5 Porifera
5 Porifera
5 Porifera
at
Porifera
Deep-sea representatives of the Porifera (sponges) belong Both hexactinellids and cladorhizids are not members of
mainly to the classes Hexactinellida and to a special family of the true vent communities. Up to now, they have never been
Demospongiae, class Poecilosclerida, the Cladorhizidae, al- observed in the immediate environment of active smokers.
though many other Demospongiae are able to live in bathyal However, a few hexactinellids and numerous cladorhizids occur
environments. Hexactinellida and Cladorhizidae display two at few distance from the active vents in high diversity, and at a
very different life strategies allowing them to survive in deep- high proportion of undescribed species in the case of
sea conditions where life is difficult for sponges, which are fil- cladorhizids (VACELET, in press). These carnivorous sponges
ter-feeders living on tiny particles. The hexactinellids have a benefit, at least in part, from a general distant enrichment
highly developed filter-feeding system with large cavities lined around the vents At the Lau sites, cladorhizid specimens caught
by extremely thin living tissue to maximize the volume of fil- tens of larval zoe of crab Austinograea alaysae (M. Segonzac,
tered water and the ability for retention of particles. Converse- pers. obs.). It cannot be excluded, however, that they belong to
ly, the Cladorhizidae have lost their filter-feeding mode and re-
a “non-vent” fauna taking advantage only of the presence of
ly on carnivory of relatively large prey, mostly crustaceans, a
hard substrata offered by the lava issued at rapidly spreading ar-
very unusual and unexpected mode of feeding in sponges. This
eas and that their high diversity is due to a higher sampling ef-
strategy allows cladorhizids to reach the hadal zone with a
fort in these areas.
depth record of 8840 m, whereas the hexactinellids and “nor-
mal” demosponges do not exceed 7000 m.
1: Abyssocladia sp. from Kilo Moana, Lau Back-Arc Basin, TUIM 05 cruise © M. Tivey.
The cladorhizids are generally small sponges with a mor- These deep-sea sponges are highly fragile. The hexactinel-
phology rather unusual in sponges. They are erect, with a pin- lids, although supported by a highly developed siliceous skele-
nate or symmetrical shape, and their aquiferous system, long ton, have very thin, partly syncytial living tissue which is very
considered as a diagnostic characteristic of the phylum Porifera, difficult to preserve properly. They must be preserved in good
has been discarded in most genera. A special mention, howev- histological or cytological fixatives as soon as possible after col-
er, should be made for the genus Chondrocladia, in which a lection. The carnivorous sponges have very fragile appendages
canal system and choanocyte chambers are maintained, but that are generally only partially preserved during collection.
serve to inflate large spheres on which the prey is captured. The Observation of their morphology from manned submersibles or
prey, mostly small crustaceans, are passively captured by means ROVs is important. For instance, a long time was needed to re-
of hook-like cheloid microscleres lining the surface of the alize that a mysterious organism with pedunculate, translucent
sponge and catching the setae or appendages of various inver- spheres actually corresponds to the genus Chondrocladia as it
tebrates, acting as a Velcro cover. In the absence of a digestive was known from preserved specimens, in which the spheres
cavity, cells act individually to digest the prey. were contracted. Their histology and reproduction is poorly
known and could be studied only from specimens preserved in
The classification of the carnivorous sponges is presently
formalin or fixatives for electron microscopy. The present un-
rather problematic (HAJDU & VACELET 2002). They all belong
certainties in the distinction of evolutionary lines in carnivo-
to the order Poecilosclerida, and most are classified in the fam-
rous sponges require more information on their molecular char-
ily Cladorhizidae, with several genera dinstinguished mainly by
acters, and the preservation of fragments in alcohol is highly
the microsclere spicules. These microscleres, however, belong
desirable.
to different evolutionary lines of the Poecilosclerida. Further-
more, it appears that a carnivorous feeding habit also exists in
some other poecilosclerids displaying a similar morphology, but
classified in different families due to their different cheloid mi-
croscleres. Molecular characters have not yet been used for the
distinction of the evolutionary lines of these sponges.
References:
HAJDU E. & J. VACELET (2002) in HOOPER J.N.A. & R.W.M VAN SOEST (Eds.) Systema Porifera: A Guide to the Classification of Sponges. Vol. 1: 636-641.
