolons and branches may develop deeply inside the sponge body, producing a
doi:10.1111/j.1439-0485.2005.00050.x skeletal network for sponge growth. This paper reviews the described relation-
ships of hydrozoans symbiotic with sponges and reports new observations.

Relationships between hydroids and sponges

Problem
The hydroids of several hydroidomedusan taxa show spe-
cialized symbiotic associations with other organisms, see
Piraino et al. (1994) and Boero & Bouillon (in press) for
lists. Sponges, with their body permeated by a complex
canal system where a continuous water flow is induced by
their filter-feeding activity, are temporary or permanent
substrata for many hydroid species. A rich associated
hydroid fauna thrives on or inside sponges; the records of
hydroid-sponge associations are scattered in the literature
and mostly report only on the co-occurrence of these ani-
mals. The techniques employed for the preservation and
study of sponges are not conducive to the identification
of hydroids: data about this symbiosis are extremely poor
in the spongological literature (Sarà & Vacelet 1973) and
most information derives from hydroid-centred studies.
In this paper, the described relationships of hydrozoans
symbiotic with sponges are reviewed, together with new
observations.
Types of sponge–cnidarian association
The Cubozoa are the only cnidarian class with no species
associated with sponges. In the Anthozoa, the Zoanthari-
an Parazoanthus axinellae (Schmidt, 1862) is often epibi-
otic on sponge species of the genus Axinella (Arndt 1936;
Lewis 1982). Host-selectivity, however, is not exclusive for
this species, which, in fact, commonly settles directly on
overhanging rocks. Symbiosis between sponges and octo-
corals is rarely recorded (Soest van & Verseveldt 1987;
Calcinai et al. 2004).
In the Scyphozoa, the polyps of Nausithoe punctata
Kolliker, 1853 are strictly associated with horny sponges
and might contribute to their skeletal structures (Uriz
et al. 1992).
The greatest diversity in the association between Cnid-
aria and Porifera, however, is found in the Hydrozoa.
The relationship between hydrozoans and sponges can be
divided into three categories:

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Hydrozoa symbiotic with Porifera Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero

