Three New Species of Carnivorous Deep-Sea Sponges from the DIVA-1 Expedition in the Angola Basin (South Atlantic)

ARTICLE IN PRESS

Organisms, Diversity & Evolution 5 (2005) 203–213 www.elsevier.de/ode RESULTS OF THE DIVA-1 EXPEDITION OF RV ‘‘METEOR’’ (CRUISE M48/1) Three new species of carnivorous deep-sea sponges from the DIVA-1 expedition in the Angola Basin (South Atlantic) Francisco Javier Cristoboa,b,Ã, Victoriano Urgorrib,c, Pilar Rı´ osa,b aEstacioń de Bioloxıá Marinã da Granã, Universidade de Santiago de Compostela, Casa do Ho´rreo, Ruá da Ribeira 1, 15590 A Granã, Ferrol, Spain bDepartamento de Biodiversidade e Recursos Marinõs, Instituto de Acuicultura, Universidade de Santiago de Compostela, 15706 Santiago de Compostela, Spain cDepartamento de Bioloxıá Animal, Facultade de Bioloxıá, Universidade de Santiago de Compostela. Campus Sur, 15782 Santiago de Compostela, Spain

Abstract

DIVA 1 is the ﬁrst in a series of expeditions dedicated to the study of benthic diversity in the deep sea of the Atlantic Ocean. On RV ‘‘Meteor’’ we collected samples along a transect of about 700 km in the Angola Basin. The collection of Porifera from this expedition is composed of 50 specimens or in some cases pieces of specimens collected at depths between 5162 and 5460 m. This work concerns only carnivorous sponges of which we collected 16 specimens belonging to four species. Here we describe three new species: Chondrocladia levii sp. nov., C. vaceleti sp. nov., and C. nicolae sp. nov.; we also found another species, C. cf. guiteli Topsent, 1904 recorded for the ﬁrst time from a very far abyssal basin (NW of the Galician coast). The main distinctive features of the three new species are the anchorate isochelae categories, the number of teeth on each, and the presence/absence of sigmas; C. levii has three unguiferous isochelae classes with 5, 5, and 9–10 teeth, respectively; C. vaceleti has two unguiferous isochelae classes with three teeth each; C. nicolae has only one unguiferous isochelae category with three teeth. The habitus of C. nicolae (orange in colour) is also very different from that of most of the known Chondrocladia species. r 2004 Elsevier GmbH. All rights reserved.

Keywords: Porifera; Deep sea; Carnivorous sponges; Angola Basin (Atlantic Ocean); New species

Introduction (Kro¨ ncke and Tu¨ rkay, 2003). Sponge fauna of the Angola Basin is not well known. Only the results of The Angola Basin is situated off the Namibian and some localities of the Valdivia expedition were published Angolan coast north of the Walvis ridge. Intensive about this area: Hexactinellida by Schulze (1904), research has been carried out on the Benguela upwelling Tetraxonia by Lendenfeld (1907), and Calcarea by system in the coastal Namibian waters but there is only Urban (1909) but these publications did not include little information available about the deep Angola Basin any carnivorous sponges. The sponge fauna of the Namibian coast at depths between 100 and 500 m shows a high degree of endemism besides an afﬁnity with the Ã Corresponding author. Estacio´ n de Bioloxı´ a Marin˜ a da Gran˜ a, Subantarctic, northern-Atlantic and Indian regions. It is Universidade de Santiago de Compostela. Casa do Ho´ rreo, Ru´ ada Ribeira 1, 15590 A Gran˜ a, Ferrol, Spain. now better known from works of Uriz (1985, 1988). E-mail addresses: [email protected] (F.J. Cristobo), [email protected] Carnivory is an extremely rare feeding strategy among (V. Urgorri), [email protected] (P. Rı´ os). recent sponges with conﬁrmed records only from the