VACELET J. (in press) Zool. J. Linn. Soc.
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
Size: 30 mm high, with a body 10 mm in diameter. Biology: On a dead smoker near an active site, water tempera-
ture 2.6°C at the site of collection. A carnivorous feeding habit
Color: White in alcohol.
has been demonstrated in another species of the genus.
Morphology: A thin peduncle bearing an ovoid or subspherical
Distribution: North-Fiji Back Arc Basin: site White Lady.
body made of radiating fascicles. No aquiferous system. Skele-
ton: Peduncle made of longitudinally arranged long fusiform
styles; radiating fascicles made of fusiform styles and smaller
styles with the tip outwardly directed. Spicules: fusiform styles,
620-2500 x 7-35 µm; arcuate isochelae, 80-170 µm; abysso-
chelae with the front alae long, nearly in contact with the op-
posite ala, 40-45 µm; anisochelae, generally twisted, one end
tridentate, the other end with fused alae, 9.5-11 µm; sigman-
cistra in two classes, 30-40 µm and 9.5-12.5 µm.
1A: View of the holotype, scale bar 3.4 mm; B: Arcuate isochela 1, scale bar 12 µm; C: Arcuate isochela 1, scale bar 22 µm;
D: Arcuate isochelae 2 (abyssochelae), scale bar 7.4 µm; E: Anisochelae, scale bar 2.5 µm; F: Sigmancistra 2, scale bar 2.2 µm;
G: Sigmancistra 1, scale bar 4.4 µm; H: Sigmancistras 1 and 2, scale bar 4.4 µm; I: Style, scale bar 47 µm; from VACELET (in press).
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
1A: View of the holotype, scale bar 4.3 mm; B: Part of a paratype, scale bar 3.4 mm; C: Paratypes, scale bar 2 mm; D: Isochelae,
scale bar 5.5 µm; E: Sigmancistra 1 and 2, scale bar 1.4 µm; F: Style of the axis, scale bar 64 µm; G: Styles of the lateral processes,
scale bar 40 µm; H: Diverse sizes of substrongyles of the base, scale bar 33.4 µm; from VACELET (in press).
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
1A: Holotype and paratype, scale bar 3.8 mm; B: Style of the axis, scale bar 83 µm; C: Head and tip of a style of the axis, scale
bar 28 µm; D: Style of the filament axis, scale bar 35 µm; E: Substrongyle of the base, scale bar 37 µm; F: Acanthotylostrongyle,
scale bar 7.5 µm; G: Head and tip of an acanthotylostrongyle, scale bar 2.3 µm; H: Anisochela 1, scale bar 4.1 µm; I: Anisochelae
2, scale bar 2 µm; J: Anisochela 2, back view, scale bar 2 µm; K: Sigmancistra, scale bar 2.9 µm; from VACELET (in press).
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
1A: Specimen from Lucky Strike, scale bar 5 mm; B: Style of the filament, scale bar 50 µm; C: Style of the filament and
head of style of the axis, scale bar 40 µm; D: Strongyle of the basis of the axis, scale bar 30 µm; E: Spinose mi-
crostrongyle, scale bar 6 µm; F: Sigmancistra and anisochela II, scale bar 4 µm; G: Anisochela I, scale bar: 8 µm; H:
Anisochela II, scale bar 2 µm; I: Basis of anisochela II and end of sigmancistras, scale bar 2 µm.
References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.
140 µm, alae 25 µm long, and 20-32 µm, alae 10-11 µm long;
Size: Up to approximately 50 cm high, with inflated spheres 3-
sigmas 45-120 x 2-3 µm.
5 cm in diameter.