1 Hydroid larvae (either planulae or actinulae) settle on

a non-sponge substrate and later a sponge overgrows
the hydrorhizae of an already formed colony. Encrust-
ing sponges are the best candidates for such
association pattern, which might be considered as
occasional (e.g. Riedl 1966; Boero 1984; Gili & Hughes
1995).
2 Hydroid larvae settle on sponges, but the hydrorhizae
of the resulting colony are not embedded in the sponge
tissues. The association is occasional and the involved
hydroid species can grow on various substrates (Boero
1984).
3 Hydroid larvae penetrate the sponge host, developing a
hydrorhizal system embedded in its tissues. The asso-
ciation is usually exclusive and obligatory.
Only the third case can be considered as a real symbio-
sis, evolved by coevolutionary processes. The hydroid–
sponge integration can differ according to the hydroid
species:
1 The hydranths protrude from the sponge surface and
cannot retract into it.
2 The hydranths protrude from the sponge surface but
can retract into the host body, completely disappearing. Fig. 1. Dipurena halterata in Chondrilla nucula. The hydranths cannot
3 The hydranths grow into the canal system of the withdraw into the sponge.
sponge and are not visible from the outside.
Most of the hydroid species strictly associated with
sponges and never found on other substrates are referred 1826) (Rees 1938 as Chalina montagui), Haliclona simu-
to the families Corynidae, Tubulariidae and Sphaeroco- lans (Johnston, 1842), Petrosia ficiformis (Poiret, 1789),
rynidae, but some are also Cytaeididae and Campanularii- Chondrilla nucula (Schmidt, 1862) and Spirastrella cuncta-
dae. trix (Schmidt, 1868).
The stolons, covered by thin perisarc, are deeply
embedded in the tissues of the sponge host (Schuchert
Family Cytaeididae 2001). In the Ligurian Sea (Italy), the hydroid is charac-
terised by an annual cycle, occurring only during a short
Genus Cytaeis
period of the year (Boero & Fresi 1986). The rest of the
Cytaeis spongicola (Haeckel, 1889) and Cytaeis abissicola year is presumably spent as resting stolons surviving in
(Haeckel, 1889) occur in association with a genus the sponge tissues (Fig. 1).
of Demospongiae, Psammoclema Marshall, 1980 (¼ Dipurena simulans Bouillon, 1965 grows in and on the
Psammopemma) and with some other genera (Stammo- sponge Haliclona simulans: the stolons, covered by thin
phyllum, Psammophyllium, Cerelasma identified by perisarc, are deeply embedded in sponge tissue. Hydrant-
Haeckel (1889) (Rees 1962) whose attribution to the hs usually have 10–13 tentacles, but those growing in the
phylum Porifera is now dubious (Hooper & Soest van sponge canal system have up to 16 tentacles. Only the
2002). polyps living in the oscula of the host become sexually
competent and develop medusa buds (Bouillon 1965).
Colonies of Dipurena halterata and Dipurena simulans
Family Corynidae (for a revision see Schuchert 2001) can live together in the same specimen of Haliclona simu-
lans. The polyps of the two species are very similar and
Genus Dipurena
distinction is based only on medusa characters (Bouillon
The hydroid of Dipurena halterata (Forbes, 1846) was 1971).
described in British waters as associated with sponges Dipurena strangulata McCrady, 1859 presents stolons
(Rees 1939). The association has also been recorded from with thin perisarc and deeply embedded in the tissue of
the Mediterranean Sea (Boero & Fresi 1986). The hydroid the sponge Clathria prolifera (¼ Microciona prolifera)
lives in and on the sponges Haliclona cinerea (Grant, (Ellis & Solander, 1786) (see Schuchert 2001).