Cladorhizidae. This is a typical deep-sea family re- Depths varied between 5162 and 5497 m. Chondrocladia stricted to the bathyal, abyssal or even hadal zones. species were found in four of these areas (Table 1). The Vacelet (1999) suggests that several other poecilosclerid trawl was run twice in each area. The mesh size in the families are also likely to contain carnivorous species. cod end measured 1 cm between stretched meshes. A Carnivory was first discovered in Asbestopluma hypogea fore-rope was let out 200 m, and a 500 kg weight was (Vacelet and Boury-Esnault 1995, 1996), from a attached before the rope was lowered further. Based on Mediterranean shallow-water cave where the habitat our earlier experience, we paid out a rope length of 1.8 resembles that of the deep sea (Vacelet et al., 1994). times the water depth to reach the sea floor. The sea bed Vacelet et al. (1995) also described carnivory in was trawled for 2–3.5 h at 2 knots, which equals a Cladorhiza sp. from the Barbados Trench. The feeding trawled area of 26,300–45,500 m2 (Kro¨ ncke and Tu¨ rkay strategy of the third genus of the Cladorhizidae, 2003). Chondrocladia was studied by Ku¨ bler and Barthel The dominant taxa were fishes, echinoderms, bivalves, (1999). Their investigations showed that Chondrocladia actinians, and sponges. Other animal groups were gigantea has developed carnivory but with a major present at times, but not regularly. difference: whereas the other genera have apparently In all areas, the temperature at depth was 2.48 1C. The lost their aquiferous system, Chondrocladia spp. still salinity at the bottom was 34.8 (Kro¨ ncke and Tu¨ rkay possess it. 2003). The sediment of all stations was white to light The ‘‘Meteor’’ expedition Me48/1-DIVA 1 (July beige. In areas 2 and 3, mud contents on the surface 6–August 2, 2000) took place in the Angola Basin to layer reached almost 99%, in deeper layers up to study the biodiversity and ecology of the benthic 99.95%. In areas 5 and 6 about 95% mud was found. communities in extreme water depths of about 5500 m. Contents decreased towards 12 cm sediment depth down to about 90% and increased again towards deeper layers up to 98% or 99%. The sediments, especially in areas 2 and 3, contained high amounts of globularian forami- Material and methods niferans. Therefore, the total carbon (TC) contents of sediments were higher (8.7% and 8.86%) in the southern The study was carried out in the Angola Basin (areas 1, 2, and 3) than in the northern stations (areas 4, (Atlantic Ocean) (Fig. 1). The megafauna was collected 5, and 6) (8.6% and 8.01%) (for details see Kro¨ ncke and in six areas with a modified Agassiz trawl of 3.5 m width. Tu¨ rkay 2003).

Fig. 1. Map of the Angola Basin (SE Atlantic) with cruise track and working area of the ‘‘Meteor’’ cruise M48-1. Numbers refer to sampling areas. ARTICLE IN PRESS F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 205

Table 1. List of stations in the Angola Basin (SE Atlantic) where specimens of Chondrocladia were found

Area Station Begin latitude Begin longitude End latitude End longitude Depth (m)

1 Me48-319AT22 122.640S03120.150E22125.340S03121.330E 5162 1 Me48-321AT22 120.640S03123.560E22126.150S03127.040E 5162 4 Me48-337AT18 118.970S04142.720E18123.970S04144.870E 5439 4 Me48-339AT18 119.430S04142.050E18124.550S04143.850E 5443 5 Me48-343AT17 107.510S04142.820E17111.230S04145.420E 5460 6 Me48-347AT16 114.990S05126.700E16120.410S05126.790E 5433 6 Me48-349AT16 115.770S05126.490E16121.490S05126.460E 5432

