J. Mar. Biol. Ass. U.K. 2001), 81,569^579 Printed in the United Kingdom

Revision of the genus Trachytedania Porifera: ) with a description of Trachytedania ferrolensis sp. nov. from the north-east Atlantic

F.J. Cristobo*and V. Urgorri O *Departamento de Biodiversidade e Recursos Marin¬ os, Instituto de Acuicultura, Universidade de Santiago de Compostela, 15706 Santiago de Compostela, Spain. E-mail: [email protected] OLaboratorio de Zoolox|¨a Marin¬ a, Departamento de Biolox|¨a , Universidade de Santiago de Compostela, 15706 Santiago de Compostela, Spain. E-mail: [email protected]

The genus Trachytedania Ridley, 1881 Porifera: Poecilosclerida), includes those characterized by the presence of acanthostyles. To date, only ¢ve species of the genus Trachytedania have been described of which only three actually belong to this genus. Other species described as could be considered as Trachytedania because of the presence of acanthostyles in their skeletons. Some authors propose retaining Trachytedania as a subgenus of Tedania because of its skeletal structure. In this work, a taxonomic revision and consideration of the validity of the genus is provided. The morphology and anatomy of Trachytedania ferrolensis sp. nov. from the R|¨a de Ferrol Spain, north-east Atlantic) are described in detail, and the types of the three valid species described as Trachytedania to date: Trachytedania spinata, Trachytedania patagonica and Trachytedania biraphidora are studied. A key to this genus is provided.

INTRODUCTION MATERIALS AND METHODS The position of the Family Tedaniidae within the The specimens were collected by SCUBA divers and by Poecilosclerida is still disputed. Species of the genus dredging using a naturalist benthic dredge Holme & Tedania Gray, 1867, have always been di¤cult to di¡er- McIntyre, 1984) in the sublittoral zones of R|¨a de Ferrol entiate clearly Bergquist & Fromont, 1988) and the value Spain). Material from the following Museum collections of the genus Trachytedania has been called into question at was also examined: types of Trachytedania biraphidora di¡erent times since 1881. In Ridley's 1881) original Boury-Esnault, 1973 from the Muse¨ um National description based on the species Trachytedania spinata,he d'Histoire Naturelle, Paris LBIM N8D. NBE 974); types remarks that this genus di¡ers from all the known of Trachytedania spinata Ridley BMNH: 1879.12.27.9) and species of Tedania in having three kinds of spicules, one Trachytedania patagonica Ridley & Dendy BMNH: of them being spined. Ridley & Dendy 1886) described 1887.5.2.205) from the British Museum Natural History), a new species, Trachytedania patagonica with entirely spined London Figure 1). acanthostyles. Hope 1889) described one species in- The methods followed were those of Ru« tzler 1978), correctly identi¢ed as Trachytedania T. e c h i n a t a ), because it Uriz 1978, 1986) and Cristobo et al. 1993). Spicules does not have onychaetes. Another erroneous identi¢ca- were examined under Hitachi S570 and Leo 435VP tion is that of Trachytedania arborea Ke« ller, 1891) that scanning electron microscopes SEMs). Underwater has oxeas and onychaetes but does not have acantho- photographs were taken with a Nikonos V camera and styles. Boury-Esnault 1973) describes a valid species, SB-102 £ash. The megasclere spicules were measured Trachytedania biraphidora with two size classes of using a light microscope which was adapted to a Kontron onychaetes from the Atlantic coasts of South America. Bildanalyse Videoplan, equipped with CPU, TV camera, Di¡erent authors have sometimes described some of the TV monitor and digitalizing tablet. This made it possible preceding species, attributing them to the genus Tedania, to take a large number of measurements. Microscleres while another Tedania could be included in Trachyedania were measured using SEM. because of the presence of acanthostyles in its skeleton. Owing to this confusion we are revising the genus begin- SYSTEMATICS ning with a new diagnosis after reviewing types of species described as Trachytedania, and have completed their Phylum PORIFERA Grant, 1836 descriptions. In this paper, we also revise those species of Class DEMOSPONGIAE Sollas 1885 Tedania with spined styles or acanthostyles in order to Order POECILOSCLERIDA Topsent, 1928 clarify the taxonomy of the group. Finally, in this work Suborder MYXILLINA Hajdu, Van Soest & Hooper, 1994 we describe a new species of Trachytedania T.ferrolensis) from the north-east Atlantic. Fa m i l y TEDANIIDAE Ridley & Dendy, 1886

Journal of the Marine Biological Association of the United Kingdom 2001) 570 F.J. Cristobo andV.Urgorri Revision of the genusTrachytedania Porifera)

Figure 1. Map of the world showing the sites of collection of Trachytedania species: *, T. spinata; *, T. patagonica; ~, T. biraphidora; *, T. ferrolensis; ~, T. spinostyla; &, T. in£ata; &, T. microrhaphidiophora.