74 Marine Ecology 26 (2005) 73–81 ª 2005 Blackwell Publishing Ltd

Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero
Dipurena spongicola Anger von, 1972 occurs in the
canal system of the sponge Halichondria panicea (Pallas,
1766). The stolons show a very thin perisarc. The stolon
tips that come in contact with the canals of the sponge
transform into hydranths and in one occasion, a young
medusa was found inside a canal of the host’s aquiferous
system. The hydroid apparently feeds on matter brought
in the sponge’s filter current, as well as on cyclopoids,
harpacticoids, ostracods, ciliates, turbellarian and nema-
todes inhabiting the canals of the sponge (Anger von
1972). Schuchert (2001) observed this hydroid living in a
sponge tentatively classified as Suberites domuncula (Olivi,
1792).
Genus Sarsia
Sarsia bella Brinckmann-Voss, 2000 is the only one
among the 10 species of the genus recorded as growing
in sponges. The unidentified sponge was encrusting the
shell margin of a specimen of Hinnites multirugosus
(Brinckmann-Voss 2000). This fact suggests that the
association between this species and the sponge might
be occasional.
Genus Bicorona
Bicorona elegans Millard, 1966 was described with the
stem embedded in the tissue of an unknown sponge (Mil-
lard 1966). The species was subsequently recorded from
several South African localities but the association with
sponges was never mentioned (Millard 1975). The only
other known species of this genus, Bicorona tricycla
(Schuchert, 1996), occurs on rock and macroalgae at low
depth (Schuchert 1996).
Family Cladonematidae
Genus Cladonema
Cladonema sp. has been recorded by Boero & Fresi
(1986) as Sarsia sp., the polyps stretching out the pores
of the sponge host and, if disturbed, retracting completely
into the sponge body, becoming invisible from the out-
side (Fig. 2). M. P. Miglietta (personal communication)
found this species along the Apulian coast and obtained a
Cladonema medusa from it.
Family Tubulariidae (for a revision see Petersen 1990)
Genus Hybocodon
The hydroids of all three species of Hybocodon have been
recorded living in sponges. Hybocodon prolifer Agassiz,
1862 is characterised by a single hydrorhizal process, bur-
ied in sponge tissue, widening toward its distal extremity,
Marine Ecology 26 (2005) 73–81 ª 2005 Blackwell Publishing Ltd
Hydrozoa symbiotic with Porifera
Fig. 2. Cladonema sp. in Ircinia sp. The hydranths are able to com-
pletely withdraw into the sponge pores.
and one or two thinner supporting tubes developed from
basal part of hydrocaulus (Petersen 1990; Schuchert
2001).
Hybocodon cryptus Watson, 1984 generally grows in
sparse clusters of up to 15 well separated stems, growing
to 8 cm in height from a hydrorhiza deeply embedded in
unidentified sponges; it occasionally also occurs on old
shells (Watson 1984).
Hybocodon unicus (Browne, 1902) has a branched or
unbranched hydrorhiza, widening to irregularly shaped
tuber-like structures embedded in the sponge Hymeniaci-
don perlevis (Montagu, 1818) (Petersen 1990).
Genus Zyzzyzus
Zyzzyzus spongicolus (Lendenfeld von, 1884) lives in
‘horny sponges’. Watson (1978) described the solitary
hydranth roots in the canal system of the host sponge as
finger-shaped hydrorhizal processes and the hydrocaulus
embedded for one-third in the host body.
Zyzzyzus warreni Calder, 1988 lives in a large variety of
sponge species (Haliclona clathrata Dendy, 1905, H. per-
mollis (Bowerbank, 1866), Clathria fasciculata Wilson,
1925, Aaptos ciliata (Wilson, 1925), Mycale angulosa
(Duchassaing & Michelotti, 1864), Mycale laxissima
(Duchassaing & Michelotti, 1864), Mycale microsigmatosa
Arndt, 1927, Tedania ignis (Duchassaing & Michelotti,
1864), characterised by different skeletal texture (Hirohito
1988). Warren (1906, p. 89) described it as Tubularia soli-
taria and suggested a parasitic interaction with the
sponge, stating ‘The sponge appears to attempt to shut
itself off as much as possible from the hydroid; the tissues
of the sponge in the immediate neighbourhood of the
hydroid are denser and more fibrous than further in,
forming a kind of cyst-wall’. Calder (1988) reported this
species on sponges, but also on the hydroid Eudendrium
carneum.
75
Hydrozoa symbiotic with Porifera Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero

Zyzzyzus calderi Petersen, 1990 was described with the

hydrocaulus embedded in a sponge of the genus Tedania.
Different from the other species, only the most distal part
of the hydrocaulus is embedded in the sponge tissue, con-
sisting of two to three small club-shaped tubers with fin-
ger-like distal processes.
Zyzzyzus floridanus Petersen, 1990 is hosted by Cally-
spongia vaginalis (Lamarck, 1814); the hydrocaulus has
globular rather than finger-shaped hydrorhizal bulbs
embedded in the sponge.

Genus Ectopleura
Ectopleura exxonia Watson, 1978, was described in associ-
ation with sponges: its hydrorhiza is a matted reticulum
penetrating 5–6 mm into the surface of an unidentified
horny sponge.

Genus Tubularia
Tubularia ceratogyne Pérèz, 1920 is associated with
Halichondria panicea. Pérèz (1925) described the rela-
tionship in detail. The symbiosis strongly influences the
ecology and the growth of the two partners. The bathy-
Fig. 3. Sphaerocoryne sp. in Aka mucosa. The hydrorhiza is filled by
metric distribution of the hydroid is determined by the
numerous nematocysts and is enveloped inside the sponge spicule
sponge distribution. The sponges colonised by T. cerato- tracts.
gyne grow along the hydroid hydrocaulus, producing
erect protrusions arising from a massive body. The
hydrocaulus, insulated by the sponge tissue, grows unti-
dily producing irregular ramifications and perisarc enveloped inside the sponge spicule tracts (Fig. 3). This
thickenings. The loss of organisation of T. ceratogyne growth structure suggests a reaction of the two partners,
when associated with the sponge suggests that environ- as recently observed in the octocoral Carijoa riisei living
mental conditions strongly modulate the processes inside the sponge Desmapsamma anchorata (Calcinai et al.
involved in the branching pattern. 2004).