The collection of Porifera from the DIVA 1 expedi- C. levii sp. nov. tion is composed of 50 specimens or, in some cases, pieces of specimens collected at depths between 5162 Type material and 5460 m. Many sponges were already broken when Holotype: Museum fu¨ r Naturkunde of Berlin no. they reached the surface because they are very delicate. ZMB Por 12629. For this reason we always consider at the beginning each Paratype 1: Museum National d’histoire Naturelle, piece as a different sample. We collected 16 specimens of Paris no. D.C.L. 3882. carnivorous sponges belonging to four species: 10 of C. Locality and habitat: Angola Basin (SE Atlantic) in levii sp. nov., one of C. vaceleti sp. nov., two of C. areas 1, 4, 5, and 6, in depths from 5162 to 5460 m in nicolae sp. nov., and three of C. cf. guiteli Topsent, 1904. white mud with globularinan foraminiferans. All live material was fixed for 48 h in 5% formalin, and afterwards in 70% ethanol. The methods followed were those of Ru¨ tzler (1978), Uriz (1978),andCristobo et al. (1993). Spicules were examined under a Leo 435VP Description scanning electron microscope (SEM). Shape and size (Fig. 2a): Sponge tulip or club shaped, Types revised in Museum National d’histoire Natur- attached to the bottom by a hollow filament of 1 mm in elle, Paris: diameter, with a short conical peduncle at the base of the body. Many tapering surface processes may get C. asigmata (Holotype) (L.B.I.M. no. D.C.L. 1401); anastomosed into each other (principally in the equa- C. dichotoma (L.B.I.M. no. D.C.L. 1402 (Holotype) torial and apical zone of the sponge of 1 mm in diameter and 1410 (Cotype)); and up to 10 mm long). Size: 55–110 mm, body C. fatimae (Holotype) (L.B.I.M. no. N.B.E. 1087); 20–25 mm 9–20 mm in diameter. Strong smell. Whitish C. gracilis (Holotype) (L.B.I.M. no. D.C.L. 1386); or light beige colour while still alive and in alcohol. C. guiteli (Holotype) (L.B.I.M. no. D.T. 1007); Consistence firm, barely compressible. Light hispid C. multichela (Holotype) (L.B.I.M. no. D.C.L. 1405); surface. C. nani (Holotype) (L.B.I.M. no. N.B.E. 1088); C. pulvinata (Holotype) (L.B.I.M. no. D.C.L. 3602); Skeleton C. scolionema (Holotype) (L.B.I.M. no. D.C.L. 3601); It consists of a central axis formed by styles which are C. verticillata (Holotype) (L.B.I.M. no. D.T. 2175); attached to the bottom by branches of transverse C. yatsui (Holotype) (L.B.I.M. no. D.T. 1568). bundles of the same spicules, in the choanosome of the sponge which radiate in order to form conical processes (Fig. 2b). Microscleres are very abundant in the sponge Results but especially in the ectosome. Order: Poecilosclerida Topsent, 1928. Spicules Family: Cladorhizidae Dendy, 1922. Twenty-five measurements per specimen of each type Genus: Chondrocladia Thomson, 1873. of spicules in light microscopy (Fig. 2c–h). Synonymy: Chondrocladia Thomson, 1873: 188. Megascleres: Styles straight or slightly curved, smooth Crinorhiza Schmidt, 1880: 83. Meliiderma Ridley fusiform. Size: 1500–2000 30–40 mm (greatest dia- and Dendy, 1887: 102. Neocladia Koltun, 1970: 193 meter). (After Hajdu and Vacelet, 2002). Microscleres: Anchorate unguiferous isochelae of Type species: C. virgata Thomson, 1873. three classes: (I) largest size, 190–212 6.2–8.7 mm, with Definition: Cladorhizidae with anchorate unguiferous five teeth at each end; (II) medium size, isochelae. 67–77 5.0–6.2 mm, with five teeth at each end; (III) ARTICLE IN PRESS 206 F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 ARTICLE IN PRESS F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 207 small size, 16.2–20.0 0.6–1.5 mm, with 9–10 teeth at Skeleton each end. It consists of axes and branches of a powerful Derivatio nominis: This species is dedicated to polyspicular fibre formed by styles. In prolongations Professor Dr. Claude Le´ vi of the Museum National of the body, the skeleton is composed of some d’histoire Naturelle of Paris, for his contribution to the polyspicular bundles of styles covered with anchorate knowledge of Chondrocladia, describing six new species. isochelae.

