Cliona, GRANT, 1826

Carballo, José Luis, Cruz-Barraza, José Antonio & Gómez, Patricia, 2004, Taxonomy and description of clionaid sponges (Hadromerida, Clionaidae) from the Pacific Ocean of Mexico, Zoological Journal of the Linnean Society 141 (3), pp. 353-397 : 354-381

GENUS CLIONA GRANT, 1826

Synonymy: see Rützler (2002a).

Type species: Cliona celata Grant, 1826 .

Diagnosis: Sponges primarily in alpha growth form (excavating chambers, communicating through papillae), some species developing beta stage by merging of papillae, very few regularly outgrowing their substratum and occurring in gamma stage. Some gamma stage species attain a large, irregular massive or cup shape but do not develop specialized incurrent or excurrent features other than, the original pori- and oscula-bearing papillae in some forms. Spicules are tylostyles as megascleres, very thin oxeas (rhaphides) as accessory scleres but never of structural importance, and spiraster-like microscleres. Spirasters are straight, bent, kinked, spiralled, or undulated spiny rhabds, including amphiastrose forms clearly derived from true spirasters; extreme forms may be entirely smooth ( Rützler, 2002a).

CLIONA PAPILLAE SP. NOV.

( FIGS 2 , 3 , 4)

Holotype: MNCN 1.01 /234, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 09.v.2003.

Paratypes: BMNH: 2003.6.27.1, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 09.v.2003. LEB-ICML-UNAM-160 , Isla de Cardones (Mazatlán, Sinaloa), 23∞11¢05¢¢N-106∞24¢07¢¢W, 8 m depth, 15.iii.2000, on rocks. LEB-ICML-UNAM-316 , Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N- 106∞27¢43¢¢W, 6 m depth, 20.ii.2000. LEB-ICML- UNAM-913 , Isla Lobos 2 (Mazatlán, Sinaloa) 23∞13¢27.7¢¢N, 106∞28¢01.6¢¢W, 7 m depth, 03.x.2003, on rocks .

Description: Boring sponge growing always in alpha stage ( Fig. 2A ). The sponge can reach up to 12 ¥ 10 cm in coverage. One of the most typical features of this species is the presence of elevated papillae that can protrude up to 8 mm on the substrate when alive. The papillae bear either grouped ostia (visible to the eye underwater) or oscula ( Fig. 2B ). The ostial papillae are regularly scattered on the surface, and they are usually spaced 1.3–10 mm from each other. Ostial papillae bear many ostia, sieve-like. They are circular or oval in section, and characteristically distally widened; the diameter in the centre is 1.8–3.5 mm, and at the end 2–6.8 mm. The oscular papillae are circular or oval-shaped, from 1 to 4.3 mm in diameter. Papillae fusion is not common. The species has a very distinctive very pale yellow colour when alive. Upon preservation in alcohol, colour changes to light brown.

Skeletal characters: Tylostyles are mostly straight or only slightly curved, with generally well-formed spherical or oval head ( Fig. 3A ). Tylostyles measures 304 ¥ 9.5 Mm on average (13 Mm of head width) ( Table 1). Tylostrongyles occasionally appear. The spirasters are very delicate, slender, with about 2–5 bends (21 Mm length in average) ( Fig. 3B ). The shaft surface is characteristically very finely spined, with nonbifurcated spines. In the periphery of the papillae there is a dense palisade of tylostyles, with tylostyle heads anchored in tissue and pointed ends piercing the surface ( Fig. 4). In the inner part of the papillae the tylostyles are irregularly scattered, there are occasional tracts of spicule bundles in the central canal. The spirasters are very scarce in the papillae. They are only abundant in the choanosome.

Etymology: The species name selected means ‘nipple’ in Latin to emphasize the characteristic papillae of this species.

Distribution: Bahía de Mazatlán (Mazatlán, Sinaloa) (present study).

Remarks: The closest species seems to be Cliona lobata sensu de Laubenfels (1954) , which assigns to this species a bright yellow specimen with very slen- der spirasters (27–40 ¥ 0.3 Mm), and tylostyles (220– 240 ¥ 5–12 Mm) with a marked neck and a very welldeveloped head (8–15 Mm in diameter). These characteristics in part agree with our specimen; however, C. lobata Hancock, 1849 , in addition to slender and longer spirasters, has additional thicker and shorter spirasters ( Topsent, 1887; Volz, 1939). Other close species seem to be Cliona caribbea Carter, 1882 (redescribed in Rützler, 1974), and Cliona paucispina Rützler (1974) . Cliona caribbea is a greenish or dark brown sponge, which has only slender spirasters similar to the C. papillae spirasters. However, important differences exist between them, like the size (32.8 ¥ 0.8 Mm in average), and the distribution and form of the spines in the spirasters of Cliona caribbea , which has singular or bifurcated actins ( Rützler, 1974; Schönberg, 2000). Cliona paucispina ( Rützler, 1974) has shorter spirasters (12.8–43.2 ¥ 0.8–3 Mm), with much smaller spines, and fewer spines and bends than C. papillae . The presence of tylostrongyles, and the morphology of the papillae are other distinguishing characteristics of C. papillae , which do not appear in the related species commented upon above.

CLIONA VALLARTENSE SP. NOV.

( FIGS 2 , 5 )

Holotype: MNCN-1.01/235, Mismaloya ( Bahía de Banderas , Jalisco), 20∞31¢56¢¢N-105∞17¢42¢¢W. 3 m depth, 06.ix.2003.

Paratypes: BMNH-2003.6.27.2, Mismaloya ( Bahía de Banderas , Jalisco), 20∞31¢56¢¢N-105∞17¢42¢¢W, 3 m depth, 06.ix.2003 . LEB-ICML-UNAM-633, Conchas Chinas ( Puerto Vallarta , Jalisco), 20∞35¢16¢¢N- 105∞14¢42¢¢W. 2 m depth, 08.x.2002 . LEB-ICML- UNAM-788, Mismaloya ( Bahía de Banderas , Jalisco), 20∞31¢56¢¢N-105∞17¢42¢¢W, 3 m depth 06.ix.2003 . LEB- ICML-UNAM-821, Antiguo Corral del Risco ( Punta Mita , Nayarit), 20∞46¢20¢¢N-105∞32¢49¢¢W, 2 m depth, 11.vi.2003 . LEB-ICML-UNAM-848, Majahuita ( Bahía de Banderas , Jalisco), 20∞29¢06¢¢N-105∞17¢42¢¢W, 10 m depth 06.viii.2003 .

Description: This species grows in alpha and beta stage, overgrowing the substrate completely and covering it with a thin (0.9–2.8 mm) layer of tissue in the beta stage ( Fig. 2C, D ). In some specimens the papillae are surrounded by patches of ectosomal tissue overgrowing the substrate. The specimens can reach a maximum surface area of 20 ¥ 10 cm. Papillae are variable in size (0.2–1.3 cm) and form (from circular to elongated), and scarce in the alpha form. Oscules and ostia frequently flush with the surface. Preserved, the sponge shrinks and mirrors the contour of the substrate beneath. Colour is green olive to pale yellow. The tissue surrounding papillae is light olive or yellow. The choanosome is ochre. This species penetrates deeper into carbonate structures (up to 9 cm depth in the holotype) than the others Cliona species in the same area.

Skeletal characters: The species presents tylostyles and spirasters ( Fig. 5 ). The tylostyles are slender, slightly curved, and gradually tapering from the middle of the spicule. The heads are characteristically narrow and elongate, sometimes ovoid, with annular swellings. Deformations are also common, and the head region of the spicule may exhibit a rather wide variety of shapes. They measure 294 ¥ 4.7 Mm on average (7.5 Mm of head width) ( Table 2). Spirasters have a thick shaft, with narrowly spaced small conic spines surrounding the entire axis of the spirasters. Sometimes the spines are not pointed but branching two, three or even more times. They may be spiral with two or more bends with almost straight shafts (24 Mm length on average). The spirasters are very scarce in the ectosome and in the papillae, but they are abundant in the choanosome around the excavating chambers.