Diagnosis is composed of tornotes with perpendicular and paratan- Encrusting, massive or digitate Poecilosclerida with a gential arrangement. Onychaetes are abundant, free in choanosomal skeleton predominantly plumo-reticulate or ectosome and choanosome. even plumose of monactinal and/or diactinal megascleres Spicules megascleres. Acanthostyles straight or slightly which are organized in tracts of styles, acanthostyles or curved with few short spines predominantly at the base of oxeas, enclosed within light or moderate spongin ¢bres, the spicule, 170^200 mm. Tornotes straight, smooth, or with no visible spongin, and spicules merely cemented isodiametric with apex abruptly pointed, 140^170 mm. together with collagen at their nodes. The ectosomal Onychaetes very ¢ne slightly curved, 130^15 0 mm. skeleton consists of diactinal megascleres which may be Habitat: on both valves of small Pecten,18.2m. distinguished from the choanosomal megascleres either in Distribution: Portland Bay, Chile. South-west coast of morphology or size; these spicules are tylotes or tornotes Patagonia. usually with basal spines, lying tangentially, paratangen- tially or erect on the surface. Microscleres are onychaetes. Remarks Chelae are absent. After revision of the holotype of Trachytedania spinata Ridley, 1881, we have observed the presence of acanthos- tyles on the skeleton of this species Figure 2B). Those GenusTrachytedania Ridley, 1881 nearest the base of the possess more spines on the Trachytedania spinata Ridley, 1881 head than those situated at the frontal part of the ecto- Material examined some, which are generally smooth. This fact is the reason Type, British Museum Natural History) BMNH: why di¡erent authorss question the validity of this genus 1879.12.27.9) Figure 2). Bergquist & Fromont, 1988; Desqueyroux-Faundez & Original description of the holotype in Ridley 1881:122). Soest, 1996). However, it is necessary to bear in mind the New data from the present authors' reexamination of fact that the specimen collected by Ridley 1881) was the holotype: found on both valves of Pecten and that the existing Shape and size: encrusting, laminar. 10.2Â9.1Â2 mm. specimen in the British Museum Natural History) Colour: yellowish white in spirit. BMNH: 1879.12.27.9) Figure 2A) is separated from the Surface: even, covered with a ¢ne membrane. Oscula shell of the mollusc. Probably the basal spongin plate with not detected; possibly represented by irregular depres- acanthostyles became stuck to the shell, and for this sions. Ostia not detected. reason it is possible that the acanthostyles closest to the Texture: smooth. base the spiniest) have disappeared on separation of the Skeleton: choanosomal architecture is composed of sponge from substrate to which it had been attached. On loosely aggregated acanthostyles in an isotropic reticula- the other hand this does not occur with other species of tion of three or four spicules, and transversal tracts of the genus that have real acanthostyles and not just spined acanthostyles with few spines. The choanosomal skeleton styles Figures 3^6).

Journal of the Marine Biological Association of the United Kingdom 2001) Revision of the genusTrachytedania Por ife ra) F.J. Cristobo andV.Urgorri 571

Figure 2. Trachytedania spinata Ridley, 1881; Holotype. A) habitus; B) acanthostyles at the base; C&D) transverse section of theskeleton; E)tractsofacanthostyles; F)estosomalspicules.

Trachytedania patagonica Ridley & Dendy, 1886 similar spicules are scattered throughout the ectosome, Material examined which together with some of the stylote spicules, form a Holotype, British Museum Natural History) BMNH: very sparse and irregular dermal reticulation. 1887.5.2.205) Figure 3A,B). Spicules: a) Megascleres. Acanthostyles rather stout Original description of the holotype in Ridley & slightly curved, spined all over with the spination most Dendy, 1886: 336; 1887: 57. marked at the base; size: 350Â12.5 mm. Tornotes, tapering Shape and size: irregularly shaped, massive, 37 mm. gradually to a slightly hastate point at each end; size: Colour: pale yellow in spirit. 245Â7 mm. b) Microscleres. Onychaetes very ¢ne slightly Surface: uneven with slight traces of hispidation. thicker at one end than at the other; length: 200 mm Oscula small, scattered. Ostia scattered throughout some spined near the broader end with ¢ne spines directed parts of the dermal membrane. towards that end of the spicule. Texture: soft and crumbling. Habitat: bottom blue mud, 320 m. Skeleton: very loose and irregular, consisting of a Distribution: south-west coast of Patagonia. somewhat isodictyal reticulation of acanthostyles but with a small admixture of tornotes. No distinct ¢bre. Remarks Immediately beneath the dermal membrane there are a The slides of the holotype re-examined correpond to number of irregular, divergent tufts of tornotes while Trachytedania patagonica but a subsample from a wet