Remarks Spicules C. levii differs from other species of this genus in the Twenty-ﬁve measurements per specimen of each type following: C. concrescens (Schmidt, 1880), C. fatimae of spicules in light microscopy (Fig. 3c–g). Boury-Esnault and Van Beveren, 1982, and C. yatsui Megascleres: Straight, smooth styles with rounded Topsent, 1930 have sigmancistres while the new species heads and abruptly pointed end. Size: 950–2250 does not have any. C. levii does not have any sigma and 10–35 mm (greatest diameter). for this reason it differs from other species that do have Microscleres: Anchorate unguiferous isochelae of two such as C. albatrossi Tendal, 1973, C. antarctica classes: (I) largest size, 60–100 6.0–8.5 mm, with a Hentschel, 1914, C. burtoni Tendal, 1973, C. clavata strong shaft, small alae, and three short teeth rounded at Ridley and Dendy, 1982, C. crinita Ridley and Dendy, the end; (II) small size, 32–52 3.0–4.2 mm, with small 1982, C. gigantea (Hansen, 1885), C. guiteli Topsent, alae, three teeth, and pointed end. Sigmas: 1904, C. michaelsarsi Arnesen, 1920, C. multichela Le´ vi, 65–86 2.1–3.0 mm. 1964, C. pulvinata Le´ vi, 1993, and C. virgata Thomson, D. nominis: This species is dedicated to Professor 1873. It also differs from C. asigmata Le´ vi, 1974,C. Dr. Jean Vacelet of the Station Marine d’Endoume dichotoma Le´ vi, 1964, C. dura Kieschnick, 1898, C. in Marseille, for the extraordinary discovery of the gracilis Le´ vi, 1964, C. guiteli Topsent, 1904, C. nani existence of carnivorous sponges and their biology. Boury-Esnault and Van Beveren, 1982, C. ramosa Kieschnick, 1898, C. scolionema Le´ vi, 1993, C. sessilis Remarks Kieschnick, 1898, and C. stipitata Ridley and Dendy, The morphology of this species is close to C. 1886, in the number of teeth and in the size classes of scolionema from New Caledonia in their conical stalk isochelae. This can be applied to differences among shape and in their ﬁlaments (Le´ vi, 1993), but it differs these species and C. cf. guiteli and C. nicolae. clearly in the kind of spicules: C. vaceleti has two different classes (in shape and size) of anchorate C. vaceleti sp. nov. unguiferous isochelae and it has sigmas. C. scolionema has only one class of microscleres. Type material C. nicolae sp. nov. Holotype: Museum fu¨ r Naturkunde of Berlin no. ZMB Por 12630. Type material Locality and habitat: Angola Basin (SE Atlantic) in area 5 in a depth of 5460 m in white mud with Holotype: Museum fu¨ r Naturkunde of Berlin no. globularinan foraminiferans. ZMB Por 12631. Paratype 1: Museum National d’histoire Naturelle, Paris no. D.C.L. 3883. Description Locality and habitat: Angola Basin (SE Atlantic) in Shape and size (Fig. 3a and b): Sponge attached to area 5 in a depth of 5460 m in white mud with soft bottom with a stalk rich in styles oriented parallel to globularinan foraminiferans. the axis. Spherical and lightly elongated body in a distal proximal direction of 14–16 mm in diameter. Distal mass is of conical shape. Body has many characteristical Description prolongations in the equatorial zone to 25 mm in length Shape and size (Fig. 4a): Body is erect with a central, and 12 mm in diameter. Fine hispid surface. Live whitish laterally compressed axis. Secondary branches were colour, rough surface, and solid consistence. always found to be on one side of the central axis.