Family Sphaerocorynidae
Genus Sphaerocoryne
Sphaerocoryne bedoti Pictet, 1893 was described as living
in two unidentified siliceous sponges: the hydrorhiza
embedded in the sponge tissue and the hydrocauli arising
from the pores (Pictet 1893). After this first record, S. bed-
oti was always observed in association with sponges, but
no additional information about the host is known
(Mammen 1963; Millard 1975; Calder 1988; Hirohito
1988).
Sphaerocoryne agassizi (McCrady, 1859) was figured by
Petersen (1990) as a single polyp collected in North Caro-
lina (USA) from a colony growing on a sponge.
In the Halong Bay, Vietnam, we collected several speci-
mens of Sphaerocoryne sp. growing on the fistules of the
boring sponge Aka mucosa (Bergquist, 1965). The hydro-
rhiza, running in the tissue of the fistule wall, is filled by
numerous nematocysts (stenoteles of two sizes) and is
Genus Heterocoryne
Heterocoryne caribbensis Wedler & Larson, 1986, the only
species of this genus, was often observed growing in the
sponges of the genus Mycale. No other detailed observa-
tion about the association are reported, but the picture
showing this species in the original description (Wedler &
Larson 1986) suggests that the sponge reacts to the
hydroid presence by growing along its hydrocaulus.
Family Campanulariidae
Genus Gastroblasta
Gastroblasta sp. (possibly G. raffaelei Lang, 1886), cited by
Boero (1980) and Boero & Fresi (1986) as Clytia sp., lives
in association with sponges. Recorded from the Ligurian
Sea, the Gulf of Naples and the Adriatic Sea (Italy), its
hydrorhiza is always embedded in the tissues of sponges
of the genera Petrosia and Ircinia (Fig. 4). This is the only
thecate hydroid exclusively associated with sponges.

76 Marine Ecology 26 (2005) 73–81 ª 2005 Blackwell Publishing Ltd

Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero Hydrozoa symbiotic with Porifera

Fig. 4. Gastroblasta cf. raffaelei in Ircinia sp. Hydranth and gonothe-

cae containing a single medusa bud.

Hydroids growing in hexactinellid sponges

Schulze (1880) described the association between the
sponge Euplectella aspergillum Owen, 1841 and a gymno-
blastic hydroid, named for this reason Bibrachium euplec-
tellae (Schulze, 1880). As Schulze observed (p. 671), the
hydrorhiza forms a ‘long-meshed net’ from which
hydranths ‘project freely into the inhalant lacunae, and
therefore toward the exterior, from the zone of ciliate
chambers’. The hydroid appeared so abundantly in the
Euplectella that, as noticed by Schulze, one or more
hydranths are found in almost every microscopic section
of the tube wall.
Fig. 5. Photographs of Nemalecium lighti growing on Carteriospon-
Another record of association between hydroid and gia foliascens (a) and Hydrichthelloides sp. growing on a unidentified
hexactinellid sponges is from Tuzet (1973) who observed sponge (b) from Bunaken Marine Park (North Sulawesi, Indonesia).
an undetermined gymnoblastic hydroid, morphologically
different but in behaviour similar to Dipurena spongicola,
occurring in the sponges Walteria flemmingii Schulze, the hydrorhiza and much hydrocaulus embedded in the
1886 and W. leuckarti Ijma, 1896. The sponge shape, tissue of an unidentified sponge.
characterised by prominent and irregularly distributed In Indonesian waters, the substrate-generalist Nemale-
oscula, appears determined by the presence of this hyd- cium lighti (Hargitt, 1924) is found only on the fan
roid. shaped horny sponge Carteriospongia foliascens (Pallas,
1766) (Fig. 5a) while Cladocoryne floccosa Rotch, 1871
lives on the fistules of the psammobiotic sponge Ocean-
Other records
apia fistulosa (Bowerbank, 1873). Hydrichthelloides reticu-
A colony of Cytaeis nuda Rees, 1962 is described partly lata Bouillon, 1978, originally described from Papua New
immersed in a sponge growing on living gastropod Fusi- Guinea, was observed on different substrates (corals,
nus perplexus. The stolon runs parallel to and a little tubes of polychaetes, rocks, shells, and sponges) (Bouillon
below the surface of the sponge (Rees 1962; Hirohito 1978). In the Bunaken Marine Park (North Sulawesi,
1988). No additional information is available about the Indonesia) we collected numerous specimens of Hydricht-
sponge. helloides sp. always living on sponge surfaces. They pro-
In Bermuda shallow waters, Calder (1988) observed duce a dense stolonal net not penetrating in the sponge
colonies of the clavid Turritopsis nutricula McCrady, tissue and it may be detached like a film from the sponge
1859, usually growing on different hard substrates, with surface (Fig. 5b).