Journal of the Marine Biological Association of the United Kingdom 2001) 572 F.J. Cristobo andV.Urgorri Revision of the genusTrachytedania Porifera)

Figure 3. A&B) acanthostyles of Trachytedania patagonica Holotype); C&D) acanthostyles of Trachytedania biraphidora Holotype); E&F) Trachytedania biraphidora Holotype), tracts of acanthostyles in the choanosomal skeleton. specimen corresponds to another species Iophon sp.). For tyles and tylotes joined by less dense lines. Onychaetes this reason there are no new data from the present sometimes in bouquets. authors' re-examination of the holotype. Spicules: acanthostyles slightly curved although they may sometimes be found with a few spines 5^12) only at the base; there may also be some isolated spines, but always Trachytedania biraphidora Boury-Esnault, 1973 near the basal zone. Size: 190^280 mmÂ2.2^6 mm. Tylotes Material examined are straight, isodiametric, completely smooth with elip- Holotype, Muse¨ um National d'Histoire Naturelle, Paris soidal ends sometimes di¡erent one from another without LBIM N8D. NBE 974) Figure 3C^F). spines although they sometimes have a small mucronated Original description of the holotype in Boury-Esnault, end. Size: 172^210 mmÂ1.2^3 mm. Onychaetes clearly 1973: 281. di¡erentiated into two size classes: 38^62 mmÂ0.9 mmand Shape and size: massive 10 cm tall. 102^174 mmÂ1.1 mm. The bigggest ones are straight or Colour: yellowish. slightly curved and have pointed tips. The smallest ones are Surface: mamelon shaped. curved, slightly sinuous or straight, generally with one of Skeleton: polygonal network of acanthostyles in the the ends more rounded and the other pointed. choanosome and tylotes on the surface; inside the sponge Habitat: 48 m. we can di¡erentiate the main primary tracts of acanthos- Distribution: south-west Atlantic.

Journal of the Marine Biological Association of the United Kingdom 2001) Revision of the genusTrachytedania Porifera) F.J. Cristobo andV.Urgorri 573

Figure 4. Trachytedania ferrolensis sp. nov.: A) living specimen in situ Paratype 1); yellowish colour, photograph F.J. Cristobo). B^D) SEM micrographs showing the arrangement of the skeleton. E&F) Tracts of acanthostyles in the choanosomal skeleton.

Remarks Trachytedania biraphidora while in Trachytedania patagonica Sometimes young very ¢ne acanthostyles may eassily there is only one size class. be confused with onychaetes. The size of spicules remeasured are slightly di¡erent from those indicated by Boury-Esnault, 1973. Trachytedania ferrolensis sp. nov. After a revision of the holostyles, we do not agree with Type material Desqueyroux-Faundez & Soest 1996: 68) who categorize Holotype: Museo Nacional de Ciencias Naturales of Trachytedania patagonica as synononymous with Trachytedania Madrid no. MNCN 1.01/184, from A Millonaria, R|¨ade biraphidora. The acanthostyles are clearly di¡erent, as we Ferrol, Galicia, Spain; 43828013 00N08815 00900W; 22 m, 14 can see in Figure 3; the ¢rst possesses tornotes and the March 2000. On coal slags and Balanus. Paratype 1: second tylotes; the onychaetes are of two size classes in Muse¨ um National d'Histoire Naturelle, Paris, no. MNHN

Journal of the Marine Biological Association of the United Kingdom 2001) 574 F.J. Cristobo andV.Urgorri Revision of the genusTrachytedania Porifera)

Galicia, Spain; 4382802500N08815 00000; 15 m, 24 November 1999. On the right valve lower) of Chlamis varia. Para- type 5: Laboratorio de Zoolox|¨a Marin¬ a, Universidade de Santiago de Compostela, from R|¨a de Ferrol, Galicia, Spain; 4382802600N08814 04700; 10 m, 21 January 1988. Para- type 6: Laboratorio de Zoolox|¨a Marin¬ a, Universidade de Santiago de Compostela; A Millonaria, R|¨a de Ferrol, Galicia, Spain; 43828013 00N08815 00900, 22 m, 14 March 2000. On both valves of Chlamis varia. Paratype 7: Labor- atorio de Zoolox|¨a Marin¬ a, Universidade de Santiago de Compostela; A Millonaria, R|¨a de Ferrol, Galicia, Spain; 43828013 00N08815 00900, 22 m, 14 March 2000. On the left valve upper) of Chlamis varia. Derivatio nominis. The name of this species derives from the toponymy of the type locality: R|¨a de Ferrol. This is a token of recognition of a ria with exceptional ecological characteristics that has unfortunately su¡ered from the adverse e¡ects of human behaviour.