Fig. 2. C. levii sp. nov.: (a) habitus of the holotype; (b) choanosomal bundles of styles; (c) large and medium anchorate unguiferous isochelae; (d) end of anchorate unguiferous isochelae of the large category and head of style; (e) medium anchorate unguiferous isochelae; (f) end of anchorate unguiferous isochelae of the medium category; (g) small anchorate unguiferous isochelae; and (h) comparison among the three classes of unguiferous isochelae. ARTICLE IN PRESS 208 F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213

Fig. 3. C. vaceleti sp. nov.: (a–b) habitus of the holotype; (c) spicules SEM; (d) end of styles; (e) large anchorate unguiferous isochela; (f) small anchorate unguiferous isochela; and (g) sigmas. ARTICLE IN PRESS F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 209

Fig. 4. C. nicolae sp. nov.: (a) habitus of the holotype; (b) surface SEM; (c) transverse section; (d–e) lengthwise cut; (f–g) style; (h) anchorate unguiferous isochela; and (i) sigmas in the choanosomal skeleton. ARTICLE IN PRESS 210 F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213

Between its branches, the body is slightly curved. The Spicules largest specimen measures 47 mm in length and 2 mm in Twenty-five measurements per specimen of each type diameter. Live colour orange. Rough surface. Solid of spicules in light microscopy (Fig. 5b–f). Consistence. Megascleres: Straight styles, smooth fusiform. Size: 1500–2100 20–22 mm (greatest diameter). Skeleton Microscleres: Anchorate unguiferous isochelae of It consists of axes and branches of a powerful three classes: (I) largest size, 175–210 5.1–7.0 mm, with polyspicular fibre formed by styles (Fig. 4b–e). three teeth at each end; (II) medium size, 42–83 3.8–5.0 mm, with five teeth at each end; (III) Spicules small size, 12.5–22.0 0.9–2.1 mm, with eight teeth at Twenty-five measurements per specimen of each type each end. Sigmas: 52–66 2.0–3.5 mm. of spicules in light microscopy (Fig. 4f–i). Megascleres: Straight styles, smooth fusiform. Size: Remarks 1180–1550 18.1–29.0 mm (greatest diameter). C. cf. guiteli is close to a species described by Topsent, Microscleres: Anchorate unguiferous isochelae: 1904 in the NW of Cap Finisterre (Galice), in a depth of 37–44 2.5–6.0 mm, with three teeth at each end. 4900 m, but the first one has a third category of Sigmas: 96–105 1.8–2.5 mm. anchorate isochelae with eight teeth. The second ones D. nominis: This species is dedicated to Professor Dr. have six teeth. In the type C. guiteli, isochelae of the Nicole Boury Esnault of the Station Marine d’Endoume largest size have normally three teeth but sometimes we in Marseille, for the extraordinary discovery of the found ends with four teeth. These differences do not existence of carnivorous sponges and their biology. seem enough at this moment to describe a new species, but as the specimens are from different and distant Remarks regions they could perhaps be different species. The Habitus of C. nicolae sp. nov. is very different from morphology is also similar to C. albatrosi but in this the other three Chondrocladia of the Angola Basin. particular case the stalk gets directly into the body Orange in colour, it grows in an axis with small (Tendal, 1973) and in C. cf. guiteli we can see a small branches. Most of the Chondrocladia species are light cone at the base. On the other hand, the anchorate grey or white; only C. asigmata from Kermadec Trench unguiferous isochelae of the large category have five (Pacific Ocean) (Le´ vi, 1964) is orange, arbuscular but teeth in C. albatrosi and three in C. cf. guiteli. does not have any sigmas as is indicated in its etymology and anchorate isochelae have seven teeth; C. nicolae has Discussion and conclusions sigmas and isochelae have three teeth. The Angola basin benthic environment reflects a C. cf. guiteli Topsent, 1904 typical deep-sea feature with extremely muddy sediment, containing high amounts of foraminiferans (Kro¨ ncke Locality and habitat: Angola Basin (SE Atlantic) in and Tu¨ rkay, 2003). areas 1 and 4 in depths from 5162 to 5439 m in white The genus Chondrocladia Thomson, 1873 has a very mud with globularinan foraminiferans. wide distribution in the deep sea. Most species are known only by very few specimens. Our collection from Description the DIVA 1 expedition is unusually large with 16 Shape and size (Fig. 5a): Erect sponge attached to the specimens (13 belonging to three new species and three soft bottom by a straight stalk of polyspicular bundles belonging to an already known species). These are the of styles of 2–3 mm in diameter and of 22–39 mm long first described Chondrocladia species of the Angola from the body to the end of the stalk (broken in all Basin. The Valdivia Expedition (1898–1899) collected specimens). A body of 12–25 mm in diameter is globular samples in this area but no carnivorous sponges were to elongated in the apical sense with the axis bearing described. abundant conical processes to 20 mm in length in the This genus includes presently 28 species of which upper part of the body or in equatorial disposition. several may be synonyms or belong to an other genus as Whitish live colour, rough surface, and fragile consis- occurs with C. alaskensis Lambe, 1894 and C. pulchra tence. Lambe, 1894 that are best referred to Crambe Vosmaer, 1880 (Hajdu and Vacelet 2002). The Chondrocladia Skeleton species have smooth, slightly curved, fusiform, and large Choanosomal skeleton formed by bundles of styles in size (subtylo)styles as megascleres and anchorate iso- axial disposition with branches towards radial processes. chelae with several teeth at each end and curved shaft Microscleres were more abundant in the choanosome. and also smooth sigmas and smooth sigmancistres can ARTICLE IN PRESS F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 211