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Hydrozoa symbiotic with Porifera
Discussion
Partners’ roles
Boero & Fresi (1986) recorded settlement on sponges by
species referred to the families Tubulariidae, Bougainvillii-
dae, Hydractiniidae, Pandeidae, Haleciidae, Hebellidae,
Campanulariidae, Aglaopheniidae and Syntheciidae. These
species, however, have been found also on other sub-
strates and can be considered as simple epibionts, taking
advantage of the sponge surface as substrate for settle-
ment. Their harm to the functioning of the sponge is
negligible, since the hydrorhizae are linear and do not
impair the flow of water generated by the sponge. The
incoming current generated by the sponge through its
pores might be of some advantage to the hydroids, as
suggested by Uriz et al. (1992) for Nausithoe. This water
is often loaded with nutrients and the hydranths might
increase the rate of prey capture because of the current
produced by the pumping activity of the sponge.
Hydroids obligatorily associated with sponges invaria-
bly have their hydrorhizae embedded into the host tis-
sues. A foreign body growing into another organism is
usually considered as a parasite; the association of hyd-
roids with sponges probably evolved as a form of para-
sitism in which some hydroid species succeeded in
overcoming the chemical defences that prevent the settle-
ment of other organisms on most sponges. The hydro-
rhizae of these species passed from the surface to the
inside of the host sponge, acquiring the same position
that can be observed in hydroids overgrown by encrust-
ing sponges. In this case, however, it is the hydroid that
takes the initiative of growing into the sponge through
specific larval settlement. For hydroids, the advantage of
this association is the already mentioned increase of
water renewal around the colony, enhancing feeding effi-
ciency. Furthermore, the sponge protects the hydroid
colony from predation by nudibranchs and from over-
growth by other benthic organisms. The price paid by
the sponge, also in this case, might be negligible and the
association might turn to be a form of mutualism, the
hydrorhizae becoming an accessory skeletal structure for
the sponge, as proposed by Uriz et al. (1992) for the
association of scyphozoan polyps with sponges. Along
the Portofino Promontory, Boero & Fresi (1986)
reported that Dipurena halterata is present on sponges
(Petrosia ficiformis, Spirastrella cunctatrix and Chondrilla
nucula) from January to June. It is probable that when
the hydroids disappear at the sponge surface their
hydrorhiza survives as a resting stage embedded in the
sponge tissues.
The top of sponge–hydroid association is reached by
Dipurena spongicola and Bibrachium euplectellae, showing
extreme cases of endosymbiosis. Dipurena spongicola pre-
78
Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero
sumably completes its life cycle inside the sponge canal
system (Anger von 1972).
The relationships among sponges and hydrozoans are
strongly unbalanced as to biomass: the sponge partner is
generally much larger than the hydroid guest. The only
documented cases of sponge shape modification owing
the presence of hydroids concern Halichondria panicea
and Tubularia ceratogyne, and Heterocoryne caribbensis
and Mycale sp. because the sponges react to the hydroid
presence by growing along the hydroid hydrocaules. In
this situation, sponges exploit the hydroid stem, becom-
ing erect. The body of massive sponges is three-dimen-
sional and the hydroid growth pattern inside their tissues
is different from that of the species that normally grow
on two-dimensional substrates. The possibility of growing
in a three-dimensional space produces a disorganised
hydrorhizal pattern. The same situation was recently
recorded in an octocoral living inside the sponge Des-
mapsamma anchorata (Calcinai et al. 2004).
Evolutionary trends in hydroid symbioses with sponges and
other organisms
Many hydroid species are substrate generalist, often grow-
ing also on other organisms. In some families, this
attitude represented a preadaptation to more strict rela-
tionships, leading to permanent associations. When the
association with a particular type of host is widespread
within a single genus or family, it is highly probable that
an adaptive radiation from a single ancestral association
occurred. In these cases, the ancestors, once established
the association, became the founders of monophyletic
clades containing species with similar host preferences.
The most evident example of this trend is found in the
Hebellidae (see Boero et al. 1997 for a revision) where all
the species of the genera Hebella and Anthohebella are
associated with hydroid hosts. Another case is the strict
association between hydroids and bivalve hosts, with the
genus Eugymnanthea that comprises species living exclu-
sively in the mantle cavity of bivalves (Kubota 2000).
Recently, Boero et al. (2000) revised the family Zanclei-
dae, containing many species strictly associated with bryo-
zoans, but also species associated with corals and algae.
Species unrelated with the Zancleidae (see Bavestrello
et al. 2000), however, have evolved symbiotic relation-
ships with bryozoans, involving behavioural patterns that
are strickingly similar to those of the Zancleidae. The
association with bryozoans, thus, is undoubtedly polyphy-
letic within the Hydrozoa. Other cases of strict association
between hydroids of a single clade and a definite host
type is to be found in the Hydractiniidae, with a sharp
preference towards gastropod shells, either living or
inhabited by hermit crabs. Cunningham et al. (1991)
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Puce, Calcinai, Bavestrello, Cerrano, Gravili & Boero Hydrozoa symbiotic with Porifera

