Revision of Calycosoma Schulze, 1899 and finding of Lophocalyx Schulze, 1887 (six new species) in the Atlantic Ocean (Hexactinellida, Rossellidae)

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1 Porifera Research: Biodiversity, Innovation and Sustainability Revision of Calycosoma Schulze, 1899 and finding of Lophocalyx Schulze, 1887 (six new species) in the Atlantic Ocean (Hexactinellida, Rossellidae) Larisa L. Menshenina (1), Konstantin R. Tabachnick (2), Daniela A. Lopes (3), Eduardo Hajdu (3) (1) Biophysical Department, Physical Faculty, MSU 2, b.2, Moscow State University, Moscow, , Russia (2) P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nahimovsky pr. 36, , Moscow, Russia. tabachnick@mail.ru (3) Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s/n, , Rio de Janeiro, RJ, Brazil. hajdu@acd.ufrj.br Abstract: New data are brought on the taxonomy of Calycosoma and Lophocalyx, two closely related rossellid genera. A specimen from the Antarctic, previously identified as Calycosoma validum, is transferred here to Lophocalyx sp. Six new species of Lophocalyx are described, found in different regions of the Atlantic Ocean, between 500 and 2860 m depth. The diagnoses of these two genera are revised, and their likely affinities are discussed. Keywords: New species, Hexactinellida, Atlantic Ocean, taxonomy Introduction The monotypic genus Calycosoma was described by Schulze (1899) on the basis of a specimen collected off the East Coast of North America (C. validum Schulze, 1899). Schulze placed it within Asconematidae. Later, however, Ijima (1903, 1904, 1927) transferred it into Rossellidae-Lanuginellinae. Schulze (1903) described another species of Calycosoma, C. gracile Schulze, 1903 off the Indonesian Archipelago. Ijima (1927) moved it to Sympagella Schmidt, Later, when Caulophacidae was abolished and synonymized with Rossellidae, Sympagella too was classified within subfamily Lanuginellinae (Tabachnick 1999, 2002). Topsent (1910, 1913) reported on C. validum finding in the Antarctic Ocean. This report was cited by Ijima (1927) and followed by Barthel and Tendal (1994), but analysis of Topsent s data was not made, in spite of his description of anchorate pentactines in the Antarctic specimen, spicules which are absent from the holotype. The examination of sponges collected by Brazilian, French, Norwegian, Russian and United States oceanographic expeditions in the Atlantic Ocean showed that Lophocalyx is widely represented in this ocean, with forms which are close to Calycosoma but for the single feature respectively, presence or absence of anchorate pentactines. The description of these materials is given below. Materials and methods The materials studied here were collected by several oceanographic expeditions. Figure 1 shows the collecting sites in the Atlantic Ocean, where all the material studied here has been gathered. The methods employed were the usual ones for preparation microscope slides of dissociated spicules and thick sections, as described elsewhere (Janussen et al. 2004, Lopes et al. 2005), with occasional substitution of nitric acid or sulfochromic solution. Abbreviations used here are: MZUB - Museum of Zoology - University of Bergen (Norway), BMNH - Natural History Museum (London), IORAS - Institute of Oceanology of Russian Academy of Sciences (Moscow), MNHN - Muséum national d Histoire naturelle (Paris), MNRJ - Museu Nacional - Universidade Federal do Rio de Janeiro (Rio de Janeiro), NMS - National Museum of Scotland (Edinbourg), REVIZEE (Evaluation of the Sustainable Potential of Life Resources in the Economic Exclusive Zone), USNM - National Museum of Natural History - Smithsonian Institution (Washington D.C.). Calycosoma Schulze, 1899 Synonymy: Calycosoma Schulze 1899: 27. Not C. gracile Schulze 1903: 14 and not C. validum from Antarctic Topsent 1910: 17; 1913: 606; Barthel and Tendal 1994: 111. Type species: Calycosoma validum Schulze, 1899 (by monotypy) Definition: Basiphytous, often pedunculate Lanuginellinae with dermalia consisting of pinular pentactines and rare hexactines; hypodermalia are of one type (orthotropal pentactines); with prostalia lateralia consisting of diactines;