Etymology: The specific epithet refers to the village of Puerto Vallarta from the Bahía de Banderas (Jalisco, Mexico).

Distribution: Bahía de Banderas (Puerto Vallarta, Jalisco) (present study).

Remarks: Cliona vallartense sp. nov. is a greenish sponge characterized mainly by skeletal details of the megascleres and by the form and spination of the spirasters. Cliona orientalis Thiele, 1900 seems to be the most similar species. However, spines on their spirasters are usually arranged at the convex side of the spicule parts ( Thomas, 1979; Calcinai et al., 2000), forming little bouquets along the shaft ( Schönberg, 2000). Our specimens have spirasters with spines arranged around the shaft, and they are commonly bifurcated. Cliona caribbea is another greenish or dark brown sponge, which has only slender spirasters with singular or bifurcated actins ( Rützler, 1974). Another close species seems to be Cliona paucispina Rützler, 1974 , which is clearly different from our specimens because its spirasters have reduced spination and nonbifurcated spines ( Rützler, 1974).

CLIONA CALIFORNIANA ( DE LAUBENFELS, 1932) COMB. NOV.

( FIGS 6 , 7 , 8 , 9 )

Synonymy: Cliona celata var. californiana de Laubenfels, 1932: 47 ; Pseudosuberites pseudos Dickinson, 1945: 38 ; syn. nov.

Paratype: Cliona celata var. californiana de Laubenfels, 1932 . BMNH-29.8.22.52a (microscope slide), BMNH-29.8.22.50 (specimen in spirit).

Holotype: Pseudosuberites pseudos Dickinson, 1945 . Allan Hancock Foundation, Sta. No. velero 553-36. MNCN-1.01/237, Estero el Bichi (Topolobampo, Sinaloa), 25∞32¢27¢¢N-109∞05¢29¢¢W, 1 m depth, 13.xi. 2002. LEB-ICML-UNAM-141, Chacala (Nayarit), 21∞09¢57¢¢N-105∞13¢38¢¢W, 4 m depth, 20.ii.2000, on sand bottoms. LEB-ICML-UNAM-224, Puente Maviri (Los Mochis, Sinaloa), 25∞34¢55¢¢N-109∞06¢52¢¢W, 2 m depth, 21.vi.2000. LEB-ICML-UNAM-285, Ensenada de Bacochibampo (Guaymas, Sonora), 27∞54¢37¢¢N- 110∞57¢12¢¢W, 5 m depth, 06.xi.2000. LEB-ICML- UNAM-295, Punta Cazón (Kino, Sonora), 28∞52¢20¢¢N- 112∞02¢01¢¢W, 3 m depth, 08.xi.2000. LEB-ICML- UNAM-300, Isla del Peruano (Guaymas, Sonora), 27∞54¢35¢¢N-110∞58¢17¢¢W, 15 m depth, 03.xi.2000. LEB-ICML-UNAM-319, Isla del Peruano (Guaymas, Sonora), 27∞54¢35¢¢N-110∞58¢17¢¢W, 12 m depth, 03.xi.2000. LEB-ICML-UNAM-336, Isla Tiburón (Kino, Sonora), 27∞54¢35¢¢N-110∞58¢17¢¢W, 15 m depth, 27.iv.2001. LEB-ICML-UNAM-337, Isla Tiburón (Kino, Sonora), 28∞47¢12¢¢N-112∞15¢6¢¢W, 15 m depth, 27.iv.2001. LEB-ICML-UNAM-338, Isla Tiburón (Kino, Sonora), 28∞47¢12¢¢N-112∞15¢6¢¢W, 15 m depth, 27.iv.2001. LEB-ICML-UNAM-369, Punta Chile (Mazatlán, Sinaloa), 23∞12¢29¢¢N-106∞25¢40¢¢W, 1.5 m depth, 08.x.2001. LEB-ICML-UNAM-444, Isla Pájaros 1 (Mazatlán, Sinaloa), 23∞15¢29¢¢N-106∞28¢25¢¢W, 3 m depth, 26.ii.2002. LEB-ICML-UNAM-494, Los Arcos (Puerto Vallarta, Jalisco), 20∞32¢73¢¢N-105∞18¢47¢W, 5 m depth, 08.iv.2002. LEB-ICML-UNAM-667, Cerro San Carlos (Topolobampo, Sinaloa), 25∞35¢33¢¢N- 109∞02¢39¢¢W, 4 m depth, 12.xi.2002. LEB-ICML- UNAM-672, Muelle de Contenedor (Topolobampo, Sinaloa), 25∞34¢55¢¢N-109∞03¢32¢¢W, 5 m depth, 12.xi.2002. LEB-ICML-UNAM-676, Punta de la Virgen (Topolobampo, Sinaloa), 25∞36¢58¢¢N- 108∞58¢12¢¢W, 1 m depth, 12.xi.2002. LEB-ICML- UNAM-681, Isla Tunosa (Topolobampo, Sinaloa), 25∞34¢58¢¢N-109∞00¢51¢¢W, 2 m depth, 12.xi.2002. LEB-ICML-UNAM-692, Islas Verdes (Topolobampo, Sinaloa), 25∞31¢47¢¢N-109∞05¢27¢¢W, 2 m depth, 13.xi.2002. LEB-ICML-UNAM-701, Estero el Bichi (Topolobampo, Sinaloa), 25∞32¢27¢¢N 109∞05¢29¢¢W, 1 m depth, 13.xi.2002. LEB-ICML-UNAM-704, Cerro Partido (Topolobampo Sinaloa), 25∞32¢7¢¢N- 109∞05¢33¢¢W, 1 m depth, 13.xi.2002. LEB-ICML- UNAM-711, Estero el Zacate (Los Mochis, Sinaloa), 25∞36¢25¢¢N-109∞04¢33¢¢W, 2 m depth, 14.xi.2002. LEB-ICML-UNAM-726, Puente el Maviri (Los Mochis, Sinaloa) 25∞34¢55¢¢N-109∞06¢52¢¢W, 6 m depth, 14.xi.2002. LEB-ICML-UNAM-738, Paraje Viejo (Guaymas, Sonora), 27∞52¢20¢¢N-110∞52¢08¢¢W, 8 m depth, 26.xi.2002. LEB-ICML-UNAM-741, Paraje Viejo (Guaymas, Sonora), 27∞52¢20¢¢N-110∞52¢08¢¢W, 5 m depth, 11/26/02. LEB-ICML-UNAM-753, Islas Gringas (Guaymas, Sonora), 27∞53¢5¢¢N-110∞57¢55¢¢W, 10 m depth, 27.xi.2002. LEB-ICML-UNAM-766, Islas Gringas (Guaymas, Sonora), 27∞53¢05¢¢N- 110∞57¢55¢¢W, 10 m depth, 27.xi.2002. LEB-ICML- UNAM-767, Cabo Haro (Guaymas, Sonora), 27∞52¢5¢¢N-110∞57¢2¢¢W, 12 m depth, 27.xi.2002. LEB- ICML-UNAM-773, Cabo Haro (Guaymas, Sonora), 27∞52¢04¢N-110∞57¢01¢¢W, 12 m depth, 27.xi.2002. LEB-ICML-UNAM-784, Estero ‘ El Bichi’ (Topolobampo, Sinaloa), 25∞32¢27¢¢N-109∞05¢29¢¢W, 1 m depth, 13.xi.2002. LEB-ICML-UNAM-785 Estero ‘ El Bichi’ (Topolobampo, Sinaloa), 25∞32¢27¢¢N- 109∞05¢29¢¢W, 1 m depth, 13.xi.2002. LEB-ICML- UNAM-833 Altata 3 (by trawling, Sinaloa), 24∞32¢42.9¢N-108∞07¢41.58¢W, 53 m depth, 18.xi.2002. LEB-ICML-UNAM-838, Altata 2 (by trawling, Sinaloa), 24∞27¢58.8¢¢N-108∞03¢37.2¢¢W, 42 m depth, 20.xi.2002. LEB-ICML-UNAM-841, Altata 4 (by trawling, Sinaloa), 24∞26¢20.4¢¢N-107∞56¢32.64¢¢W, 32 m depth, 16.xi.2002. LEB-ICML-UNAM-843, Altata 1 (by trawling, Sinaloa), 24∞24¢19.92¢¢N- 107∞53¢17.58¢¢W, 31 m depth, 25.xi.2002. LEB-ICML- UNAM-933, Isla Hermano Sur (Mazatlán, Sinaloa), 23∞10¢59¢¢N-106∞26¢24.1¢¢W, 8 m depth, 22.x.2003. LEB-ICML-UNAM-895, Isla el Crestón (Mazatlán, Sinaloa), 23∞11¢02¢¢N-106∞25¢37¢¢W, 7 m depth, 12.ix.2003. LEB-ICML-UNAM-958, Isla Hermano Norte (Mazatlán, Sinaloa), 23∞11¢16.2¢¢N- 106∞25¢11.5¢¢W, 8 m depth, 24.x.2003. LEB-ICML- UNAM-995, Isla Cardones (Mazatlán, Sinaloa), 23∞11¢05¢¢N-106∞24¢07¢¢W, 6 m depth, 26.xi.2003. LEB-ICML-UNAM-1008, Cerro Pelón (Isla Isabel), 21∞51¢21¢¢N-105∞53¢33¢¢W, 20 m depth, 10.xii.2003. LEB-ICML-UNAM-1014, Cerro de la Cruz ( Isla Isabel), 21∞50¢32¢¢N-105∞52¢58¢¢W, 13 m depth, 11.xii.2003. CNPGG-93, Golfo de Tehuantepec , 15∞36¢0¢¢N-94∞04¢0¢¢W, i.1989. CNPGG-579, Bahía Petacalco (Guerrero), 17∞52¢12¢¢N-102∞12¢48¢¢W, 37 m depth, 11.ii.1982. CNPGG-642, Bahía de los Ángeles (Baja California), 19.ii.1999. CNPGG-682, Isla Coronado (Baja California), 19.ii.1999. CNPGG 698 ( Michoacán ), 18∞40¢12¢¢N-103∞46¢12¢¢W, 30 m depth, 10.i.1983. CNPGG-699, Bahía Concepción ( Baja California Sur ), 26∞35¢0¢¢N-111∞47¢0¢¢W, 10.ii.1999. CNPGG-700 ( Guerrero ), 17∞12¢30¢¢N-100∞55¢12¢¢W, 20 m depth, 20.iv.1982. CNPGG-701, La Bufadora ( Ensenada , Baja California ), 11.v.2001. CNPGG-702 (Guerrero), 16∞35¢18¢¢N-99∞07¢0¢¢W, 25 m depth 15.ii.1982. CNPGG-703 (Guerrero), 17∞55¢30¢¢N- 102∞01¢12¢¢W, 46 m depth, 23.iv.1982.