Diagnosis The living sponge is a yellow encrusting thin sheet-like coating of the substrate. The choanosomal skeleton consists of acanthostyles of two size classes arranged with their heads ¢xed to a basal plate of spongin forming point directed outward columns. Right and thin ectosomal tylotes in palisade or few de¢ned bouquets. Onychaetes abundant of two size classes.

Description Shape and size: the sponge is encrusting, forming a sheet-like coating of rocky substrates and Balanus.Patches 530 cm in diameter and 3 mm thick have been seen. The sponge is ¢rmly attached to the substratum by its whole undersurface Figure 4A). Colour: the in situ colour is bright yellow. After ¢xation in formalin, the alcohol-preserved specimens are white, cream or light pink. It may give the alcohol a slightly yellowish tint. The surface is smooth but irregular as the sponge closely follows the irregularities of the substratum. Ostia scattered fairly evenly all over. Single circular oscules of 41mm in diameter with small super¢cial canals. This aquiferous system that characterizes the sponge surface in the ¢eld is no longer detectable after collection. Texture: smooth. The consistency is quite cartilaginous, Figure 5. Camera lucida drawings of Trachytedania ferrolensis although easy to tear. sp. nov. spicules. A) Primary acanthostyle; B) secondary Skeleton: the principal skeleton consists of acantho- acanthostyle; C) tylote; D) large onychaete; E) small styles of two size classes arranged with their heads ¢xed onychaete. to a basal plate of spongin forming pointed outward directed columns of 4^5 spicules. Right and thin ecto- DCL 3667 from A Millonaria, R|¨a de Ferrol, Galicia, somal tylotes in palisade or few de¢ned bouquets. Spain; 43828013 00N08815 00900W; 22 m, 14 March 2000. On Onychaetes of two size classes more abundant in the coal slags and Balanus. Paratype 2: Museo de Historia ectosome Figure 4B^D&F). Natural Luis Iglesias Universidade de Santiago de Spicules Figures 5 & 6). Acanthostyles of two size Compostela no. 10005; from A Millonaria, R|¨a de Ferrol, classes: primary acanthostyles measure 167^261 mmin Galicia, Spain; 4382802500N08815 00000; 15m, 30 August length. They are straight or slightly curved with identical 1999. On the right valve lower) of Chlamis varia. Para- heads; scattered spines extend along the shaft but are type 3: Laboratorio de Zoolox|¨a Marin¬ a, Universidade much less frequent in the distal half and disappear in the de Santiago de Compostela; from A Millonaria, R|¨ade last quarter. Secondary acanthostyles measure 67^109 mm Ferrol, Galicia, Spain; 4382802500N08815 0000;15m,30 in length, almost always straight, conspicuously spined August 1999. On both valves of Chlamis varia. Paratype 4: along the whole length. The spines on the head are fairly Laboratorio de Zoolox|¨a Marin¬ a, Universidade de long, sometimes slightly recurved ending abruptly, Santiago de Compostela; A Millonaria, R|¨a de Ferrol, sometimes deformed. Overall size: 67- 160)-261 mmby