Fig. 5. C. cf. guiteli Topsent, 1904: (a) habitus; (b) spicules SEM; (c) anchorate unguiferous isochelae; (d) medium anchorate unguiferous isochela; (e) small anchorate unguiferous isochelae; and (f) sigma. ARTICLE IN PRESS 212 F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213

Table 2. Comparison between the size of spicules of four described species: Measurements of spicules are given in mm

Species Styles Isochelae I Teeth Isochelae II Teeth Isochelae. III Teeth Sigmas

C. levii 1500–2000 30–40 190–212 6.2–8.7 5 67–77 5.0–6.2 5 16.2–20.0 0.6–1.5 9–10 C. vaceleti 950–2250 10–35 60–100 6.0–8.5 3 32–52 3.0–4.2 3 65–86 2.1–3.0 C. nicolae 1180–1550 18–29 37–44 2.5–6.0 3 96–105 1.8–2.5 C. cf. guiteli 1500–2100 20–22 175–210 5.1–7.0 3 42–83 3.8–5.0 5 12.5–22.0 0.9–2.1 8 52–66 2.0–3.5

be found as microscleres. The habitus and ecology of electron microscope and Dr. Carsten Lueter from the each species are important as diagnostic characters but Museum fu¨ r Naturkunde of Berlin. Special thanks to these deep-sea sponges differ from those seen or two anonymous referees for their helpful comments. photographed in natural environments and in collected specimens. The species described here are C. levii, C. vaceleti, and C. nicolae. We found one more species, perhaps new, but References we cannot differentiate it (attending only to its skeleton components) from the described C. guiteli Topsent, 1904 Cristobo, F.J., Urgorri, V., Solorzano, M.R., Rı´ os, P., 1993. off the Galician coast (NW Iberian Peninsula). Me´ todos de recogida, estudio y conservacio´ n de las The described species of this work can be differen- colecciones de porı´ feros. International Symposium and tiated by their spicules (Table 2): C. levii does not have First World Congress on Preservation and Conservation of sigmas while the other three do have them; sigmas of C. Natural History Collections, vol. 2, pp. 277–287. cf. guiteli are 52–66 mm long, those of C. vaceleti are Hajdu, E., Vacelet, J., 2002. Family Cladorhizidae Dendy, 65–86 mm long, and in C. nicolae they are 96–105 mm 1922. In: Hooper, J.N.A., Van Soest, R.W.M. (Eds.), long. With regard to the anchorate isochelae categories, Systema Porifera. A Guide to the Classiﬁcation of Sponges. C. nicolae has only one category with three teeth at each New York, pp. 636–641. end; C. vaceleti has two categories of different isochelae Kro¨ ncke, I., Tu¨ rkay, M., 2003. Structural and functional aspects of the benthic communities in the deep Angola with three teeth, and C. levii and C. cf. guiteli have three Basin. Mar. Ecol. Prog. Ser. 260, 43–53. anchorate isochelae categories with 5, 5, and 9–10 