Neophrissospongia galapagoensis, Schuster & Cárdenas & Pisera & Pomponi & Kelly & Wörheide & Erpenbeck, 2018

NEOPHRISSOSPONGIA GALAPAGOENSIS View in CoL SP. NOV.

( FIGS 2–4; TABLE 1)

D i a g n o s i s: M a s s i v e e a r - s h a p e d, f l a b e l l a t e, Neophrissospongia with rhabdome of dichotriaenes showing a large variation from 172 to 617 µ m length with spined cladome and blunt to acute tips.

Type material: Holotype: HBOM 003 View Materials :02010, Coll. Johnson Sea-Link I ( JSL-I) dive 3923 [27 October 1995, Galápagos , 4 nautical miles ( NM) SE of Tortuga Island, 01°04 ′ 59 ″ S, 90°51 ′ 56 ″ W, 241 m] GoogleMaps . Paratypes: HBOM 003 View Materials :01097 (227 m) , HBOM 003 View Materials :01098 (189 m) , Galápagos, NE of Rocas Gordin, Santa Cruz Island, 0°32 ′ 53 ″ S, 90°7 ′ 47 ″ W, Coll GoogleMaps . JSL-I dive 3900. HBOM 003 View Materials :02002 (204 m) , HBOM 003 View Materials :02003 (120 m) , Galápagos, 5 NM NE of Kicker Rock at San Cristobal Island, 01°43 ′ 8 ″ S, 89° 27 ′ 14 ″ W, Coll GoogleMaps . JSL-I dive 3905. HBOM 003 View Materials :01099 (177 m) , Galápagos, N-Side 1.25 NM NW of Cormorant PT at the Santa Maria Island (Floreana), 01°12 ′ 6 ″ S, 90°26 ′ 10 ″ W, Coll GoogleMaps . JSL-I dive 3901. HBOM 003 View Materials :02008 (235 m) , Galápagos, West Coast 2 NM NW of PTA Tortuga at Isabela Island, 0°14 ′ 31 ″ S, 91°24 ′ 56 ″ W, Coll GoogleMaps . JSL-I dive 3915. HBOM 003 View Materials :02005 (328 m) , Galápagos, NW Coast, 66 NM offshore, Isabela Island , 0°8 ′ 35 ″ S, 91°23 ′ 42 ″ W, Coll GoogleMaps . JSL-I dive 3912.

Comparative material: Neophrissospongia nolitangere Schmidt, 1870 , HBOI 2-VI-91-1-005 (358 m), Portugal, Madeira, Savage Islands, Selvagem Grande, South Coast, 30°07 ′ 21 ″ N, 15°52 ′ 7.8 ″ W, Coll. JSL-I dive 3006, June 1991, identified by Michelle Kelly. Neophrissospongia microstylifera Lévi & Lévi, 1983 0CDN7083-J (111 m), Palau, Koror Reef, W-Side of Uchelbeluu Reef, wall, rock, 7°16 ′ 25 ″ N, 134°31 ′ 25 ″ E, 2001, identified by Michelle Kelly; 0CDN5043-Q (109 m), Palau, Koror Reef, W-Side of Uchelbeluu Reef, overhang, rock, 7°16 ′ 25 ″ N, 134°31 ′ 26 ″ E, 1997, identified by Michelle Kelly.

Type locality: Tortuga Island, Galápagos Islands (241 m) ( Fig. 1) .

D i s t r i b u t i o n: To r t u g a I s l a n d, I s a b e l a I s l a n d, Fernandina, Santa Cruz Island, San Cristobal, Floreana Island ( Fig. 1).

Habitat: Attached to hard substratum, depth range 120–328 m.

