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

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


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 Hydrozoa symbiotic with Porifera

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).
Fig. 2. Cladonema sp. in Ircinia sp. The hydranths are able to com-
pletely withdraw into the sponge pores.
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 and one or two thinner supporting tubes developed from
shell margin of a specimen of Hinnites multirugosus basal part of hydrocaulus (Petersen 1990; Schuchert
(Brinckmann-Voss 2000). This fact suggests that the 2001).
association between this species and the sponge might Hybocodon cryptus Watson, 1984 generally grows in
be occasional. sparse clusters of up to 15 well separated stems, growing
to 8 cm in height from a hydrorhiza deeply embedded in
Genus Bicorona unidentified sponges; it occasionally also occurs on old
Bicorona elegans Millard, 1966 was described with the shells (Watson 1984).
stem embedded in the tissue of an unknown sponge (Mil- Hybocodon unicus (Browne, 1902) has a branched or
lard 1966). The species was subsequently recorded from unbranched hydrorhiza, widening to irregularly shaped
several South African localities but the association with tuber-like structures embedded in the sponge Hymeniaci-
sponges was never mentioned (Millard 1975). The only don perlevis (Montagu, 1818) (Petersen 1990).
other known species of this genus, Bicorona tricycla
(Schuchert, 1996), occurs on rock and macroalgae at low Genus Zyzzyzus
depth (Schuchert 1996). 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
Family Cladonematidae finger-shaped hydrorhizal processes and the hydrocaulus
embedded for one-third in the host body.
Genus Cladonema
Zyzzyzus warreni Calder, 1988 lives in a large variety of
Cladonema sp. has been recorded by Boero & Fresi sponge species (Haliclona clathrata Dendy, 1905, H. per-
(1986) as Sarsia sp., the polyps stretching out the pores mollis (Bowerbank, 1866), Clathria fasciculata Wilson,
of the sponge host and, if disturbed, retracting completely 1925, Aaptos ciliata (Wilson, 1925), Mycale angulosa
into the sponge body, becoming invisible from the out- (Duchassaing & Michelotti, 1864), Mycale laxissima
side (Fig. 2). M. P. Miglietta (personal communication) (Duchassaing & Michelotti, 1864), Mycale microsigmatosa
found this species along the Apulian coast and obtained a Arndt, 1927, Tedania ignis (Duchassaing & Michelotti,
Cladonema medusa from it. 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
Family Tubulariidae (for a revision see Petersen 1990) sponge, stating ‘The sponge appears to attempt to shut
itself off as much as possible from the hydroid; the tissues
Genus Hybocodon
of the sponge in the immediate neighbourhood of the
The hydroids of all three species of Hybocodon have been hydroid are denser and more fibrous than further in,
recorded living in sponges. Hybocodon prolifer Agassiz, forming a kind of cyst-wall’. Calder (1988) reported this
1862 is characterised by a single hydrorhizal process, bur- species on sponges, but also on the hydroid Eudendrium
ied in sponge tissue, widening toward its distal extremity, carneum.

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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).

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

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

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


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 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
Dipurena simulans Haliclona simulans Bouillon (1965), Schuchert (2001)
Dipurena strangulata Microciona prolifera Schuchert (2001)
Dipurena spongicola Halichondria panicea, Suberites domuncula(?) Anger von (1972), Schuchert (2001)
Sarsia bella Unidentified Brinckmann-Voss (2000)
Cladonema sp. Petrosia ficiformis Boero & Fresi (1986)
Ircinia sp.
Bicorona elegans Unidentified Millard (1966)
Hybocodon prolifer Unidentified Agassiz (1862), Petersen (1990)
Hybocodon unicus Hymeniacedon perlevis Browne (1902), Petersen (1990)
Hybocodon cryptus Unidentified Watson (1984)
Zyzzyzus warreni 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
Zyzzyzus spongicolus horny sponge Lendenfeld von (1885), Watson (1978)
Zyzzyzus floridanus Callyspongia vaginalis Petersen (1990)
Zyzzyzus calderi Tedania sp. Petersen (1990)
Ectopleura exxonia Unidentified Watson (1978)
Tubularia ceratogyne Halicondria panicea Pérèz (1925)
Sphaerocoryne agassizi Unidentified McCrady (1859), Petersen (1990)
Sphaerocoryne bedoti Unidentified Pictet (1893), Millard (1975), Petersen (1990)
Sphaerocoryne sp. Aka mucosa Puce S., personal observation
Heterocoryne caribbensis Mycale sp. Wedler & Larson (1986)
Cytaeis spongicola Stammophyllum, Psammophyllum, Haeckel (1889), Rees (1962)
Psammopemma
Cytaeis abissicola Stammophyllum, Psammophyllum Haeckel (1889), Rees (1962)
Gastroblasta cf. raffaelei Petrosia ficiformis Boero & Fresi (1986)
Ircinia sp.
Bibrachium euplectellae Euplectella aspergillum Schulze (1880)
gymnoblastic hydroid Walteria flemmingi, Walteria leuckarti Tuzet (1973)

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

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


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

Dictyoceratida (1 species), and to the Hexactinellid order Boero F. (1984) The ecology of marine hydroids and effects of
Lyssacinosida (3 species). These species do not share any environmental factors: a review. Pubblicazioni della Stazione
peculiar morphological or skeletal characters suggesting Zoologica di Napoli I: Marine Ecology, 5, 93–118.
selectivity in the hydroid choice. Recently, the selectivity Boero F., Bouillon J. (in press) Cnidaria and Ctenophora. In:
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of actinulae is different in different species of sponges sug- Boero F., Fresi E. (1986) Zonation and evolution of a rocky
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Conclusion
Boero F., Bouillon J., Gravili C. (2000) A survey of Zanclea,
The integration of hydroid colonies into sponge hosts is Halocoryne and Zanclella (Cnidaria, Hydrozoa, Anthomedu-
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