Table 1. List of the records of hydroid species living with the colonies partially embedded in sponge tissue.

Hydroid Sponge host Author

Dipurena halterata
Dipurena simulans
Dipurena strangulata
Dipurena spongicola
Sarsia bella
Cladonema sp.
Bicorona elegans
Hybocodon prolifer
Hybocodon unicus
Hybocodon cryptus
Zyzzyzus warreni
Zyzzyzus spongicolus
Zyzzyzus floridanus
Zyzzyzus calderi
Ectopleura exxonia
Tubularia ceratogyne
Sphaerocoryne agassizi
Sphaerocoryne bedoti
Sphaerocoryne sp.
Heterocoryne caribbensis
Cytaeis spongicola
Cytaeis abissicola
Gastroblasta cf. raffaelei
Bibrachium euplectellae
gymnoblastic hydroid
Haliclona cinerea, Haliclona simulans, Rees (1938) as Chalina montagui Bouillon (1971)
Petrosia ficiformis, Chondrilla nucula, as Adocia simulans Boero & Fresi (1986), Schuchert (2001)
Spirastrella cunctatrix
Haliclona simulans Bouillon (1965), Schuchert (2001)
Microciona prolifera Schuchert (2001)
Halichondria panicea, Suberites domuncula(?) Anger von (1972), Schuchert (2001)
Unidentified Brinckmann-Voss (2000)
Petrosia ficiformis Boero & Fresi (1986)
Ircinia sp.
Unidentified Millard (1966)
Unidentified Agassiz (1862), Petersen (1990)
Hymeniacedon perlevis Browne (1902), Petersen (1990)
Unidentified Watson (1984)
Haliclona clathrata, Haliclona permollis Calder (1988), Hirohito (1988),
Clathria fasciculata, Aaptos ciliatus C.J.A. Campos, personal communication
Mycale angulosa
Mycale laxissima
Mycale microsigmatosa, Tedania ignis
horny sponges
horny sponge Lendenfeld von (1885), Watson (1978)
Callyspongia vaginalis Petersen (1990)
Tedania sp. Petersen (1990)
Unidentified Watson (1978)
Halicondria panicea Pérèz (1925)
Unidentified McCrady (1859), Petersen (1990)
Unidentified Pictet (1893), Millard (1975), Petersen (1990)
Aka mucosa Puce S., personal observation
Mycale sp. Wedler & Larson (1986)
Stammophyllum, Psammophyllum, Haeckel (1889), Rees (1962)
Psammopemma
Stammophyllum, Psammophyllum Haeckel (1889), Rees (1962)
Petrosia ficiformis Boero & Fresi (1986)
Ircinia sp.
Euplectella aspergillum Schulze (1880)
Walteria flemmingi, Walteria leuckarti Tuzet (1973)