Description: Three typical growth stages of C. californiana (alpha, beta and gamma) have been found ( Fig. 6 ). The alpha stages (papillae) are pale yellow (dark brown or pale yellow in alcohol) ( Fig. 6A ). Their ostial papillae are more frequently circular, 0.2– 3.4 mm in diameter, and 0.5–(0.9)- 1.8 mm apart. They protrude 1–5 mm above the sponge surface. Oscular papillae are less abundant than ostial papillae; they are also circular, slightly elevated, and open to 0.7– 2.3 mm in diameter. Fusions of papillae have not been observed. The alpha stage is frequently boring into bivalve shells and rocks, reaching sizes up to 8.5 ¥ 14 cm. The beta stage overgrows the surface of the substrate reaching a maximum coverage of 9 ¥ 12 cm, and the specimens exhibited no tendency to undergo papillary fusion ( Fig. 6B ). The gamma stage has a characteristic mammillate surface with mainly circular papillae, uniformly distributed, about 1– 2 mm apart. They protrude above the substratum surface. Gamma specimens can grow up to 1 m in length, 33 cm high, and 55 cm wide (at the base). In this case, oscular papillae are frequently at the top of the sponge, and the ostial papillae are scattered on the lateral sides. Specimens are globular or semiglobular (4.8 ¥ 4.4 ¥ 1.5 cm), and volcano-shaped specimens are also common (22 ¥ 25 ¥ 13 cm) with an inner diameter of 8 cm. In the latter form, the ostial papillae are mainly located on the external surface and the oscular papillae are in the interior of the volcano. The ostial papillae are more frequently circular, very homogeneous in size, 0.75–(2)- 4.5 mm in diameter, and they are spaced 5–(1.4)- 2.2 mm from each other, without a tendency to fuse. The edges of the papillae are raised to about 5–(1.47)- 2.5 mm above the substratum. In preserved specimens papillae protrude from the body of the sponge, but in some specimens preserved by freezing the papillae are very often sinking, forming openings of 0.75–(2.3)- 4.5 mm in diameter. The oscular papillae are less abundant than ostial ones, they are also circular-shaped, and they can reach up to 10 mm in diameter. The surface is smooth, but the spicules protrude the surface by up to 70 Mm. The ectosome is not detachable, from 0.9 to 1.5 mm thick. In the choanosome there are canals from 0.3 to 1 cm wide. The consistency is firm, slightly compressible. The colour in the alpha and beta stages is golden yellow. In the gamma stage it is golden yellow, light pink, reddish brown and ochre ( Fig. 6C, D ). In some specimens the colour is not uniformly distributed, presenting both yellow and light pink areas. Alcohol turns them very dark brown.

Skeletal characters. The spiculation is formed exclusively by slightly curved tylostyles ( Fig. 7 ), with a well-differentiated globular head, mucronate or ovoid, sometimes with an apical knob. The pointed end is thin. The tylostyles in the gamma stage are characterized by having a shaft that is thinner in the middle part than at the extremes. They are straight or slightly curved with a well differentiated head, sometimes they are also mucronate. They measure 256 ¥ 7.8 Mm on average (8.3 Mm head width). Styles appear very occasionally ( Table 3). The ectosome has the typical clionaid tylostyle palisade that is 0.75– 1.57 mm wide. The choanosomal skeleton is made of ascending tracts of tylostyles (120–210 Mm thick), branching and anastomosing towards the ectosome from the base of the sponge ( Fig. 8 ).

Distribution: Sea of Cortez ( Dickinson, 1945; Hofknecht, 1978; as Pseudosuberites pseudos , and

A B C

Green & Gómez, 1986 as Cliona celata ) ( Fig. 1 ). California ( de Laubenfels, 1932 as C. celata var. californiana ). Baja California, Sonora, Sinaloa, Nayarit, Jalisco, Guerrero and the Gulf of Tehuantepec. This is the most typical sponge species in the north of the Sea of Cortez due to its abundance and its large size .