Journal of the Marine Biological Association of the United Kingdom 2001) Revision of the genusTrachytedania Porifera) F.J. Cristobo andV.Urgorri 575 , 4 ^ 1 2 ^ 6 Â ^ Trachytedania 0.5 4 180 Â Â ^ 180 220 ^ ^ Uni-plurispicular reticulate ¢bres of acanthostyles. Tylotes on surface Brown Dry) 6 Ty: 160 Trachytedania microphaphidophora ^ 4 13 m 4 ^ ^ 6.5 160 Â ^ 4 280 2202 Â ^ ^ 143 100 260 72 ^ ^ ^ acanthostyles. Pali- sade of tylotes Spt) Ty: 169 , Ridley 122); 1881: Burton 1932: 306). ,Cristobo 1997:197). 12 190 3mm Notseen ^ ^ 7 6 Ty: 230 Â Â 390 194 m On algae: Int 81 35 ^ ^ ^ Trachytedania spinata Loose reticulation Plumose tracts of Trachytedania ferrolensis 1978: 59). ¨ 4.0 To: 320 ^ , Sara 34 m 28 1.6 10.4 228 ^ ^ Â 45 220 Â ^ 205 115 261 117 50 ^ ^ 109 ^ ^ Basal lamina of spongin. Columns of acanthostyles. Tylotes in palisade 67 rcyeai in£ata Trachytedania , Boury-Esnault 1973: 281). 3 Ty: 148 6167 Â ^ 3 278 Â ^ 260 148 322 143 39 ^ ^ ^ Primary tracts of acanthostyles and tylotes on surface Trachytedania biraphidora , Bergquist & Fromont 1988: 59). 7 Ty: 228 Â 12.5 265 Â species. Massive Massive Encrusting Subspherical Encrusting Massive Loose, irregular reticulation of acanthostyles with tufts of tornotes 7 To: 245 m. L, large; S, small; To, tornote; tylote; Ty, Spt, in spirit; Int, intertidal. Sources: ^ m 5 mm Small, scattered Not seen Circular: 1 mm Not seen Not seen Dispersed: 1 mm 4 207 m Mud: 320 m 48 m On stones: 10 ^ 2 200 220 rcyeai spinostylota Trachytedania ^ 11 350 Â ^ ^ .18 Trachytedania 1.2 05 240 ^ Â Â 28 Pecten 210 ^ 6 acanthostyles. ^ ^ 3 branching or massive vertical ¢bre of Tornotes on surface T.spinata T.patagonica T.biraphidora T.ferrolensis T.microrhaphidophora T.spinostylota T.in£ata Comparison between , Ridley & Dendy 1886: 336); Ridley & 57). Dendy 1887: OsculesPores Irregular: 0.2 Not found Scattered Not seen Scattered Circular: 1 Shape Encrusting, £abellate, Colour Yellowish Spt) Pale yellow Spt) Yellowish Yellow Yellowish-white Spt) Red. Creamy-white SurfaceTextureHabitat Glabrous Pitted On Uneven Soft. Crumbling Mameloned Uneven Uneven Firm Di¡. Distinguish Glabrous Elastic Rough DistributionMeasurements of spicules are given in South-west Atlantic South-west Patagonia South-west Atlantic North-east Atlantic Sea of Japan New Zealand Patagonia patagonica Burton 70); 1935: Koltun 156). 1959: Skeleton With basal lamina and Onychaetes S 62 Onychaetes L 52 Tylote/Tornote S Tylote/Tornote L To: 180 Acanthostyles S Acanthostyles L 165 Table 1.

Journal of the Marine Biological Association of the United Kingdom 2001) 576 F.J. Cristobo andV.Urgorri Revision of the genusTrachytedania Porifera)

Figure 6. Scanning electron micrographs of Trachytedania ferrolensis sp. nov. spicules. A) Head of a primary acanthostyle; B) head of a secondary acanthostyle; C) head of a tylote; D) large and small onychaetes and a tylote; E) small onychaete; F) end of a large onychaete; G) end of a small onychaete.

4.5- 7)-10.5 mm. The tylotes are straight, thin, with analysis of sediment reveals the following contents: sand strongylote to subtylote and unequal ends. Size: 148- mid^high 64.62%), pelitic low 5.56%); organic mate- 187)-220 mmby1.6- 3)-4mm. The onychaetes are very rial 2.85%, and nitrogen 0.15%; carbonates low 33.66% thin, straight or slightly curved, strongly microspined Parapar, 1991). This type of muddy bottom is poor in with spines always bent toward one of the ends, two size sponges although there are fragments of shells and small classes: 39^117 mmand148^205 mm. Small onychaetes stones which form a peculiar habitat for a ¢xation base with the end blunt instead of pointed as in the bigger for some species. In spite of the presence of Trachytedania forms Figure 6F,G). Size altogether: 39.52- 116)- ferrolensis, its quantitative importance is exiguous. 205 mmÂ0.3- 1)-2 mm. Locality and habitat: holotype was found in A Millonaria, R|¨a de Ferrol, Galicia, Spain; 4382802500N Typelocality 08815 00000;10^22 m. Sandy and muddy bottoms with Dredging station no. 36 Cristobo, 1997), is in front of gravels and coal slags. In areas of sedimentary tubes of Concepcio¨ n Arenal key, R|¨a de Ferrol, Spain 4382802600N polychaetes Chaetopterus variopedatus Renier, 1804) often 08814 04700) 10 m, it has a muddy bottom with gravel and occur and Diopatra neapolitana Delle Chiaje, 1841. The coal slags.