Biology: The collected specimen was rooted in sediment be-
Color: White in life, yellowish white to clear brown in alcohol.
tween pillow lava, near active hydrothermal sites, but in an
Morphology: Large stalked sponge, composed of a rhizoid fixa- area still with low density of animal life. Sponges of similar
tion system, a cylindrical stalk ending in an enlarged, ovoid morphology have been often observed on various sites of the
body from which radiate in all directions secondary branches, East Pacific Rise, either rooted in sediment or attached to pil-
each ending in translucent sphere in the living animal, in an ir- low lava, always at some distance from the rich animal com-
regular swelling including crustacean debris on the preserved munities of the active hydrothermal sites. Their identification
specimen. Aquiferous system present, with large choanocyte to C. lampadiglobus cannot be ascertained from external mor-
chambers and canals ending in the inflatable spheres. Skeleton: phology alone, and several species may be present in this large
stalk and branches made of large fusiform styles longitudinally geographic zone. Carnivorous mode of feeding.
arranged, covered with a feltwork of rugose tylostyles; terminal
Distribution: East Pacific Rise: collected at 17°S. Sponges of
swellings made of smaller styles with an outer cover of numer-
similar morphology observed in various sites extending from
ous microscleres. Spicules: fusiform styles, 700-4750 x 15-75 µm
23°S to 13°N, 2600-3000 m deep.
and 510-580 x 17-30 µm; rugose tylostyles, 300-535 x 5-6 µm;
anchorate isochelae in two sizes, with six alae at each end, 123-
1: ROV Tiburon/2003, dive 556, 20°47.03’N, 109°08.98’W, 2555 m; by R. 2A: Holotype; scale bar 17 mm; B: Tylostyle of the
Vrijenhoek © MBARI. cover of the stalk; scale bars: 42 µm and 4.2 µm; C:
Style of the stalk; scale bar 30 µm; D: Style of the
body surface; scale bar 63 µm; E: Anchorate isochelae
1; scale bar 14 µm; F: Anchorate isochelae 1; scale bar
9.6 µm; G: Two anchorate isochelae 2; scale bar 7 µm;
H: Sigma; scale bar 10 µm; from VACELET (in press).
3A: Holotype, East Pacific Rise: 17°S, 2714 m © Ifremer/Naudur; inset: Unidentified worm gliding on the lower left sphere;
B: Collection of the holotype by the “Nautile“ submersible © Ifremer/Naudur; C: Presumed C. lampadiglobus, Geocyarise 3
(CY 30), 2622 m, 12°54’N, 103°58’W; cruise Geocyarise © Ifremer; D: Presumed C. lampadiglobus, Geocyarise 1 (CY 07), 2623 m;
© Ifremer; modified from VACELET (in press).
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
43
© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
1A: Type specimen from Lofoten, 550 m, approximately 60 mm high, from SARS (1872); B: Specimen from MAR, Logatchev
vent site, DiversExpedition, dive 3668, 3012 m, with oocytes and embryo; scale bar 3 mm; C: Style and anisochelae; scale
bar 30 µm; D: Anisochela; scale bar 5 µm; E: Sigmas; scale bar 25 µm; F: Sigmancistra; scale bar 8 µm.
References:
LUNDBECK W. (1905) The Danish Ingolf-Expedition 6(2): 1-219.
SARS G.O. (1872) in Kongelige Norske Universitet (Ed.) Spongiae. Volume I. Brøgger & Christie, Christiania, Norway: 1-82.
1A: View of the holotype (left), two paratypes and a fragment of a paratype; scale bar 3 mm; B: Style; scale bar 90 µm;
C: Developmental stage of anisochela; scale bar 3.1 µm; D: Anisochela; scale bar 3.7 µm; E: Anisochela, back view; scale bar
3.2 µm; F: Sigma; scale bar 12 µm; G: Sigmancistra; scale bar 5.3 µm; H: Sigmancistra; scale bar 5.3 µm; from VACELET (in press).
Reference:
VACELET J. (in press) Zool. J. Linn. Soc.
700 x 6-14 µm; palmate isochelae, 75-130 µm, in two size cate-
Size: 14-120 mm high, shaft 3-4 mm wide, lateral filaments up
gories in some specimens; placochelae I, 60-90 µm; placochelae
to at least 8 mm long and 70-100 µm in diameter.
II, 100-130 µm, absent in the type specimen; sigmancistra I, 11-
Color: Clear brown in alcohol. 14 µm; sigmancistra II, 22-30 µm, sometimes absent.