Remarks: A lot of specimens of a Cliona species very close to C. celata in form and colour were collected along the Mexican Pacific coast. In the north-east Pacific coast these specimens are commonly assigned to C. celata var. californiana de Laubenfels 1935 ( Sim & Bakus, 1986) or to C. celata ( Green & Gómez, 1986) . We compared our specimens with those from localities of Cliona celata along the east Atlantic and Mediterranean Sea coasts ( Rützler, 1973; Carballo et al., 1994; Carballo, 1994; Bavestrello, Calcinai & Sarà, 1995), and found important differences between them, mainly in the size of the megascleres ( Fig. 9 ). The tylostyles from Pacific specimens are significantly shorter (P <0.0001, F -ratio 30.04) and wider (P <0.0001, F -ratio 66.98), than those from the Mediterranean. Moreover, Pacific specimens have better formed termi- East Atlantic and Pacific Mediterranean specimens specimens

East Atlantic and Pacific Mediterranean specimens specimens nal heads, although small styles/subtylostyles with a slight swelling or knob that can be displaced along the shaft, are present in some specimens. On the basis of our results, we propose that Cliona celata var. californiana be considered as a valid species different from Cliona celata . Specimens at gamma stage are generally found in deeper waters than those at alpha stage (Carballo, 1994), but we have found important populations of Cliona californiana in small coastal lagoons where the alpha, beta and enormous gamma specimens appear simultaneously in shallow waters (from 0.5 to 1.5 m depth). This feature is very interesting and led us to clearly determine and compare the characteristics of the three growth forms in this species. A similar situation has been described along the northern coast of Spain ( Rodriguez-Solórzano, 1990). The specimens largely considered as Pseudosuberites pseudos Dickinson, 1945 in the Gulf of California matched perfectly with the gamma stage of the Cliona californiana . In areas where this species is very abundant we have observed massive golden yellow, brown, pink and salmon-coloured specimens growing together with yellow and pinkish brown specimens. The spiculation (size and form of the tylostyles) of all these specimens was similar and of the same characteristics as the holotype of Pseudosuberites pseudos . In fact, dark brown and dark yellow specimens of C. celata var. californiana have been described from the central California coast ( Hartman, 1975). Consequently, on the basis of both external and skeletal features, P. pseudos is here considered synonymous with C. californiana . The species Pseudosuberites pseudos was considered a synonym of Pseudosuberites melanos de Laubenfels 1934 by Desqueyroux-Faúndez (1972), and more recently of Cliona chilensis Thiele, 1905 by Desqueyroux-Faúndez & Soest (1997). Cliona chilensis has spirasters ( Thiele, 1905; Burton, 1932; Bergquist, 1968; among others), a characteristic missing from our specimens and the holotype. Moreover, other differences exist between the specimens assigned previously to C. pseudos and C. chilensis , in colour, surface, tylostyle measurements, geographical location and habitat.

CLIONA RAROMICROSCLERA DICKINSON, 1945 COMB. NOV.

( FIGS 2 , 10 )

Synonymy: Delaubenfelsia raromicrosclera Dickinson, 1945: 34 .

Material examined: MNCN-1.01/238, Ensenada de Bacochibampo (Guaymas, Sonora), 27∞54¢37¢¢N- 110∞57¢12¢¢W, 5 m depth, 06.xi.2000. LEB-ICML- UNAM-284, Ensenada de Bacochibampo (Guaymas, Sonora), 27∞54¢37¢¢N-110∞57¢12¢¢W, 5 m depth, 06.xi.2000. LEB-ICML-UNAM-302, Isla del Peruano (Guaymas, Sonora), 27∞54¢35¢¢N-110∞58¢17¢¢W, 15 m depth, 03.xi.2000. LEB-ICML-UNAM-734, Paraje Viejo (Guaymas, Sonora), 27∞52¢20¢¢N-110∞52¢08¢¢W, 5 m depth, 26.xi.2002. LEB-ICML-UNAM-746, Paraje Viejo (Guaymas, Sonora), 27∞52¢20¢¢N-110∞52¢08¢¢W, 5 m depth, 26.xi.2002. LEB-ICML-UNAM-755, Islas Gringas (Guaymas, Sonora), 27∞53¢05¢¢N- 110∞57¢55¢¢W, 10 m depth, 27.xi.2002. CNPGG-63, Bahía Concepción (Baja California Sur), 26∞35¢N- 111∞47¢W, 30.x.1998. CNPGG-237, Puerto Libertad (Sonora), 29∞59¢N-112∞45¢W, intertidal 11.x.1985. CNPGG-400, Santispac (Bahía Concepción, Baja California), 26∞35¢N-111∞52¢W. 30.x.1998. CNPGG-640, La Silica (Bahía de los Ángeles, Baja California), intertidal.

Description: Thickly encrusting to massive sponge (0.5–4 cm thick), covering a maximum area of 10 ¥ 15 cm ( Fig. 2E, F ). The surface is smooth or wrinkled in some parts, but it can be irregularly tuberculate or verrucose. The oscules are circular or oval, from 2 to 5 mm in diameter. These are elevated up to one millimetre from the surface. The choanosome presents abundant calcareous debris (mollusc shells, tubes of polychaetes, barnacles) and sand. The consistency is slightly compressible but firm. Colour in life is orange and olive green, and pale orange, beige or pink in alcohol.

Skeletal characters: The species has tylostyles and spirasters ( Fig. 10 ). The tylostyles are straight or slightly curved, with a generally round, oval or malformed head with annular swellings. They are usually bent in the upper third of their length. They measure 341 ¥ 9 Mm on average (12.2 Mm of head width) ( Table 4). Reduced tylostrongyles are also common in some specimens; they are straight, thick and with a round head, sometimes with incipient heads in the distal extreme ( Fig. 10B ). Tylostrongyles measurements are (as length ¥ width shaft; head width): 80–(126)-181 ¥ 16.5–(20.7)-24 Mm; 19.5–(25)- 30 Mm. The spirasters are sinuous or straight, and some are like anthosigmas (20 Mm average length). The spines are not pointed, but branching to two, three or even more spines. In the choanosome the tylostyles are arranged in vague tracts or in ascending bundles 130–(232)-350 Mm in diameter, which brush (325–425 Mm wide) out toward the surface in a palisade ( Fig. 11 ). The spirasters are scattered throughout the choanosome, and absent in the ectosome.

Distribution: Gulf of California ( Dickinson, 1945). Baja California, Baja California Sur and Sonora (present study).

Remarks: There is no other species similar to C. raromicrosclera . Anthosigmella vagabunda Ridley sensu de Laubenfels, 1954 from the Central Pacific, has ramose processes, it is spongy, and has different spicule sizes: tylostyles 500–600 ¥ 8–27 Mm, ‘C’ shape spirasters measure 13–20 Mm. We found three species with reduced tylostrongyles: C. desimoni Bavestrello et al. (1995) , C. argus var. laevicollis Thiele 1898 , and C. ensifera Sollas, 1878 . In C. desimoni megascleres are smaller in size, tylostyles are 110–(165)-225 ¥ 5– (7.8)-11 Mm, reduced tylostrongyles are 45–145 ¥ 7– 19 Mm, but the spiraster is a straight microrhabd (13– 25 Mm). In C. argus var. laevicollis the spiculation is larger; tylostyles measure 400–500 ¥ 18 Mm, tylostrongyles measure 240 ¥ 5 Mm, and the spiraster type has a verrucose surface instead of slender spines. C. ensifera has several malformations on the tylostyles different from the reductions of our species, besides smaller measurements and a different spiraster shape. These characteristics support the separation between them.

Cliona raromicrosclera has been also cited in Japan ( Hoshino, 1981). The spicule complements, the measurements and the morphological characteristics of the Japan specimens match the Mexican material closely. However, we do not consider those registers valid; the reduced tylostrongyles were not found in Japan specimens, and moreover, C. raromicrosclera has a typical distribution in the inner part of the Gulf. In spite of having sampled in a lot of localities along the Mexican Pacific Ocean, we have not found that species out of their typical area of distribution.