Journal of the Marine Biological Association of the United Kingdom 2001) Revision of the genusTrachytedania Porifera) F.J. Cristobo andV.Urgorri 577

DISCUSSION ¢gure 52); Tedania fuegiensis Thiele, 1905: ¢gure 53); Tedania murdochi Topsent, 1913: 629); Tedania toxicalis The systematic position of the genusTrachytedania Ridley, Laubenfels, 1932: 85) and Tedania gurjanovae Koltun, 1881 Porifera, Poecilosclerida) has already been discussed 1959: ¢gure 113); ii) Tedania coulmani has acanthostyles at length. It was proposed for classi¢cation in the Tedania but does not have onychaetes Kirkpatrick, 1907: 280; species, in which the styles are partly or entirely spined. 1908: 33); and iii) Tedania bispinata has a kind of Whether this characteristic is in itself su¤cient to sepa- strongyles with microspined bases, and these structural rate the two genera was questioned by some authors after macroscleres di¡er clearly from acanthostyles Hentschel, the ¢rst genus description Ridley & Dendy, 1887; 1911: ¢gure 26). Topsent, 1892; Topsent, 1928; Topsent, 1929). Currently for Another incorrect identi¢cation is the presence of some authors Trachytedania is a synonym of Tedania Gray, Trachytedania spinata in the Adriatic Sea Szymanski, 1867, in view of the occasional occurrence of spining in 1904; Vatova, 1928). Unfortunately these authors neither the styles of di¡erent specimens Bergquist & Fromont, describe nor illustrate the specimen. Anyhow, if it is a 1988), while others propose retaining Trachytedania as a Trachytedania species, it must be di¡erent from the subgenus of Tedania Desqueyroux-Faundez & Soest, 1996) Trachytedania spinata from south-west Atlantic. because the ectosomic spicules di¡er rather signi¢cantly In our opinion Trachytedania is a valid genus. Occur- from Tedania anhelans in being mucronate-oxeote, without rence of acanthostyles is not sporadic or rare in the spination at the apices, the skeletal structure is very loose Trachytedania species and choanosomal megascleres are not and no distinct meshes are recognized. only a few styles with spines but true acanthostyles as we To date, only ¢ve species of the genus Trachytedania can see in Figure 2B. Ridley's 1881) type is encrusting, Ridley, 1881, have been described: Trachytedania spinata laminar on both valves of Pecten but the specimen is now Ridley, 1881; Trachytedania patagonica Ridley & Dendy, detached from the shell and it is possible that most of the 1886; Trachytedania echinata Hope, 1889; Trachytedania arborea acanthostyles were missing from the basal lamina. Ke« ller, 1891; and Trachytedania biraphidora Boury-Esnault, According to Burton 1932), the signi¢cance of these 1973, of which only three belong to this genus and two spined spicules must be appreciated. In sections taken at have been erroneously identi¢ed: Hope 1889: 338) has right angles to the substratum it was found that the incorrectly identi¢ed his Trachytedania echinata because it acanthostyles at the base of the vertical columns are the has two size classes of acanthostyles and tylotes but does most conspicuously spined and those towards the surface not have onychaetes, therefore it might also be a species are hardly spined at all. Moreover, small acanthostyles of Stylopus but not a species of Trachytedania; another case found in the embryos of his specimens are very occasionally is Trachytedania arborea Ke« ller, 1891: 314) that has oxeas, associated with the normal basally spined styles set on the styles and onychaetes but does not have acanthostyles, substratum. It seems probable therefore that the holotype of therefore it may be a Tedania but not a Trachytedania.A this species represents nothing more than the immediate survey of the literature of those species described as post-¢xation form of the species; that there is retained in Tedania with spined styles or acanthostyles, shows that the adult a tendency to form spines on the styles, and that some of them could be included in Trachytedania: Tedania this tendency, although almost wholly suppressed in the microrhaphidophora Burton, 1935, Tedania in£ata Sara¨ ,1978 adult, is still occasionally found in spicules. and Tedania spinostyla Bergquist & Fromont, 1988, giving, The synapomorphic value of acanthostyles is a wide- with Trachytedania ferrolensis, a total of seven valid species spread characteristic which leads to con£icting inter- belonging to Trachytedania, and others which cannot be pretations. They do not occur in the whole order of included:Tedania coulmani Kirkpatrick,1907;Tedania bispinata Poecilosclerida, so that postulating symplesiomorphic Hentschel, 1911; Tedania murdochi Topsent, 1913; Tedania status on the one hand implies accepting their loss higher toxicalis Laubenfels, 1930; Tedania gurjanovae Koltun, 1958; in the tree. Another possible interpretation is that and Tedania mucosa Thiele, 1905 synonymyzed with Poecilosclerida without acanthostyles are primitive and Tedania pectinicola Thiele, 1905; Tedania fuegiensis Thiele, that the latter developed subsequently Hajdu et al., 1994). 1905; and Tedania excavata Thiele, 1905 by Desqueyroux- In our opinion the presence of true acanthostyles is Faundez & Soest, 1996). Tedania microrhaphidophora as we su¤cient to uphold the separate categorization of the can see in Burton 1935, ¢gure 1) and in Koltun 1959, genus. There is, however, a further di¡erence from species ¢gure 114), has slightly curved acanthostyles with sparse of Tedania:alltheTrachytedania species revised have spines usually found only at the base, dermal tornota ectosomal spicules without spination on the apices. Type and onychaetes raphides). Tedania in£ata Sara¨ ,1978, species of the genus Tedania Tedania anhelans)havetylotes ¢gure 37) has acanthostyles with spines only at the with well developed microspined heads. base, and it also has tylotes and onychaetes. Tedania Categorization of Trachytedania patagonica Ridley & spinostyla has typical acanthostyles with spines on its Dendy and Trachytedania biraphidora Boury-Esnault as the head and spining which extends a short distance down same Desqueyroux-Faundez & Soest, 1996: 68) is an the shaft and tylotes and onychaetes Bergquist & error. The di¡erences are su¤cient for categorization as Fromont, 1988: ¢gure 26a). Spicule dimensions and the di¡erent species. The ¢rst one has spined acanthostyles all main characteristics of the three species are shown in over except at the extreme apex, tornotes with hastate Ta b l e 1. points and onychaetes of one size class. The second has On the other hand, the reasons for placing the other slightly curved acanthostyles with marked spination only species outside Trachytedania are the following: i) most of at the head, tylotes and onychaetes of two size classes. the species do not have acanthostyles: Tedania excavata Trachytedania ferrolensis di¡ers from all otherTrachytedania Thiele, 1905: ¢gure 51); Tedania pectinicola Thiele, 1905: species because of the presence of two size classes of