2 450 Fig. 1: Distribution of Calycosoma and Atlantic species of Lophocalyx. A. C. validum. B. L. pseudovalida sp. nov. C. L. biogasi sp. nov. D. L. oregoni sp. nov. E. L. brasiliensis sp. nov. F. L. atlantiensis sp. nov. G. L. reiswigi sp. nov. oxyoidal microscleres and strobiloplumicomes (slightly modified from Tabachnick 2002). Remarks: An emphasis has been put on the lack of a second category of hypodermal pentactines. As argued below, this is the main feature separating Calycosoma from Lophocalyx. Calycosoma validum Schulze, 1899 (Fig. 1; Table 1) Synonymy: Calycosoma validum Schulze 1899: 27. Not C. validum from Antarctic - Topsent 1910: 17; 1913: 606; Barthel and Tendal 1994: 111. Material examined: Holotype. USNM USFCS Albatross, stn. 2573, 40 o N 66 o W (eastern North American continental slope, N of the New England seamounts), 3186 m. A fragment of holotype. BMNH Other materials. USNM (kt3005; kt522.1; kt522.2; kt523; kt524.1; kt527.1; kt528; kt529.1; kt529.2; kt531; kt533; kt534; kt535.1; kt535.2; kt536; kt537) - USFCS Albatross, stn. 2117, 15 o N 63 o W (Aves Ridge, W of the Lesser Antilles), 1250 m. USNM (kt1123; kt1124; kt1126; kt1128; kt1130; kt1131; kt1132; kt1133; kt1134) - USFCS Albatross, stn. 2126, 13 o N 70 o W, 3111 m. USNM (kt135; kt200; kt202; kt204; kt205.1; kt205.2) - USFCS Albatross, stn. 2127, 19 o N 75 o W, 2997 m. USNM (kt1405; kt1406; kt1407; kt1409; kt1420; kt1421) - R.V. Bartlett , stn. 58, 13 o N 67 o W (N of the Netherlands Antilles), 5008 m. USNM (kt1306) - R.V. Bartlett , stn. 90, 13 o 27.4 N 64 o 43.9 W (N Table 1: Some measures of spicules of Calycosoma validum Schulze, 1899 (in mm). USNM holotype USNM (kt1306) USNM (kt1123) USNM (kt1458) n avg min max std n avg min max std n avg min max std n avg min max std L dermal pentactine distal ray L dermal pentactine tangential ray L atrial pentactine proximal ray L atrial pentactine tangential ray D oxyhemihexaster and oxyhexaster d oxyhemihexaster and oxyhexaster D oxyhexactine D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette.

3 451 of Isla Margarita), m. USNM (kt1458; kt1460) - R.V. Alaminos, stn.68 A3-5B (Gulf of Mexico), 3840 m. USNM (kt 328) - R.V. Oregon, stn. 2199, 24 o 42 N 92 o 18 W, 3658 m. IORAS 5/2/414; 5/2/415 - R.V. Akademik Mstislav Keldysh - 1, stn. 54, 21 o 04.5 N 82 o 27.6 W (S of Cuba), m. We do not present the description of these specimens as they are given in the Family Rossellidae chapter of the Systema Porifera (Tabachnick 2002). Only a table (Table 1) with spicule micrometric data for several specimens is given below. Lophocalyx Schulze, 1887 Synonymy: Lophocalyx Schulze 1887: 514. Part of Rossella - R. philippinensis Gray 1872: 137; 1873a: 234; 1873b: 361; Carter 1875: 118; Marshall and Meyer 1877: 261. Polylophus (preoccupied name) Schulze 1885 (nomen nudum): 451; 1886: 47: 1887: 132. Calycosoma (part) - C. validum Topsent 1910: 17; 1913: 606; Barthel and Tendal 1994: 111. Type species: Rossella philippinensis Gray, 1872 (by monotypy) Definition: Lophophytous or basiphytous Lanuginellinae with hypodermal pentactines of two types: with normal, straight tangential rays and with short-toothed anchorate rays, microscleres with discoidal outer ends are rare microdiscohexasters (modified from Tabachnick 2002). Remarks: The definition of Tabachnick (2002) has been slightly modified for greater clarity. It is unclear how to classify the large hexactines seen in Lophocalyx spp.: choanosomal or hypoatrial. The radial rays in these spicules are different; the distal ray is pinular or carries some spines, while the tangential rays are similar in size and shape to those in hypodermal pentactines. Such spicules are known from several genera of Rossellidae. Lophocalyx pseudovalida sp. nov. (Fig. 1, 2, 3; Table 2) Etymology: The species is named pseudovalida because of its resemblance to Calycosoma validum. Material examined: Holotype - IORAS 5/2/ R.V. Akademik Mstislav Keldysh - 43, stn. 3988, submarine Mir, 44 o 57,40 N 28 o 00,90 W (Mid Atlantic Ridge, N of the Azores), 2800 m. Paratype - IORAS 5/2/ Ibid. Description: Body. The sponge is tubular with relatively small osculum, basiphitous with conules of prostalia lateralia of diactines gathered in tufts. The holotype is 55 mm high, about 30 mm in diameter, the walls are about 5 mm thick, the osculum is 11 x 16 mm in diameter, the conules are 2-3 mm in length, the largest prostalia lateralia protrude 5 mm. The paratype is smaller with broken upper part: 40 mm high, about 25 mm in diameter, the walls are about 3 mm in thickness. Spicules: The choanosomal skeleton is composed of diactines from 1.5 to several mm long and mm in diameter. Fig. 2: Lophocalyx pseudovalida sp. nov., holotype (IORAS 5/2/3111). External shape (scale 20 mm). The smaller ones of these have a widening in the middle; the larger ones are stout; their outer ends conically pointed, rounded, rarely clavate or lanceolate, smooth or rough. The diactines that serve as prostalia lateralia are about 18 mm long and mm in diameter with rough surface. Hypodermal pentactines are of two types for Rossellidae, normal and rare anchorate, which serve as prostalia lateralia. The normal hypodermal pentactines have tangential rays / mm (here and bellow measures offered are smallest length largest length/smallest diameter close to the centre largest diameter close to the centre); the proximal ray is mm long. They are smooth with conically pointed outer ends. The anchorate pentactines have short tangential rays / mm; the proximal ray is about 8 mm long, it is smooth at base and rough in proximal part. Hypoatrialia are mostly hexactines, sometimes diactines and rarely pentactines and stauractines. The choanosomal hexactines have proximal ray mm long, tangential rays are mm long, distal ray is mm long, and their diameter is mm. These rays are smooth, with conically pointed outer ends either smooth or