teeth Ku¨ bler, B., Barthel, D., 1999. A carnivorous sponge Chon- and with 3, 3, and 8 teeth, respectively. drocladia gigantea (Porifera: Demospongiae: Cladorhizi- The habitus of C. nicolae (erect with an axis and dae), the giant deep-sea clubsponge from the Norvegian secondary branches) is very different to the other known Trench. Mem. Queensl. Mus. 44, 289–298. Chondrocladia species. The other three species described Lendenfeld, R.V., 1907. Die Tetraxonia. In: Fisher, G. (Ed.), in this work are tulip shaped. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expe- The three new species are from localities, which are dition auf dem Dampfer ‘‘Valdivia’’ 1898–1899, vol. 4, different from the ones of the Chondrocladia species Jena, pp. 1–374. known in the world. Le´ vi, C., 1964. Spongiaires des zones bathyale, abyssale et We found no crustacean on the surface of the hadale. Galathea Rep. 7, 63–112. Le´ vi, C., 1993. Porifera Demospongiae Spongiaires bathyaux sponges. de Nouvelle-Cale´ donie, re´ colte´ s par le ‘‘Jean Charcot’’ Campagne BIOCAL, 1985. Re´ sultats des Campagnes Musorstom. 11. Me´ m. Mus. Natl. Hist. Nat. A 158, 9–87. Acknowledgements Ru¨ tzler, K., 1978. Sponges on coral reef. In: Stoddart, D.R., Johanness, R.E. (Eds.), Coral Reefs: Research Methods, These are results of the DIVA 1 expedition (cruise vol. 5. Unesco, Paris, pp. 81–120. M48/1 of RV ‘‘Meteor’’) supported by the Deutsche Schulze, F.E., 1904. Hexactinellida. In: Fisher, G. (Ed.), Forschungsgemeinschaft and a part of project number Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expe- PGIDT01PXI20008PR sponsored by the Xunta de dition auf dem Dampfer ‘‘Valdivia’’ 1898–1899, vol. 11, Galicia, Spain. We are warmly grateful to Claude Le´ vi, Jena, pp. 1–226. Jean Vacelet and Nicole Boury-Esnault for their friend- Tendal, O.S., 1973. Sponges collected by the Swedish Deep Sea Expedition. Zool. Scr. 2, 33–38. ship and help whenever we needed them. We want to Topsent, E., 1904. Spongiaires des Ac¸ores. Re´ sult. Camp. Sci. mention specially Dr. Michael Tu¨ rkay (Senckenberg Prince Albert de Monaco 25, 1–280. Museum, Frankfurt) leader of the expedition and Prof. Urban, F., 1909. Die Calcarea. In: Fisher, G. (Ed.), Wolfgang Wa¨ gele (Ruhr-Universita¨ t Bochum). We Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expe- thank Ramiro Barreiro of the Santiago de Compostela dition auf dem Dampfer ‘‘Valdivia’’ 1898–1899, vol. 19, University for technical assistance with the scanning Jena, pp. 1–40. ARTICLE IN PRESS F.J. Cristobo et al. / Organisms, Diversity & Evolution 5 (2005) 203–213 213

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