CLIONA AMPLICAVATA RÜTZLER, 1974

( FIGS 12 , 13 )

Synonymy: Cliona amplicavata Rützler, 1974: 26 ; Cliona amplicavata Rosell & Uriz, 2002: 68 .

Material examined: MNCN-1.01/308, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 5 m depth, 19.ix.2001, boring into a bivalve shell ( Arcidae ). LEB-ICML-UNAM-89, Isla de la Peña (Guayabitos, Nayarit), 21∞32¢53¢¢N-105∞17¢59¢¢W, 14 m depth, 23.xi.1999. LEB-ICML-UNAM-103, Isla de la Peña (Guayabitos, Nayarit), 21∞32¢53¢¢N-105∞17¢59¢¢W, 14 m depth, 11.xi.1999. LEB-ICML-UNAM-201, Estero El Zacate (Los Mochis, Sinaloa), 25∞36¢25¢¢N- 109∞04¢33¢¢W, 2 m depth, 21.vi.2000. LEB-ICML- UNAM-240, Isla Patos (Bahía de Ouhira, Topolobampo, Sinaloa), 25∞37¢12¢¢N-109∞00¢56¢¢W, 2 m depth, 22.vii.1999, on mollusc shell. LEB-ICML- UNAM-254, Isla de la Peña (Guayabitos, Nayarit), 21∞32¢53¢¢N-105∞17¢59¢¢W, 14 m depth, 23.vii.2000. LEB-ICML-UNAM-286, Ensenada de Bacochibampo (Guaymas, Sonora), 27∞54¢37¢¢N-110∞57¢12¢¢W, 5 m depth, 06.xi.2000, boring into a bivalve shell. LEB- ICML-UNAM-290, Isla El León Echado (Guaymas, Sonora), 27∞55¢34¢¢N-110∞57¢12¢¢W, 17 m depth, 06.xi.2000. LEB-ICML-UNAM-319, Isla Peruano (Guaymas, Sonora), 27∞54¢35¢¢N-110∞58¢17¢¢W, 15 m depth, 03.xi.2000. LEB-ICML-UNAM-348, Punta Cazón (Kino, Sonora), 28∞52¢20¢¢N-112∞02¢01¢¢W, 2 m depth, 28.iv.2001. LEB-ICML-UNAM-365, Isla Lobos 1 (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 5 m depth, 19.ix.2001, boring into an Arcidae bivalve shell. LEB-ICML-UNAM-461, Isla Redonda (Islas Marietas, Nayarit), 20∞42¢04¢¢N-105∞33¢89¢¢W, 10 m depth, 05.iv.2002. LEB-ICML-UNAM-484, Antiguo Corral del Risco (Punta de Mita, Nayarit), 20∞46¢20¢¢N- 105∞32¢49¢¢W, 4 m depth, 06.iv.2002. LEB-ICML- UNAM-582, Tecuchitan (Bahía de Banderas, Nayarit), 20∞43¢54¢¢N-105∞24¢44¢¢W, 10 m depth, 05.x.2002. LEB-ICML-UNAM-598, Isla redonda (Islas Marietas, Nayarit), 20∞42¢04¢¢N-105∞33¢89¢¢W, 9 m depth, 05.iv.2002, near the entrance of a small cave.

Description: Species found growing in alpha stage ( Fig. 12A ). Papillae are regularly distributed on the surface, circular in form, and protruding 0.1– 0.5 mm from the substrate. Papillary fusion is not frequent. The ostial papillae are 1.45–3.5 mm in diameter, and are more abundant than oscular papillae (0.2–0.7 mm in diameter). Colour alive is bright yellow.

Skeletal characters: The species presents tylostyles and rhaphides ( Fig. 13 ). The tylostyles are usually curved, with the shaft slightly bent in the upper third. They measured 266 ¥ 7 Mm on average. The heads are typically mucronated (8.5 Mm in average). In some specimens there is a small percentage of styles. The rhaphides are slender and slightly curved in the middle (Table 5).

142.5 142.5 132.5 110 -) 182.5 137.5

-) -) 112.5 105 117.5 - -) - 140) - 147.5)

Rhaphides - 80 (95) 110 – -100 85) (93 – (120 – 133.3 125 112.5 (– - 80 – () 95.8 - 92 –() 80 - 93) (80 – - 106 (90) – – () 92 101.8 – (137.5 154.3 – (112.5 82.5 87.5 – (120.3 – 117.5 97.5 (- – 87.5 96.8) 110 (means -) 5 4.1 - 5 - 5 -) 5 are (– 3.8) 4.1 4 () – 4.1 (– parentheses - ¥ 197.5) ¥ - 3.8– (185 ¥ - 162.5) 3 ¥ - 2.5) 180 in 172.6) 169 154.5 164.8 Values Styles (– 162 – (150 – 140 ((– 130.

specimens amplicavata -) 12.5 - 12.5 - 10) - 7.4 – ()

12.5 - 9.9 12.5) - 12.5) (9 - 8) 12.5 - 15 () 9.9 – 10 - 7.5) - 8.8) Cliona of;

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9.6 5) -)

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10.5

–

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5

–) 8

;

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-

5– 7.8; 8.8 (-

6.7

; 2.5 10) (– 3.8;

12.5 -

; – 6.3 12.5

-

; 3.8 –

10

(- 3.8

7.4

;) 10 - 7) –; 3 (

7.5 -

3.8 (

5

6.5 –;

7.5

; 7.5 6.3 (– - –

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10) 8.4

10

5 (

) 7.5) ()) () M

m

in

(

- (7 – 3.8) -) – (7.3 - 9 6 () 7; - 6 –) (¥ 2.5 – 7 (¥ 1.3 – -) 9.2 – (– (8.5 2.5 (2.5 7.3 – ¥ 3.8 – - 5.8) (2– – 1.3 4.4 (- (–) 5 6.5 - – 5 () 7.1

dimensions ¥ - 251.5) 285 - ¥) 305 3.8 - ¥) – 3 315 ¥ - 300) 2.5 -) 249.8 322.5 - 248.6 307.5) - ¥ 3.8 350) ¥ -) 325 258.8 - ¥) 357 271.3 - 257.3) 342.5 - ¥) 295 253.3 ¥ - 197.3) 245 - ¥) 290 271.5 ¥ -) 302.5 spicule Tylostyles 165 (– – 140 257 (200 273 (– 160 (– 240 (– 180 – 131.3 (268 150 (– – (165 – 155 (152.5 –(172 – ((145 – – 240 (– (216 180 the for data

Comparative

Table

.

5

LEB-ICML-UNAM- 89 103 LEB-ICML-UNAM- 201 LEB-ICML-UNAM- 240 LEB-ICML-UNAM- LEB-ICML-UNAM- 254 LEB-ICML-UNAM- 286 LEB-ICML-UNAM- 290 LEB-ICML-UNAM-319 348 LEB-ICML-UNAM- LEB-ICML-UNAM- 365 461 LEB-ICML-UNAM- 484 LEB-ICML-UNAM- 582 LEB-ICML-UNAM- 598 LEB-ICML-UNAM-

Distribution: Caribbean ( Rützler, 1974), Mediterranean Sea (Rosel & Uriz, 2002), and east Pacific Ocean (present study).