4 452 Table 2: Some mesures of spicules of Lophocalyx pseudovalida sp. nov. (in mm). IORAS 5/2/ holotype IORAS 5/2/ paratype n avg min max std n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray D oxyhemihexaster and oxyhexaster d oxyhemihexaster and oxyhexaster D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette. rough; the outer ends of proximal rays are often tuberculated. The hypodermal/hypoatrial pentactines and choanosomal hexactines have similar dimensions. The diactines located in the vicinity of the atrial surface have rays similar in shape to those of hexactines. These diactines are mm long and mm thick. Dermalia are pinular hexactines, rarely pentactines with the unpaired ray distally directed. The distal ray of dermal hexactines is clavate with short spines, it is mm in length, the tangential rays are mm long, and the proximal ray is mm long. The rays of dermal spicules, except the dermal one, are rough with conically pointed outer ends; they are mm in diameter. Atrialia are mostly hexactines, sometimes pentactines. The proximal ray of atrial hexactines is mm long, tangential rays are mm long, the distal one is mm long. They are mm in diameter, the proximal ray is nearly equal to the others (which are rough) but it has short spines. Microscleres: The microscleres comprise oxyhemihexasters, some oxyhexasters and strobiloplumicomes. The oxyhemihexasters and oxyhexasters are mm in diameter; their primary rosettes are mm in diameter. Their secondary rays are slightly rough mm in diameter. The strobiloplumicomes are mm in diameter, with primary rosette mm in diameter; their primary rays have a terminal spine. Remarks: The new species is distinguished on the basis of specific dermal spicules, mostly hexactines, and presence of hypoatrial hexactines. The rare anchorate pentactines and composition of microscleres permits referring this sponge to Lophocalyx. Lophocalyx biogasi sp. nov. (Fig. 1, 4; Table 3) Etymology: The name of the species is derived from Biogas, the name of the French expedition during which it was collected. Material examined (slope between the Celtic Sea and Biscay Plain): Holotype - MNHN HCL Biogas V, stn. CW 40, 47 o N 9 o W, 2860 m. Paratypes: MNHN HCL 591; HCL 592; HCL Ibid. MNHN HCL 594; HCL 595- Byocyan II, stn. Pl. 18, submarine Cyana, 47 o N 8 o W, 2000 m. MNHN HCL EPI I, R.V. Suroit, stn. CP 39, 47 o 32 N 8 o 38 W, 2100 m. MNHN HCL Biogas VII, CP 26, 47 o N 8 o W, 2115 m. Description: Body. Various fragments represent this species: tubular or lamellate. The holotype is 90 mm high, 170 x 120 mm in diameter; the walls are about 2 mm in thickness. The paratype MNHN HCL 594 is 22 mm in length, 20 x 40 mm in diameter; the walls are 4 mm thick. The paratype MNHN HCL 595 is 150 mm long, 60 mm in diameter; the walls are about 4 mm thick. The paratype MNHN HCL 597 is 210 mm long, 70 mm in diameter; the walls are 2-3 mm thick. The other fragments are lamellate, some as large as 300 x 300 mm (MNHN HCL 591), 2-5 mm in thickness. Hence the sponges may be large. Spicules: The choanosomal skeleton is composed of diactines with rounded or conically pointed rough outer ends. The smaller diactines have a widening in the middle; the large spicules are stout. Some of the largest diactines have traces of synapticular junctions. The diactines are up to 20 mm in length, mm in diameter. Hypodermal pentactines are of two types: normal and rare anchorate. The normal hypodermal pentactines have tangential rays / mm; the proximal ray is mm long. These rays are smooth with rounded, rough outer ends. The anchorate pentactines have curved tangential rays /0.038 mm. Hypoatrialia are represented by a few pentactines. Dermalia are pinular hexactines and rare pentactines. The distal ray of dermal hexactines is clavate with short spines, it is mm long, the tangential rays are mm long, and the proximal ray is mm long. The rays of dermal spicules, except the distal one, are rough with conically pointed or rounded outer ends; they are mm in diameter. The rare dermal pentactines are similar to hexactines but they have a short rudimentary tubercle instead of the proximal ray. The distal ray of these pentactines is mm long, the tangential rays are about mm

5 453 Fig. 3: Spicules of the holotype of Lophocalyx pseudovalida sp. nov. (IORAS 5/2/3111). A. dermal hexactine. B-C. atrial hexactines. D. atrial pentactine. E-F. hypoatrial pentactines. G-H. choanosomal hexactines. I. anchorate pentactine. J. hypodermal pentactine. K. hypoatrial diactine. L-O. fragments of large choanosomal diactines. P- Q. fragments of small choanosomal diactines. R-S. oxyhemihexasters. T. strobiloplumicome. long, and the reduced proximal ray is mm long. Atrialia are pentactines and hexactines occurring in similar proportions. The proximal pinular ray of these spicules is conically pointed, its outer end protrudes far beyond the last spines, and the spines are sparse and longer than those of dermal spicules are. The other rays are rough. The proximal ray of atrial hexactines is mm long, tangential rays are mm in length, distal one is mm long, and these rays are mm in diameter. The proximal ray of atrial pentactines is mm in length; tangential rays are mm long, distal rudimental one is mm long. Microscleres: Oxyhemihexasters, oxyhexactines and their derivatives with reduced rays, rare oxyhexasters and strobiloplumicomes represent the microscleres. The oxyhemihexasters and oxyhexasters are mm in diameter; their primary rosettes are mm in diameter. The oxyhexactines and their derivatives are mm in diameter. The oxyoidal spicules have