Journal of the Marine Biological Association of the United Kingdom 2001) 578 F.J. Cristobo andV.Urgorri Revision of the genusTrachytedania Porifera) acanthostyles. Moreover it di¡ers fromTrachytedania spinata, Cristobo,F.J.,Urgorri,V.,Solorzano,M.R.&R|¨ os, P., 1993. Trachytedania patagonica,Trachytedania microrhaphidophora, and Me¨ todos de recogida, estudio y conservacio¨ n de las Trachytedania in£ata by the presence of two size classes of colecciones de por|¨ feros. International Symposium & First World onychaetes while these species have only one size class. Its Congress on Preservation and Conservation of Natural History form, encrusting, and its skeleton with basal lamina of Collections, 2,277^287. Desqueyroux-Fau¨ ndez, R. & Soest, R.W.M. van, 1996. A review spongin, columns of acanthostyles and ectosomal diactines of Iophonidae, Myxillidae and Tedaniidae occurring in the is similar to that of other species such Trachytedania South East Paci¢c Porifera: Poecilosclerida). Revue Suisse spinata and Trachytedania spinostylota and di¡ers from Zoologie, 103,3^79. Trachytedania patagonica massive), Trachytedania biraphidora Hajdu, E., Soest, R.W.M. van & Hooper, J.N.A., 1994. Proposal massive), Trachytedania microrhaphidophora subspherical) for a phylogenetic subordinal classi¢cation of poecilosclerid and Trachytedania in£ata massive). With regard to distribu- sponges. In Sponges in time and space ed. R.W.M. van Soest tion, most Trachytedania species are from the south-west et al.), pp. 141^150. Rotterdam: A.A. Balkema. Atlantic while Trachytedania microrhaphidophora is from the Hentschel, E., 1911. Tetraxonida 2. In Die Fauna Su« dwest- Sea of Japan, Trachytedania spinostylota from New Zealand, Australiens ed. W. Michaelsen and R. Hartmeyer), pp. 342^ and Trachytedania ferrolensis from the north-east Atlantic. 393. Jena: Gustav Fischer. Holme, N. & McIntyre, A., 1984. Methods for the study of meio- benthos. Oxford: Blackwell Scienti¢c Publication. Key to the species of Trachytedania Hope, R., 1889. On two new British species of sponges, with short notices of an ovigerous specimen of Hymeniacidon 1. Onychaetes of two size classes ...... 2 dujardinii Bowk., and of a fossil Toxite. Annals and Magazine of ö Onychaetes of one size class ...... 4 Natural History, 4 23), 333^342. 2. Acanthostyles of two size classes...... T.ferrolensis Ke« ller, C., 1891. Die Spongienfauna des rothen Meeres. ö Acanthostyles of one size class ...... 3 Zeitschrift fu« r Wissenschaftliche Zoologie, 52,294^368. 3. Tylotes of two size classes. Straight acanthostyles with Kirkpatrick, R., 1907. Preliminary report on the Mono- spines on the head and shaft ...... T.spinostyla axinellida of the National Antarctic Expedition. Annals and ö Tylotes of one size class. Slightly curved acanthostyles Magazine of Natural History, 20,271^291. with spines only at the head...... T.biraphidora Kirkpatrick, R., 1908. Porifera. Tetraxonida. In National Antarctic Expedition 1901 1904 Natural History),pp.156. London: 4. Tylotes present...... T.in£ata ^ ^ British Museum Natural History. ö Tornotes present ...... 