showed that both host and guest lineages underwent par-
allel speciation patterns.
All the above mentioned associations involve exclu-
sively athecate (i.e. Anthomedusae) hydroids, suggesting
that the biology of this group is conducive to the estab-
lishment of symbiotic relationships. The hydroid–sponge
association is consistent with this trend and almost all
species belong to the Anthomedusae. The only known
exception is Gastroblasta raffaelei, a Campanulariid
species with polygastric medusae and hydroids living
embedded in sponge tissues. The array of genera with
sponge-inhabiting species is rather vast. Some genera are
represented by species that are all associated with sponges,
such as Gastroblasta, Heterocoryne, Hybocodon, Zyzzyzus
and Sphaerocoryne, whereas in others the association is
occasional, as it might be argued for Tubularia, Dipurena,
Cladonema, Sarsia, Cytaeis.
Sponge-inhabiting hydroids, thus, are referable to a dis-
parate assortment of phylogenetically unrelated genera.
The association with sponges evidently evolved several
times along the phylogeny of Hydroidomedusae and is,
thus, polyphyletic. In some cases, sponge-associated ances-
tors initiated adaptive radiations, leading to monophyletic
clades of sponge-associated species of the same genus,
whereas other associations arose isolately within clades
that have no tendency to evolve symbiotic relationships, as
is the case of the campanulariid Gastroblasta raffaelei.
Host diversity
We know 26 identified species of hydroid-inhabited
sponges (Table 1), belonging to the demosponge orders
Hadromerida (4 species), Poecilosclerida (8 species),
Halichondrida (2 species), Haplosclerida (8 species),

Marine Ecology 26 (2005) 73–81 ª 2005 Blackwell Publishing Ltd 79

Hydrozoa symbiotic with Porifera
Dictyoceratida (1 species), and to the Hexactinellid order
Lyssacinosida (3 species). These species do not share any
peculiar morphological or skeletal characters suggesting
selectivity in the hydroid choice. Recently, the selectivity
of Z. warreni against different sponge species was experi-
mentally tested. This experiment showed that the settling
of actinulae is different in different species of sponges sug-
gesting an important role of the sponge chemical defences
on the viability of hydroid larvae (C. J. A. Campos,
personal communication).
Conclusion
The integration of hydroid colonies into sponge hosts is
sometimes total. Some species can retract entirely into the
sponge body (Cladonema sp.) and others live completely
embedded into the host (Dipurena spongicola). Close
inspection of the surface and the inside of massive sponge
species, thus, will surely lead to the discovery of new
species of hydroids, or, at least, to the elucidation of life
cycles for species in which only the medusa stage is
described.
Acknowledgements
Financial support was provided by MIUR (PRIN and
FIRB projects), NSF (PEET project), the MARBEF Net-
work of Excellence ‘Marine Biodiversity and Ecosystem
Functioning’ funded in the EU Sixth Framework Pro-
gramme (contract no. GOCE-CT-2003-505446).
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