6 454 Table 3: Some measures of spicules of Lophocalyx biogasi sp. nov. (in mm). MNHN HCL holotype MNHN HCL 595 paratype MNHN HCL 597 paratype MNHN HCL 594 paratype n avg min max std n avg min max std n avg min max std n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L dermal pentactine distal ray L dermal pentactine tangential ray L dermal pentactine rudimental ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray L atrial pentactine proximal ray L atrial pentactine tangential ray L atrial pentactine distal rudimental ray D oxyhemihexaster and oxyhexaster d oxyhemihexaster and oxyhexaster D oxyhexactine D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette

7 455 Fig. 4: Spicules of Lophocalyx biogasi sp. nov.: A-G, J, L- P, S, holotype (MNHN HCL 590); H, I, Q, paratype (MNHN HCL 591); K, paratype (MNHN HCL 597); R, paratype (MNHN HCL 594). A. dermal hexactine. B. dermal pentactine. C. atrial pentactine. D. atrial hexactine. E. hypodermal pentactine. F. hypodermal stauractine. G. hypodermal tauactine. H- I. anchorate pentactine. J-K. fragments of large choanosomal diactines. L. small choanosomal diactine. M. oxyhexaster. N. oxyhemihexaster. O oxyhexactine. P and R. abnormal oxyoidal spicules. Q. oxystauractine. S. strobiloplumicome. nearly smooth rays about mm in diameter. The strobiloplumicomes are mm in diameter, with primary rosette mm in diameter; their primary rays have each a terminal spine. Remarks: Lophocalyx biogasi sp. nov. differs from L. pseudovalida sp. nov. by the following features: (1) the dimensions of oxyoidal microscleres ( mm in diameter in L. biogasi sp. nov., mm in L. pseudovalida sp. nov.), (2) the rays of oxyoidal microscleres (thinner in L. biogasi sp. nov., about mm in diameter; vs mm in L. pseudovalida sp. nov.), (3) rarity of oxyhexasters and predominance of derivatives of oxyhexactines and oxyhemihexasters with reduced ray number in L. biogasi sp. nov., (4) smaller size of strobiloplumicomes ( mm in diameter in L. biogasi sp. nov., mm in L. pseudovalida sp. nov.). Very similar specimens to this new species were collected off Peru by the French expedition NAVTIPERC-2, R.V. Naudir. Lophocalyx oregoni sp. nov. (Fig. 1, 5; Table 4) Etymology: The name of the species, oregoni, is derived from the name of the United States research ship R.V. Oregon from which all the type series was collected. Material examined: Holotype - USNM (kt 286) - R.V. Oregon, stn , 24º27 N 87º38 W (Gulf of Mexico), 500 m. Paratypes: USNM (kt 287; kt 288) - Ibid. Description: Body. Lamellate fragments, all of which likely belong to a single specimen, represent this sponge. The holotype is about 1.5 mm in thickness with notable prostalia, probably basalia. The other fragments are 2-4 mm thick without the notable prostalia. It is likely that this sponge is lophophitous. Spicules: The choanosomal skeleton is composed of diactines mm long and mm thick, with conically pointed, rounded or clavate, rough outer ends. The large diactines are stout and the small ones have a widening in the middle, some of them have traces of synapticular junctions.

8 456 Fig. 5: Spicules of the holotype of Lophocalyx oregoni sp. nov. (USNM kt 286). A. dermal hexactine. B. atrial pentactine. C. atrial hexactine. D. hypodermal pentactine. E- F. anchorate pentactines. G-H. large choanosomal diactines. I. small choanosomal diactine. J. oxyhexaster. K. oxyhexactine. L. oxypentactine. M-N. oxystauractines. O. oxydiactine. P. abnormal oxyoidal spicule. Q. strobiloplumicome. Table 4: Some measures of spicules of Lophocalyx oregoni sp. nov. (in mm). USNM(kt286) - holotype USNM(kt288) paratype n avg min max std n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactin tangential ray L atrial hexactine distal ray L atrial pentactine proximal ray L atrial pentactine tangential ray L atrial pentactine rudimental ray D oxyhemihexaster and oxyhexaster d oxyhemihexaster and oxyhexaster D oxyhexactine D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette Hypodermal pentactines are of two types: normal and anchorate, the latter of which act as prostalia. The normal hypodermal pentactines have tangential rays / mm; the proximal ray is about 1.5 times the length of the tangential ones. The rays are smooth with conically pointed rough outer ends. The anchorate pentactines have short tangential rays / mm; the proximal ray is mm long. Some of the anchorate pentactines near the dermal surface (mostly with short proximal ray) have hook-like tangential rays and smooth proximal rays.