5 Koltun, V.M., 1959. Corneosiliceous sponges of the northern and 5. Onychaetes 515 0 mm ...... T.microrhaphidophora far eastern seas of the USSR. In Keys for the identi¢cation of the ö Presence of onychaetes 415 0 mm and acanthostyles fauna of the USSR, vol. 67 ed. Zoological Institute of the 5300 mm ...... T.spinata Academy of Sciences of the USSR), pp. 1^236. Moscow: ö Presence of onychaetes 415 0 mm and acanthostyles Zoological Institute Akademia Nauk CCCP. 5300 mm ...... T.patagonica Laubenfels, M.W., 1932. The marine and fresh-water sponges of California. Proceedings of the United States Natural History Museum, 81,1^140. ¨ ¨ ¨ We are particularly grateful to Claude Le¨ vi of Muse¨ um Parapar, J., 1991. Anelidos poliquetos bentonicos de la R|adeFerrol National d'Histoire Naturelle of Paris for giving us access to type Galicia). PhD thesis, Universidad de Santiago de collections and his many positive comments, to Nicole Boury- Compostela. Servicio de Publicaciones e Intercambio Esnault for critical advice on earlier drafts of this paper and Cient|¨ ¢co Universidade de Santiago de Compostela. Chantal Be¨ zac for her competent technical assistance in scanning Ridley, S.O., 1881. Spongida. In Account of the zoological collections electron microscopy, both work at the Station Marine d'En- made during the survey of HMS Alert in the Straits of Magellan and doume, Marseille. To Claire Valentine of the British Museum the coast of Patagonia. Proceedings of the Zoological Society of London, Natural History), London for giving access to type collections. I 107^137, 14 0^141. also thank Alfonso Naveiro and Xaqu|¨ nLo¨ pez Serantes for Ridley, S.O. & Dendy, A., 1886. Preliminary report on the providing material for examination. This research is part of pro- Monaxonida collected by HMS Challenger. Annals and ject number XUGA20005B95 sponsored by the Xunta de Magazine of Natural History , 18,325^351, 470^493. Galicia and the contribution no. 003/2000 of the Instituto de Ridley, S.O. & Dendy, A., 1887. Report on the Monaxonida Acuicultura, Universidade de Santiago de Compostela. collected by HMS `Challenger' during the years 1873^1876. Report. Scienti¢c Results Challenger Zoology, 20,1^275. Ru« tzler, K., 1978. Sponges on coral reef. In Coral reefs: research REFERENCES methods, vol. 5 ed. D.R. Stoddart and R.E. Johanness), pp. Bergquist, P.R. & Fromont, J., 1988. The marine fauna of New 299^313. Paris: Unesco. Zealand: Porifera, Demospongiae, Part 4 Poecilosclerida). Sara¨ , M., 1978. Demospongie di acque super¢ciali della Terra Memoirs. New Zealand Oceanographic Institute, 96,1^197. del Fuoco. 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Topsent, E., 1928. Spongiaires de l'Atlantique et de la Va¨ tova, A., 1928. Compendio della Flora e Fauna del Mare Me¨ diterrane¨ e, provenant des croisie©res du Prince Albert Ier Adriatico presso Rovigno. Istituto di Biologia Marina per de Monaco. Re¨ sultats des Campagnes Scienti¢ques Prince Albert I l'Adriatico Rovigno, 14,107^128. Monaco, 74,278^280. Topsent, E., 1929. Remarques sur Tedania spinata. Revista Chilena de Historia Natural, 33, 282^287. Uriz, M.J., 1978. Contribucio¨ n a la fauna de esponjas Demospongia)de Catalunya.Tesis Universidad de Barcelona. Submitted 19 September 2000. Accepted 3 July 2001.

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