Remarks: Four Cliona species with rhaphides are currently known: C. radiata Hancock, 1849 ; C. linearis Sollas, 1878 , C. raphida Boury-Esnault, 1973 , and C. amplicavata Rützler, 1974 . Cliona radiata Hancock, 1849 (originally described without rhaphides), and C. linearis Sollas, 1878 were synonymized with C. celata by Topsent (1900). However, Rützler & Stone (1986) found rhaphides in C. radiata after revision of the type material and considered it a good species. In the same way, Dendy (1921) suggests the possibility that C. linearis is a species different from C. celata . The other species with rhaphides, C. raphida Boury- Esnaut, 1973, was described from a unique specimen in the Atlantic coast of South America, and we do not know of any record after that. However, Cliona raphida has tylostyles similar in size (260–355 ¥ 6– 9 Mm), but smaller rhaphides (32–62 ¥ 0.5 Mm) than C. amplicavata . Cliona amplicavata was described in the Caribbean by Rützler (1974), and later cited in the Mediterranean Sea by Rosell & Uriz (2002), who considered this species to be a synonym of C. linearis . However, these authors considered C. linearis as a nomen dubium, giving validity to C. amplicavata . Our specimens are in perfect agreement with the Caribbean specimens in form, size of the tylostyles (190– 290 Mm in length and 5–8 Mm in width) and rhaphides (117–150 Mm), and in the sporadic presence of small styles ( Rützler, 1974). In contrast, the Mediterranean specimens have larger tylostyles (230–460 in length and 6–12 Mm in width) and rhaphides (125–250 Mm), and the small styles are not present.

Besides these species, specimens with rhaphides assigned to C. celata are known in the Indian Ocean ( Dendy, 1921; Topsent, 1932; Thomas, 1973), Gulf of Mexico ( Topsent, 1887), and off Australia ( Schönberg, 2000); these would need to be reviewed to clarify their taxonomic status.

CLIONA FLAVIFODINA RÜTZLER, 1974

( FIGS 14 , 15 )

Synonymy: Cliona flavifodina Rützler, 1974: 9 .

Material examined: MNCN-1.01/309, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 5 m depth, 24.iv.2003. LEB-ICML-UNAM-373, Punta Chile (Mazatlán, Sinaloa), 23∞12¢29¢¢N-106∞25¢40¢¢W, intertidal, 08.x.2001. LEB-ICML-UNAM-376, Cerritos 2 (Mazatlán, Sinaloa), 23∞18¢51¢N-106∞29¢31¢W, 1.5 m depth, 10/15/01. LEB-ICML-UNAM-555, Isla Lobos 1 (Mazatlán, Sinaloa), 23∞13¢49¢¢N- 106∞27¢43¢¢W, 5 m depth, 24.iv.2003. LEB-ICML- UNAM-654, Isla Pájaros (Mazatlán, Sinaloa), 23∞15¢29¢¢N-106∞28¢25¢¢W, 5 m depth, 31.x.2002. LEB- ICML-UNAM-912, Isla Lobos 2 (Mazatlán, Sinaloa), 23∞13¢27.7¢¢N, 106∞28¢01.6¢¢W, 7 m depth, 03.x.2003.

Description: This species tends to overgrow the substrate, covering a maximum surface of 8 ¥ 14 cm ( Fig. 14A, B ). The papillae (oscular and inhalant) are scarce and irregularly distributed over the surface. They are well-spaced (from 0.25 to 1.6 cm apart), measuring from 1.4 to 5 mm in diameter, and without a tendency to fuse. They are mainly circular, and when the sponge is alive the papillae protrude on the substrate from 1.5 to 5 mm. The colour is commonly purple brown, although golden-yellow specimens also appear. The choanosome is yellowish brown; after fixation the surface and choanosome turn brownish.

Skeletal characters: Species having tylostyles and spirasters ( Fig. 15 ). The tylostyles are mostly straight or slightly bent, with the shaft characteristically widest near the midregion. The point is sharp. They measure 295 ¥ 7.5 Mm on average. The heads are well set off and usually spheroid (average 10.8 Mm) ( Table 6). The spirasters (average 35 Mm) are very characteristic, with three or four bends and with few and slender prominent spines. The actins are robust, single or bifurcated, and end in a sharp point. Tylostyles are in loose bundles in the choanosome. In the ectosome, the tylostyles form a compact palisade.

Distribution: The species was described from Bermuda (Caribbean Sea) ( Rützler, 1974). This constitutes the first record from the Pacific Ocean, and it is the second time that the species has been found. It is distributed from the intertidal zone to 12 m depth, on dead corals, pelecypod shell and rocks. The species has been found alive in the intertidal zone during low tide.

Remarks: Our specimens agree with the description of Cliona flavifodina in Bermuda by Rützler (1974). However, they differ in the shape of the tylostyle heads, which are mainly rounded in the Pacific specimens and droplet-shaped or ovoid in the Caribbean, sometimes with secondary swellings. Moreover, the spirasters have singular spines in the Caribbean but they can bifurcate in the Pacific specimens. Cliona aprica Pang, 1973 from Jamaica is another similar brown-black sponge with a tendency to overgrow the substrate. However, the spirasters in this species possess shorter spines (1–2 Mm in length), and they are located mainly in the convex side of the axis of the spiraster. Cliona langae Pang, 1973 has spirasters with rather widely spaced spines like C. flavifodina , and they also tend to overgrow the substrate completely ( Pang, 1973). However, this species differs from Cliona flavifodina in spicular dimensions and in the form and spination of the spirasters. Recently, C. langae and C. aprica have been synonymized with C. caribbaea by Rützler (2002b).

CLIONA EURYPHYLLA TOPSENT, 1887

( FIGS 14 , 16 , 17 )

Synonymy: Cliona euryphylla Topsent, 1887: 82 ; Cliona euryphylla de Laubenfels, 1954: 218 ; Cliona euryphylla Bergquist, 1968: 30 .

Material examined: MNCN-1.01/310, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 09.v.2003 . LEB-ICML-UNAM-123, Cerritos (Mazatlán, Sinaloa), 23∞18¢27¢¢N-106∞29¢25¢¢W, Intermareal, 18.ii.2000, on rocks. LEB-ICML-UNAM-276, Isla San Pedro Nolasco (Sonora), 27∞57¢24¢¢N- 111∞22¢34¢¢W, 20 m depth, 15.xi.2000 . LEB-ICML- UNAM-313, 02/20/01, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 20.ii.2001 . LEB- ICML-UNAM-323, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 20.ii.2001 . LEB- ICML-UNAM-355, Cerritos 2 (Mazatlán, Sinaloa), 23∞18¢51¢N-106∞29¢31¢W, 1.5 m depth, 28.v.2001. LEB- ICML-UNAM-366, Isla Lobos (Mazatlán, Sinaloa), 23∞13¢49¢¢N-106∞27¢43¢¢W, 6 m depth, 18.ix.2001 . LEB-ICML-UNAM-431, Isla Pájaros (Mazatlán, Sinaloa), 23∞15¢29¢¢N-106∞28¢25¢¢W, 4 m depth, 15.ii.2002 . LEB-ICML-UNAM-443, Antiguo muelle de atraque (Mazatlán, Sinaloa), 23∞11¢57¢¢N- 106∞25¢15¢¢W, 4 m depth, 24.i.2002. LEB-ICML- UNAM-445, Isla Pájaros (Mazatlán, Sinaloa), 23∞15¢29¢¢N-106∞28¢25¢¢W, 4 m depth, 26.ii.2002 . LEB- ICML-UNAM-481, Careyeros (Punta Mita, Nayarit), 20∞47¢13¢¢N-105∞71¢13¢¢W, 4 m depth, 6.iv.2002. LEB- ICML-UNAM-498, Los arcos (Puerto Vallarta, Jalisco), 20∞32¢73¢¢N-105∞18¢47¢W, 4 m depth, 04.viii.2002. LEB-ICML-UNAM-521 Playa los Muertos (Sayulita, Nayarit), 20∞52¢29 N-105∞26¢72¢¢, 5 m depth, 04.ix.2002. LEB-ICML-UNAM-522, Los Arcos (Puerto Vallarta, Jalisco), 20∞32¢73¢¢N-105∞18¢47¢W, 2 m depth, 10.iv.2002. LEB-ICML-UNAM-592, Tecuchitan (Bahía de Banderas, Nayarit), 20∞43¢54¢¢ N-105 24 ¢44¢¢W, 10 m depth, 05.x.2002 . LEB-ICML-UNAM- 625, Conchas Chinas (Puerto Vallarta, Jalisco), 20∞35¢16¢¢N-105∞14¢42¢¢W. 2 m depth, 08.x.2002. LEB- ICML-UNAM-641, Playa los Muertos (Sayulita, Nayarit), 20∞52¢29 N-105∞26¢72¢¢, 8 m depth, 09.x.2002. LEB-ICML-UNAM-433, Isla Pájaros 1(Mazatlán, Sinaloa), 23∞15¢29¢¢N, 106∞28¢25¢¢W, 10 m depth, 15.ii.2002 . LEB-ICML-UNAM-854, Majahuita (Jalisco), 20∞29¢6.66¢¢N, 105∞35¢3.42¢¢W, 7 m depth, 08.vi.2003. LEB-ICML-UNAM-914, Isla Lobos 2 (Mazatlán, Sinaloa), 23∞13¢27.7¢¢N, 106∞28¢01.6¢¢W, 7 m depth, 03.x.2003 . LEB-ICML-UNAM-959, Isla Hermano Norte (Mazatlán, Sinaloa), 23∞10¢59¢¢N, 106∞26¢24.1¢¢W, 8 m depth, 24.x.2003 . LEB-ICML- UNAM-1010, Cerro de la Cruz ( Isla Isabel), 21∞50¢32¢¢N-105∞52¢58¢¢W, 13 m depth, 11.xii.2003 .