9 457 The largest pentactines located above the dermal surface and likely functioning as basalia have straight tangential rays bent proximally (the tangential rays have dimensions similar to the hook-like ones) and shafts which become rough at a certain distance from the distal part of the spicule. Hypoatrialia are likely to be absent; some rare orthotropal pentactines are very similar to dermal ones and may be allochthonous. Dermalia are pinular hexactines, rarely pentactines with the unpaired ray distally directed and a rudimental tubercle as a reduced proximal ray. The distal ray of dermal hexactines is mm long, it is clavate with relatively long spines, the tangential rays are mm long, and the proximal ray is mm long. The rays of the dermal spicules, except the dermal one, are rough with conically pointed outer ends; they are mm in diameter. Atrialia are mostly pinular pentactines, sometimes hexactines. Their pinular ray is conically pointed; less spiny and thinner than those of dermal spicules, the reduced distal ray in pentactines is represented by a short tubercle; the tangential rays are rough with conically pointed outer ends. The proximal ray of atrial pentactines is mm in length, tangential rays are mm long, and the reduced distal ray is mm long. The distal ray of atrial hexactines is mm in length. These rays are mm in diameter. Microscleres: The microscleres are abnormal oxyoidal spicules (with some reduced rays) derived from oxyhexactinic forms and oxyhemihexasters, oxyhexactinal microscleres are rare oxyhexasters and strobiloplumicomes. Most of the oxyoidal spicules have hook-like outer ends, rarely straight; their rays are slightly rough. The oxypentactines, oxystauractines and oxyhexactines prevail over the other forms. Sometimes they have small tubercle-like rudiments in place of the absent ray. These spicules are mm in diameter. Their rays are rough and mm in diameter. The strobiloplumicomes are mm in diameter, with a primary rosette mm in diameter; the primary rays each have a short terminal spine. Remarks: Lophocalyx oregoni sp. nov. is distinguished by its peculiar hook-like oxyoidal microscleres mostly comprising abnormal forms of oxyhexactines with some reduced rays; specific dermal spicules are mostly hexactines, and atrial spicules are mostly pentactines. The anchorate pentactines with straight tangential rays in addition to curved ones are known also in L. philippinensis (e.g. Tabachnick, 2002) but in L. oregoni sp. nov. these spicules prevail in the likely basalia, while the hook-like ones do not protrude far and act as prostalia lateralia. Lophocalyx brasiliensis sp. nov. (Fig. 1, 6; Table 5) Etymology: The name of the species is derived from its type locality, off the southeastern Brazilian Coast. Material examined: The holotype is distinguished formally as a fragment which together with the other fragments (paratypes), probably belong to a single specimen. Holotype - MNHN HCL R.V. Marion Dufresne - MD 55, sta. 64 CB 105, 23 o 46 S 42 o 9 W (southeastern Brazilian continental slope), m. Paratypes: MNHN HCL Ibid. Description: Body. The studied material is composed of relatively small fragments. The holotype was chosen on the basis of its possession of both dermal and atrial spicules in the same fragment, a single conule with a tuft of prostalia lateralia 15 mm in length, 12 x 3 mm in diameter. The other fragments (paratypes) have a similar shape. Prostalia lateralia protrude about 20 mm. Spicules: The choanosomal skeleton is composed of diactines from 1.8 mm to several mm in length; their diameter is mm. The diactines, of the prostalia are longer (about 25 mm long). The choanosomal diactines are conically pointed, rarely clavate, with rough or rarely smooth outer ends. The smaller diactines have a widening in the middle, the larger ones are stout. Hypodermal pentactines are of two types: normal and anchorate, both as prostalia. Some of the normal hypodermal pentactines have smooth rays with conically pointed rough outer ends; their tangential rays are / mm, the proximal ray is mm long. The other normal pentactines with regular orthotropal tangential rays have all the rays rough with conically pointed outer ends. Their tangential rays are smaller and thinner, mm long and mm thick, than those in smooth pentactines, the proximal ray is usually about 1.5 times longer. The anchorate pentactines are very rare; rays are smooth, with straight or slightly bent tangential ones. The tangential rays are mm long; the proximal ray is very long, being mm in diameter at base and mm in diameter in some distance. Hypoatrialia seem to be similar to the hypodermal spicules, but anchorate forms are lacking and hexactines with short proximal ray and rounded rough outer end are present. The proximal ray of these spicules is mm long and mm thick; tangential rays are mm long, the distal ray is mm long and is equal in diameter to the other rays at its base, but thicker (about 0.04 mm in diameter) at some distance from the base. Dermalia are pinular hexactines with slightly clavate pinular distal ray bearing relatively long spines; all other rays are rough. The distal ray is / mm, the tangential rays are mm long, and the proximal ray is mm long. Atrialia are hexactines with spiny or rough rays. Their proximal ray is / mm, the tangential rays are mm long, and the distal ray is mm long. Microscleres: The microscleres comprise oxyoidal, asterous or nearly asterous (with very short primary rays) spicules derived from oxyhemihexasters, oxyhexactines and strobiloplumicomes. Most oxyoidal spicules have minute spines, but sometimes a few rays may have long spines. These spicules are mm in diameter, and morphotypes with and without secondary rays are present. The central portions of these microscleres vary from minute to slightly spherical, mm in diameter. The rays of these spicules are mm in diameter. The strobiloplumicomes are mm in diameter, with primary rosette mm in diameter; the primary rays have each a short terminal spine.