Morphology: Erect, pennaceous, composed of a short peduncle Biology: Type specimens collected from 91 m. Fixed on solid
and a flattened shaft bearing symmetrically paired lateral fila- substrata up to 4960 m deep. The presence of lateral filaments
ments, broken and reduced to their basis in most specimens. No covered by microscleres regularly arranged with the teeth up-
aquiferous system. Skeleton a spicular axis of styles, condensed wards, and the absence of canal system and apertures in the col-
at the basis and divided in parallel fibres upward; lateral fila- lected specimens suggest a carnivorous mode of feeding, similar
ments with a twisted axis of subtylostyles. Microscleres most of- to that developed in Cladorhizidae. This is supported by the
ten regularly arranged at the surface of the shaft and of the fil- presence of debris of small crustaceans in the best preserved
aments, with the teeth outwardly directed. Spicules: Styles of specimens from Barbados (cruise Manon © Ifremer).
the axis, fusiform with an obtuse point, 900-1550 x 15-40 µm;
Distribution: Cape Verde Islands (91 m); Barbados (4960 m);
strongyles in the basis of the peduncle, slightly curved or flexu-
Mid-Atlantic Ridge: Lucky Strike; Rainbow.
ous, 210-490 x 14-22 µm; subtylostyles in the filaments, 330-
References:
TOPSENT E. (1909) Bull. Inst. Océanogr. Monaco 151: 1-23.
TOPSENT E. (1928) Résult. Camp. Sci. Prince Albert Ier de Monaco 74: 1-376.
2: Spicules (SEM); after BOURY-ESNAULT & DE VOOS (1988). 3: A preserved specimen; by P. Briand @ Ifremer.
Reference:
BOURY-ESNAULT N. & L. DE VOS (1988) Oceanol. Acta 8: 51-60.
The leptolid fauna of hydrothermal vents of oceanic ridge Collecting activities at or near Mid-Atlantic vents areas
sites is still poorly investigated and understood. Fragmentary have currently provided material for proper identification of 3
colonies of leptolids from oceanic ridges in the Pacific have so anthoathecates and 23 leptothecates; 4 more leptothecates
far made it possible to identify Zygophylax cervicornis (NUTTING, could only be identified to the genus (CALDER & VERVOORT
1905), Zygophylax sp. and Halecium tenellum HINCKS, 1861. 1998). One more species (Hydrallmania falcata) was dead when
While from vents on the Mid-Atlantic Ridge only a single an- collected and accidentally introduced; all remaining leptothe-
thoathecate leptolid, Candelabrum serpentarii SEGONZAC & cates belong to the deep-water fauna and are only party re-
VERVOORT, 1995 can be considered to be a true companion of stricted to the Atlantic.
the vent community. A second species, Candelabum phrygium
Because of the technique used to collect animals at ridge
(FABRICIUS, 1780) is a North Atlantic deep-sea inhabitant oc-
sites, only rarely do complete animals or colonies become avail-
casionally found associated with the hydroythermal vent fauna.
able for scientific study. Anthoathecates in particular are hard
Another anthoathecate, Bouillonia sp. was found on the to collect and they must be narcotized prior preservation. So
Mid-Atlantic Ridge in the Gulf of Guinea area; it probably rep- far, only a single complete animal of Candelabrum serpentarii has
resents the deep-water leptolid community without direct been obtained, in addition to many incomplete individuals,
affinities to hydrothermal vents (SVOBODA, STEPANJANTS & which have been studied. Anthoathecates are best preserved in
LJUBENKOV, in press). Photographs taken during surveys of Mid- 6% formalin (after prior relaxation by adding MgSO4 solution
Atlantic vent areas show a third species of anthoathecate, oc- to the seawater), but Leptothecates get very brittle in formalin
casionally in great numbers; it may represent a species of Tubu- and are best preserved in 70% ethanol. These preservation
laria or Ectopleura. There is presently no material available to techniques only apply to material for routine taxonomic inves-
substantiate that guess. A probably undescribed species of Hy- tigations; more sophisticated methods are necessary for modern
dractinia has also been observed. DNA techniques.
References:
CALDER D.R. & W. VERVOORT (1998) Zool. Verh., Leiden 319: 1-65.
SEGONZAC M. & W. VERVOORT (1995) Bull. Mus. Natl. Hist. Nat., Paris (4)17(1-2): 31-64.
SVOBODA A., STEPANJANTS S. & J. LJUBENKOV (in press) Zool. Meded., Leiden.