Description: The most characteristic trait of this species is that papillae are surrounded by and connected with patches of ectosomal tissue. The sponge typically grows in alpha stage, but it is able to spread over the substratum by lateral growth of the ectosome next to the papillae, reaching a beta stage (from 1 to 3 mm thick) ( Fig. 14C, D, E ). The size of the specimens reaches up to 12 ¥ 23 cm coverage. Specimens growing in beta stage are very irregular in form. Fusions between papillae are not common. The papillae can be circular and oval, from 3 to 6 mm in diameter, and they can protrude over the substrate up to 3 mm in a live sponge. They are very irregularly distributed, and can be spaced up to 12 mm. Oscules are usually circular, up to 2 mm in diameter, and can protrude over the substrate up to 5 mm. The consistency is firm. The papillae range from yellow to light orange in life, and turn pale in alcohol. The orange colour is more typical in the beta specimens.

Skeletal characters: The megascleres are tylostyles, each with well-formed rounded head ( Fig. 16A, D ). They measure 249 ¥ 7.3 Mm on average (9.7 Mm head width) ( Table 7). The microscleres are short, stout spirasters, with many heavy spines uniformly distributed along the shaft ( Fig. 16B, E ). Short amphiasters sometimes appear. The mean length of the spirasters is 18 Mm. The skeleton has the typical arrangement of Cliona species , with a palisade of tylostyles in the periphery of the sponge, and tylostyles in disorganized bundles in the choanosome ( Fig. 17 ). Sometimes tylostyles are disposed in tangential form in the ectosome. Differences in the skeletal characters were not found between the orange and yellowish specimens.

Distribution: Gulf of Mexico (Atlantic Ocean) ( Topsent, 1887), New Zealand ( Bergquist, 1968), central Pacific ( de Laubenfels, 1954). On rocks, boring into shells of bivalves, etc. These are the first records in the east Pacific.

Remarks: Cliona euryphylla is included in the group of Cliona species with thick and short spirasters like the species C. chilensis Thiele, 1905 , C. burtoni Topsent, 1932 and C. aethiopicus Burton, 1932 ; which are considered conspecific species by de Laubenfels (1954). We agree with Bergquist (1968) and consider the four species valid on the basis of different characteristics, mainly the spiraster sizes ( Thiele, 1905; Burton, 1932). Cliona euryphylla Topsent (1887) was described in Campeche (Gulf de Mexico) in the Atlantic coast, and later was cited by de Laubenfels (1954) in the central Pacific, and by Bergquist (1968) in New Zealand. De Laubenfels (1954) and Bergquist (1968) mistakenly thought that Topsent (1887) had described this species in the eastern Pacific Ocean. However the measurements given by Bergquist (1968) are larger (344 ¥ 12.5 Mm) than our specimens which match very well with the tylostyle measurements given by Topsent (300 ¥ 5 Mm), and by de Laubenfels (300 ¥ 7 Mm). The spirasters of our specimens are also smaller than those from Bergquist (24 ¥ 6.3 Mm) and Topsent (35 ¥ 5 Mm). The closest species is Cliona dioryssa ( de Laubenfels, 1950) , which has two types of spirasters: stout spirasters with a thick shaft densely set with large, strong spines, most of which are joined at the base, and more delicate spirasters with a long, slender shaft and small spines, which are usually well spaced ( Rützler, 1974). However, the SEM shows different fine details in the spiraster types, and in the size, which is slightly smaller in C. euryphylla .

Two different expressions of colour are found in C. euryphylla ; however, the morphology and size of the spicules are very similar in the two forms.

CLIONA VERMIFERA HANCOCK, 1867

( FIGS 12 , 18 )

Synonymy: Cliona vermifera Hancock, 1867: 239 ; Bernatia vermifera Rosell & Uriz, 1997: 352 .

Material examined: MNCN-1.01.311 Antiguo Corral del Risco (Punta de Mita, Nayarit), 20∞46¢20¢¢N- 105∞32¢49¢¢W, 4 m depth, 06.iv.2002, on dead coral. LEB-ICML-UNAM-364, Los Arcos (Puerto Vallarta, Jalisco), 20∞32¢73¢¢N-105∞18¢47¢W, 2 m depth, 23.i.2001, on dead coral. LEB-ICML-UNAM-470, Careyeros (Punta Mita, Nayarit), 20∞47¢13¢¢N- 105∞71¢13¢¢W, 4 m depth, 06.iv.2002, on dead coral. LEB-ICML-UNAM-477, Antiguo Corral del Risco (Punta de Mita, Nayarit), 20∞46¢20¢¢N-105∞32¢49¢¢W, 4 m depth, 06.iv.2002, boring into dead coral. LEB- ICML-UNAM-800, Isla Redonda (Islas Marietas, Nayarit), 20∞42¢04¢¢N-105∞33¢89¢¢W, 14 m depth, 10.vi.2003, boring into coral Pavona gigantea . LEB- ICML-UNAM-807, Isla Redonda (Marietas, Nayarit), 20∞42¢04¢¢N, 105∞33¢89¢¢W, 12 m depth, 10.iv.2003, boring into dead coral. LEB-ICML-UNAM-847, Majahuita (Jalisco), 20∞29¢6.66¢¢N, 105∞35¢3.42¢¢W, 10 m depth 08.vi.2003. LEB-ICML-UNAM-857, Antiguo Corral del Risco (Punta Mita, Nayarit), 20∞46¢20¢¢N-105∞32¢49¢¢W, 3 m depth 06/08/03. LEB- ICML-UNAM-865, Chacala (Nayarit), 21∞09¢57¢¢N- 105∞13¢38¢¢W, 4 m depth, 12.vi.2003. LEB-ICML- UNAM-1016, Cerro de la Cruz ( Isla Isabel), 21∞50¢32¢¢N-105∞52¢58¢¢W, 13 m depth, 11.xii.2003. LEB-ICML-UNAM-1040, Las Monas ( Isla Isabel), 21∞50¢58¢¢N-105∞52¢46¢¢W, 6 m depth, 10.xii.2003. LEB-ICML-UNAM-1049, Punta Bobo ( Isla Isabel), 21∞50¢35¢¢N-105∞52¢44¢¢W, 12 m depth, 9.xii.2003. CNPGG-666, Manzanillo (Zihuatanejo, Guerrero), 17∞37¢0¢¢N-101∞31¢0¢¢W, 3.5 m depth, 05.iii.1982. CNPGG-667, Contramar (Zihuatanejo, Guerrero), 17∞35¢0¢¢N-101∞32¢0¢¢W, 6 m depth, 17.viii.1982. CNPGG-672 (Guerrero), 17∞54¢24¢¢N-101∞53¢42¢¢W, 10 m depth, 13.ii.1982.