10 458 Fig. 6: Spicules of Lophocalyx brasiliensis sp. nov.: A-G, I-W, holotype (MNHN HCL 598); H, paratype (MNHN HCL 605). A-B. dermal hexactines. C-D. atrial hexactines. E. choanosomal hexactine. F. hypodermal rough pentactine. G-I. hypodermal pentactine. J-L. large choanosomal diactines. M. small choanosomal diactine. N-Q. asters derived from oxyhemihexasters. R. oxystauractine. S-T. oxyoidal microscleres derived from hexactines. U-V. outer ends of oxyoidal microscleres. W. strobiloplumicome. Remarks: Lophocalyx brasiliensis sp. nov. is distinguished from other species in the genus by its characteristic oxyoidal microscleres which have asterose forms with thick secondary rays mm in diameter. These microscleres are derived mostly from oxyhemihexasters. Lophocalyx atlantiensis sp. nov. (Fig. 1, 7, 8; Table 6) Etymology: The name of the species indicates its central north Atlantic type locality. Material examined (Mid Atlantic Ridge, Charlie-Gibbs Fracture Zone): Holotype - MZUB n Mar-Eco, superstation 70, local station 385, N, W, m. Paratype - MZUB n Mar-Eco, superstation 60, local station 380, N W, m. Description: Body. The holotype is ovoid, 23 mm high, its diameter is 19 mm in the middle, the osculum is about 4 mm in diameter, several tufts of prostaslia lateralia protrude about 5 mm above the body. The paratype is a small fragment. Spicules: Choanosomal spicules are diactines mm long and mm thick, with a widening in the middle portion, surrounded by four rudimentary tubercles; their outer ends being conically pointed, rough.

11 459 Table 5: Some measures of spicules of Lophocalyx brasiliensis sp. nov. (in mm). MNHN HCL holotype n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray D oxyhemihexaster and oxyhexaster d oxyhemihexaster and oxyhexaster D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette Prostalia in tufts are mostly diactines about 7 mm long and thick mm. Hypodermal pentactines are of two types: normal and anchorate. The anchorate pentactines have tangential rays about 0.12 mm long, the proximal ray is more than 3 mm long, their diameter is 0.04 mm, and the tangential rays are usually smooth. The normal pentactines have tangential rays mm long, the proximal ray is mm long, their diameter is mm, the rays and outer ends of these spicules are usually smooth, rarely rough. Choanosomal hexactines were found in large amounts in the paratype, their distal ray is rough while all the other rays are smooth. The distal ray in hypodermal hexactines is about 0.30 mm long, other rays are about 0.5 mm long, and their diameter is about mm. Dermalia are pinular hexactines, their pinular ray is stout, mm long, tangential rays are mm long, the proximal ray is mm long, their diameter is mm. Atrialia are also pinular hexactines, their proximal ray is mm long, tangential rays are mm long, distal ray is mm long, Fig. 7: Lophocalyx atlantiensis sp. nov., holotype (MZUB n14822). External shape (scale 10 mm). their diameter is mm. The rays of dermal and atrial spicules are rough or spiny and have conically pointed outer ends. Microscleres: Oxyoidal microscleres are mostly oxyhexactines and a few oxyhemihexasters, the latter have one ray branching into two, rarely three secondary rays. The diameter of oxyhexactines and oxyhemihexasters is mm; the primary ray in the oxyhemihexasters is mm. The strobiloplumicomes are mm in diameter with primary rosette mm in diameter; the primary rays have each a short, conical terminal spine. Table 6: Some measures of spicules of Lophocalyx atlantiensis sp. nov. (in mm). MZUB n holotype MZUB n paratype n avg min max std n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray D oxyhexactine or oxyhemihexaster D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette

12 460 Fig. 8: Spicules of Lophocalyx atlantiensis sp. nov.: A-E, G- N, holotype (MZUB n14822); F, paratype (MZUB n15318). A-B. dermal hexactines. C. atrial hexactine. D. anchorate pentactine. E. hypoatrial pentactine. F. choanosomal hexactine. G. prostalia lateralia (diactine). H-J. choanosomal diactines (an outer end and central parts). K. oxyhexactine. L-M. oxyhemihexasters. N. strobiloplumicome. Remarks: The specific features of this new species are that oxyoidal microscleres are in the majority oxyhexactines; oxyhemihexasters (with one ray branching only) are only rarely seen, and the pinular ray of dermal hexactines is stout. Lophocalyx reiswigi sp. nov. (Fig. 9, 10; Table 7, 8) Etymology: The specific epithet honours Dr. Henry M. Reiswig (Royal British Columbia Museum, Victoria, Canada), one of the greatest specialists on hexactinellid systematics, who has just turned 70. Material examined: The holotype is distinguished formally as a fragment, which together with the others fragments (paratypes), probably belong to a single specimen. Holotype - MNRJ 3339C R.V. Thalassa, Programme REVIZEE Bahia II, stn.0496, 07.VI.2000, 13º S 38º W (off Bahia State, Brazil), 1717 m. Paratypes - MNRJ 3339A, MNRJ 3339B, MNRJ 3339D, MNRJ 3339E, MNRJ 3339G and MNRJ 3339I Ibid.

13 Fig. 9: Lophocalyx reiswigi sp. nov.: A-B. holotype (MNRJ 3339C); C-F. paratypes (MNRJ 3339A, MNRJ 3339B, MNRJ3339D, MNRJ 3339E, respectively). External shape (scale 50 mm). 461

14 462 Fig. 10: Spicules of Lophocalyx reiswigi sp. nov.: A-O, holotype (MNRJ 3339C); P, paratype (MNRJ 3339B). A-B. dermal hexactines. C-F. atrial hexactines. G. anchorate pentactine. H-I. hypodermal pentactines. G. prostalia lateralia diactine. K. choanosomal diactine. L. oxyhemihexaster. M. oxyhexactine. N. secondary ray of oxyoidal microsclere. O. strobiloplumicome. P. choanosomal hexactine. Table 7: Some measures of spicules of Lophocalyx reiswigi sp. nov. (in mm). MNRJ 3339 A paratype MNRJ 3339 B paratype MNRJ 3339 C - holotype n avg min max std n avg min max std n avg min max std L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray D oxyhexactine or oxyhemihexaster D strobiloplumicome d strobiloplumicome L - length, D - diameter, d - diameter of primary rosette

15 463 Table 8: Some measures of spicules of new species of Lophocalyx (in mm). L. pseudovalida L. biogasi L. oregoni L. brasiliensis L. atlantiensis L. reiswigi sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. min max min max min max min max min max min max L dermal hexactine distal ray L dermal hexactine tangential ray L dermal hexactine proximal ray L atrial hexactine proximal ray L atrial hexactine tangential ray L atrial hexactine distal ray D oxyoidal microsclere D strobiloplumicome L - length, D - diameter, d - diameter of primary rosette Description: Body. The lamellate holotype is a fragment 200 mm long, 145 mm in diameter; the walls are 1-4 mm thick, and with tufts of prostalia marginalia piercing the surface for 9 mm at most. Other specimens are also lamellate fragments, and these could all possibly belong to a single specimen. Both surfaces of the holotype are clearly reticulated to the naked eye, with meshes 1-5 mm in diameter. The paratype MNRJ 3339A is the largest fragment, with 295 x 135 mm in area. MNRJ 3339B has thick walls (10-19 mm), and a detachable atrial surface membrane. MNRJ 3339D is harder than the other specimens. MNRJ 3339I comprises several fragments, one of them with tufts of prostalia marginalia protruding for 15 mm over the dermal surface, and meshes of up to 7 mm in diameter. Spicules: The choanosomal skeleton is composed of diactines with microspined terminations which can be rounded or conically pointed. The diactines of the prostalia lateralia are the stouter ones (1-3/ mm), while the choanosomal diactines are slender, slightly curved or straight, and have four rudimentary tubercles in the middle (up to at least 19 mm in length and mm in thickness). Many choanosomal diactines are fused by secondary synapticular junctions. Hypodermal pentactines occur in two types: normal and rare anchorate. The normal hypodermal pentactines have short tangential rays 0.5-1/ mm and proximal rays from 1 mm up to at least 4 mm long, mm in diameter; the rays have conical terminations, and can be spined or rough, all over, or on a variably long distal section. Anchorate pentactines have smooth tangential rays, / mm. Hypoatrial pentactines are equal to dermal orthotropal ones. The choanosomal hexactines have slightly curved, rough rays (at least in the distal and proximal rays) and gradually tapering ends. The distal ray in the choanosomal hexactines is about / mm. Dermalia are pinular hexactines with slender pinular distal rays (sometimes reduced) bearing short spines; all other rays are rough or microspinouse. The distal rays are / mm, the tangential rays are mm, and the proximal rays are / mm. Atrialia are pinular hexactines with slender or slightly clavate distal rays bearing short spines; all other rays are rough. The distal rays are / mm, the tangential rays are / mm, and the proximal rays are / mm. Microscleres: Thin rayed oxyoidal microscleres are oxyhexactines and oxyhemihexasters mm in diameter. The primary rays of the latter may branch into 2 secondary rays. The strobiloplumicomes are about mm in diameter with primary rosettes mm in diameter. Remarks: Lophocalyx reiswigi sp. nov. is closely related to L. atlantiensis sp. nov. on the basis of the marked similarity of their spicule complement. Main points of distinction are the tangential rays of the anchorate hypodermal pentactines, bent, becoming parallel to the main axis in L. reiswigi sp. nov. and flat, perpendicular to the main axis in L. atlantiensis sp. nov. The normal pentactines are always rough in the former species but only rarely so in the latter species; the pinular ray is larger in the dermal hexactines of L. reiswigi sp. nov. ( mm vs mm); and the oxyhemihexasters may have more than one branching ray. Lanuginellinae indet. An interesting representative of this subfamily was captured together with L. pseudovalida sp. nov. (R.V. Akademik Mstislav Keldysh - 43, stn. 3988). Unfortunately it is a strongly damaged fragment which contains many allochthonous spicules of other Hexactinellida collected simultaneously, and which is not possible to assign confidently to a genus. Its spicules strongly differ from all known species of the genera Lophocalyx and Calycosoma to one of which this fragment should be referred on the basis of its microsclere complement. The microscleres of this fragment resemble much those of L. biogasi sp. nov. by having many oxyhexactines, while the hypodermal and/or hypoatrial pentactines are mostly large, rough spicules, similar in shape to the small, rough pentactines of L. brasiliensis sp. nov. The possibly dermal and atrial spicules of this sponge have no peculiar features, but are clearly different from those of L. biogasi sp. nov. - proximal rays of the likely atrial spicules of this fragmented sponge are entirely covered by spines.