Description: Boring sponge in alpha stage, covering a maximum surface of 8 ¥ 11 ¥ 6 cm. Papillae are very small ( Fig. 12C, D ). Their shape is circular, from 0.3 to 1.2 mm, or oval, from 0.6 to 1.5 mm in diameter. They are regularly distributed, lying 0.1–6.5 mm apart. No fusion tendency was observed. Oscules have not been observed. Colour is orange in life, beige in alcohol.

Skeletal characters: The species have tylostyles and ‘smooth spirasters’ ( Fig. 18 ). The tylostyles are mostly straight with spheroid heads, sometimes mucronated, with hastate or acerate ends. They measure 201 ¥ 6 Mm on average (8.8 Mm head width) ( Table 8). The microscleres are smooth, spiral or undulated rods, with 2–4 bends and obtuse extremities. Ectosomal arrangement of tylostyles in a palisade, accompanied along the shafts by intermingled spirasters.

Distribution: Mediterranean Sea ( Vosmaer, 1933; Volz, 1939), east Atlantic Ocean ( Carballo et al., 1994), Caribbean ( Topsent, 1889; Hechtel, 1965; Rützler, 1974), east Pacific Ocean ( Salcedo et al., 1988), Indian Ocean ( Topsent, 1932). Mexican Pacific: Nayarit, Jalisco and Guerrero State (present study). There is a high incidence of the species in corals of the species Pavona gigantea . More than 70% of the specimens of this coral were found with C. vermifera ( Fig. 12D ).

Remarks: Volz (1939) and later Carballo et al. (1994) mistakenly thought that Carter (1882) had cited the species C. vermifera in Acapulco ( México) in the eastern Pacific Ocean. However, the only previous register of C. vermifera in the eastern Pacific Ocean is from Salcedo et al. (1988) in Guerrero ( Mexico).

Rosell & Uriz (1997) erected the genus Bernatia to harbour Cliona species with two size classes of tylostyles. However, this character could be insufficient to justify the formation of a new genus, because the presence of two size classes of tylostyles seems to appear in different boring sponge genera ( Schönberg, 2000). We accept the concept of the genus Cliona sensu Rützler (2002a) considering Bernatia a synonym of Cliona until the true importance of this character can be conveniently evaluated.

Figures

Figure 1. Location of the sampling stations along the coast of Mexico. The numbers show the distribution of the species: (1) Cliona papillae sp. nov.; (2) Cliona vallartense sp. nov.; (3) Cliona californiana comb. nov.; (4) Cliona raromicrosclera comb. nov.; (5) Cliona amplicavata; (6) Cliona flavifodina; (7) Cliona euryphylla; (8) Cliona vermifera; (9) Pione carpenteri; (10) Pione mazatlanensis; (11) Thoosa calpulli sp. nov.; (12) Thoosa mismalolli sp. nov.; (13) Spheciospongia ruetzleri sp. nov.; (14) Spheciospongia incrustans sp. nov.

Figure 2. External morphologies. A, Cliona papillae sp. nov., boring into rock and calcareous algae at 5 m depth. B, Detail of the papillae of Cliona papillae sp. nov. The arrow shows the fusion of oscular and ostial papillae. The typical sieve-like ostial papillae are visible to the eye underwater. C, Cliona vallartense sp. nov. growing in beta stage with scattered oscular papillae (arrow) at 4 m depth. D, Cliona vallartense sp. nov. growing in beta stage with numerous oscula (arrow) at 7 m depth. E and F, Cliona raromicrosclera comb. nov. massive orange and beige specimens.

Figure 3. Cliona papillae sp. nov.: A, heads and tips of the tylostyles, some showing annular swelling and a rounded end; B, different forms of spirasters; C, tylostyles.

Figure 5. Cliona vallartense sp. nov.: A, heads and tip of the tylostyles showing different modifications and secondary annular swelling; B, tylostyles; C, different forms of spirasters.

Figure 6. External morphology at different growth stages of Cliona californiana comb. nov. A, Specimen in alpha stage boring into a bivalve shell at 15 m depth. The arrow shows oscular papillae. B, Specimen in beta stage growing on rocks and calcareous algae, 5 m depth. C, Large, massive yellow specimen (gamma stage) at 1 m depth. D, Pink specimen growing in gamma stage, at 10 m depth. E, Specimen (C) out of the water. The arrows show collapsed papillae. F, Pink gamma stage with closed papillae.

Figure 7. Cliona californiana comb. nov.: tylostyles and detail of the heads in (A) alpha stage, (B) beta stage and (C) gamma stage. Same scale bar for tylostyles (in the centre) and for the heads (below).

Figure 8. Cliona californiana comb. nov.: skeletal arrangement close to the surface of a gamma stage.

Figure 9. Multiple ‘Box-and-Whisker’ plot for tylostyle shaft width (A), and length (B) for Mediterranean and Pacific specimens.

Figure 10. Cliona raromicrosclera comb. nov.: A, heads and tip of the tylostyles; B, small tylostrongyles with secondary swelling; C, tylostyle; D, different forms of spirasters.

Figure 11. Cliona raromicrosclera comb. nov.: cross section of the body close to the surface.

Figure 12. External morphology of A, Cliona amplicavata. The arrow shows an oscule. B, Pione mazatlanensis boring into different calcareous material. C, Cliona vermifera boring into the coral Pavona gigantea. The papillae are only visible in the base of the coral. D, Excavating pattern of the papillae can be observed after breaking a fragment of the coral. E, Spheciospongia incrustans sp. nov. at 5 m depth; the arrow shows an oscule. F, Spheciospongia incrustans sp. nov.; the arrows show the pore sieves.

Figure 13. Cliona amplicavata: A, mucronated heads and tip of the tylostyles - in some specimens there is a small percentage of styles; B, rhaphide and its magnification; C, tylostyles, styles and rhaphides.

Figure 14. External morphology of A, Cliona flavifodina; yellow specimen living in a small intertidal cornice. B, Cliona flavifodina at 5 m depth; arrow shows an oscule. C, Cliona euryphylla; Light orange specimen growing in alpha stage. Arrow shows ostial papillae. D, Orange Cliona euryphylla specimen; the arrow shows an ostial papillae surrounded by a patch of ectosomal tissue. E, Cliona euryphylla; light orange specimen growing in alpha stage at 6 m depth. Arrow shows an ostial papillae.

Figure 15. Cliona flavifodina: A, heads and tip of the tylostyles; B, different forms of spirasters.

Figure 16. Cliona euryphylla: A, heads and tip of the tylostyles; B, spirasters and D, tylostyles from orange specimens; C, heads and E, spirasters from yellow specimens.

Figure 17. Cliona euryphylla: Cross section of the body close to the surface.

Figure 18. Cliona vermifera: A, heads and tips of the tylostyles; B, smooth spirasters.