16 464 Discussion Remarks on the new taxa of Lophocalyx The species of Lophocalyx may be divided into three groups by characteristics of their dermal skeleton. 1. Sponges with dermal stauractines: L. philippinensis (e.g. Carter, 1875, Schulze, 1886) and L. suluanus (Ijima, 1927) (e.g. Ijima 1927, Lévi 1964). 2. Species where the stauractines are supplemented by pentactines with distally directed unpaired ray: L. spinosa (Schulze, 1900) (e.g. Schulze 1900, 1902). 3. Species with predominance of dermal hexactines (the pentactines are rare): L. moscalevia Tabachnick, 1988 (e.g. Tabachnick 1988, Tabachnick and Lévi 2004), L. sp. from the Antarctic (Topsent 1910, 1913, Barthel and Tendal 1994) and all species of Lophocalyx from the Atlantic Ocean described in this paper. Lophocalyx sp. from the Antarctic [probably a new species; Calycosoma validum sensu Topsent (1910)] is easily distinguished from the other species of the 3 above groups by peculiar, very large dermal hexactines with pinular distal ray covered by spines almost everywhere (Topsent 1910, 1913). Other specimens of Lophocalyx from the Antarctic collected by the R.V. Eltanin (deposited in the USNM, reexamined by KT) are very similar to the specimen of Topsent after analysis of his material stored in the NMS. They all seem to have large hypoatrial hexactines and their interpretation by Topsent as dermal may be erroneous. Unlike the hexactines described above, those in L. pseudovalida sp. nov. have spines or tubercles situated close to the upper end. Lophocalyx moscalevia is more similar to the Atlantic species but its dermal and atrial spicules differ considerably in dimensions from each other. The other differences in the microscleres composition seem to be less important. Lophocalyx moscalevia has oxyhemihexasters, oxyhexactines and derivatives of the latter with reduction of ray number. The features, which allow recognition of the Atlantic species of Lophocalyx, are given in the remarks after their descriptions. Some microdiscohexasters were reported by Ijima (1927) for one of three specimens of L. suluanus. Some discohexasters were found in several specimens of L. philippinensis (Tabachnick 2002). But these discoidal spicules are absent in all the Atlantic species. All the new species of Lophocalyx from the Atlantic Ocean have many common features: dermalia are hexactines, the spicule s measures mostly overlap between all these species (see Table 8). The features which permit their differentiation are the shape of dermal pinular rays and the type and shape of oxyoidal microscleres. Remarks on the affinities of Calycosoma and Lophocalyx The genus Lophocalyx (then Polylophus) was created by Schulze (1887) for a single specimen, named L. philippinensis (Gray, 1872). Ijima (1927) later mostly accepted his diagnosis. Several species of Lophocalyx have been described since and its diagnosis required clarification. The genus Calycosoma never had a diagnosis (before Tabachnick 2002), being differentiated in keys from allied Lophocalyx by their dermal skeleton construction. Schulze (1904) proposed to differentiate both genera on the basis of presence of hexactines and pentactines in the former, and stauractines in the latter. At the same time, Ijima (1904) proposed the basiphytous habit, prostalia diactines, dermalia pentactines and hexactines as diagnostic for Calycosoma; while the lophophitous habit, pentactine anchors, dermalia stauractines and pentactines as diagnostic for Lophocalyx. If the criteria by Schulze were the only ones available, Calycosoma would better be regarded as a junior synonym of Lophocalyx through newly found transitional forms: L. moscalevia Tabachnick, 1988 and the new species described above. Furthermore, the criteria by Ijima are not strong enough since a basyphitous form of Lophocalyx is already known. Lophocalyx moscalevia Tabachnick, 1988 is fixed to a dead Hexactinosida directly by the base, but numerous anchorate hypodermal pentactins are also present. Besides it is also possible to imagine lophophytous Calycosoma fixed by diactines only (without anchorate spicules) to soft substrata. Thus, the most important feature, which still allows the distinction of these two genera, is the presence in Lophocalyx of a second category of hypodermal pentactines, which are extended and serve as prostalia anchorate spicules, and are absent in Calycosoma. Additionally, Calycosoma has prostalia diactines gathered in tufts. Calycosoma validum described from the Antarctic by Topsent (1910, 1913) should be transferred to Lophocalyx, as it has these anchorate spicules and also due to other differences from C. validum from the type location. The Antarctic material possibly belongs to a new species, the description of which depends on re-analysis of C. validum s type specimen and other Antarctic materials. After description of the new species of Lophocalyx with hexactines in dermalia and especially the new species from the Atlantic Ocean, the definition of Calycosoma also required a revision. The new species of Lophocalyx described here approach closely the spicule set known from C. validum; most species of Lophocalyx from the Atlantic have few anchorate pentactines and walls which are smooth (without prostalia lateralia) or with tufts of prostalia lateralia which are mostly diactines (L. atlantiensis sp. nov., L. brasiliensis sp. nov., L. pseudovalida sp. nov.) as it is known for Calycosoma. But the external shape of the body and the shape of the hypodermal pentactines seem to be important for generic definitions within the Lanuginellinae (Tabachnick 2002). The presence of a peduncle defines three doubtless basiphytous genera: Calycosoma (very short peduncle), Lanugonychia von Lendenfeld, 1915 and Sympagella Schmidt, Lanuginella Schmidt, 1870 is a basiphytous sponge but with tendency to form a veil of outwardly protruded hypodermal pentactines in some specimens and it may thus have lophophytous forms. Most Atlantic species of Lophocalyx (except L. oregoni sp. nov.) seem to be basiphytous. The shape of the common hypodermal pentactines seems to be an important feature for the recognition of Mellonympha Schulze, 1897: it has hypodermal pentactines with paratropal tangential rays, besides anchorate ones (Tabachnick 2002). The only remaining Lanuginellinae genus, Dochonestes Topsent, 1928,