Description: Morphology, in the shape of an ear, flared cup or vase, or foliose with folded margins when mature; walls 1–1.5 cm thick, margins flattened with rounded edges ( Fig. 2); dimension of holotype, 12 cm diameter, 16 cm high ( Fig. 2A); paratypes range 20–30 cm diameter, 30–45 cm high or larger ( Fig. 2C, D). Surface, with horizontal growth lines; concave inhalant surface with numerous ostia, 25–35 µ m diameter; convex (exhalent surface) has numerous, barely visible, oscules. Texture, stony, especially towards the base of the sponge. Colour, beige in life, brown in ethanol preservative. Ectosomal skeleton, 1.2–1.5 mm thick, outermost surface a densely packed, 100- µ m-thick layer of microscleres, within which lies the cladomes of the dichotriaenes, the rhabdomes of which penetrate deeply into the choanosome ( Fig. 4A, B). Choanosomal skeleton composed of zygosed dicranoclone desmas and acanthose microtylostyles, often located around the pore openings. Megascleres, desmas and triaenes; dicranoclone desmas, 280– 450 µ m (N15) maximum diameter, with a massive, arched, central core, from which four to six arms splay away from the underside of the arch, both the central core and the arms are covered irregularly with tubercles ( Fig. 3B, C, F), the tubercules on the desma core are larger, fungiform and multituberculate; dichotriaenes, rhabdome 172– 422 –617 µ m (N30) long, cladome, 175– 227 –372 µ m (N30) wide, upper surface of clads irregularly spined ( Fig. 4G, H). Microscleres, irregular amphiasters with short, thick rays, 15.2– 11.2 –18 µ m long × 4.9– 7.2 –10.7 µ m (N25) wide; acanthose microtylostyles, 44– 91 –133 µ m (N25) long.

Etymology: Named after the locality and distribution of the species, the Galápagos Islands.

DNA barcodes: We sequenced partial COI, 28S (C1-D2) and complete 18S of the holotype and the paratypes from different localities of the Galápagos Islands ( Fig. 1). COI sequences of all type material sequenced were identical with the exception of HBOM 003 View Materials :02005 (no COI sequenced). 28S and 18S sequences of all type material were identical with the exception of HBOM 003 View Materials :02005 (no 18S sequenced). GenBank accession numbers: COI KY652805 View Materials – KY652811 View Materials ; 28S

KY652771 View Materials – KY652778 View Materials ; 18S KY652826 View Materials – KY652832 View Materials ; SBP No. 1702–1709.

Remarks: According to the World Porifera Database (WPD) ( Van Soest et al., 2017, http: //www. marinespecies.org/porifera/), only six different species of Neophrissospongia have been described: N. nolitangere ( Table 1; Pisera & Vacelet, 2011) and N.tubulata Van Soest & Stentoft, 1988 from the tropical Western Atlantic; N. endoumensis Pisera & Vacelet, 2011 , N. nana Manconi & Serusi, 2008 and N. radjae Pisera & Vacelet, 2011 , from the Mediterranean Sea, and N. microstylifera Lévi & Lévi, 1983 from New Caledonia in the Pacific Ocean.

Neophrissospongia galapagoensis sp. nov., which is cup-shaped to foliose, clearly differs from the encrusting N. nana , discovered in a Mediterranean marine cave. The club-shaped N. radjae ( Table 1; Pisera & Vacelet, 2011) from a cave in the Mediterranean with an apical oscule is also distinguished from our species by gross morphology. Minor differences between the dichotriaenes of N. galapagoensis sp. nov. and N. radjae ( Table 1) also distinguish both species: N. radjae has a more distinctly tuberculated cladome than in our new species.

Neophrissospongia nolitangere View in CoL , from the Azores, Cape Verde Islands, off Portugal (e.g. Pisera & Lévi, 2002a), Madeira ( Carvalho et al., 2015), submarine caves in the Mediterranean ( Table 1; Pisera & Vacelet, 2011) and the Bahamas ( Pomponi et al., 2001), is distinguished from N. galapagoensis View in CoL sp. nov. in the possession of smaller streptasters with longer rays; 28S sequences of N. nolitangere View in CoL by Chombard, Boury-Esnault & Tillier (1998) and here (HBOI 2-VI-91-1- 005) are both 100% identical and differ by 1% from our new species. Molecular analyses ( Fig. 5) further separate N. galapagoensis View in CoL sp. nov. from N. nolitangere View in CoL . Sequence comparison with material from the West Pacific (WAM Z35946, 36053; probably both new species, A. Pisera, personal observation) disclose a 1.6% difference.

Neophrissospongia tubulata View in CoL from Barbados is characterized by small, single or clustered tubes, in which the dichotriaenes have short rhabds (30–50 µ m) and a non-tuberculate cladome. Rather, the clads have dentate margins, clearly distinguishing them from the dichotriaenes of N. galapagoensis View in CoL sp. nov. ( Table 1) and all other members of the genus. This lateral dentation of the dichotriaene clads of N. tubulata View in CoL points to the need for a revision of this species.

The dichotrianes of N. endoumensis , from the Mediterranean (cladome width: 248–283 µ m; rhabdome length: 531–615 µ m) are larger and thicker and are less variable in size ( Table 1; Pisera & Vacelet, 2011) compared to those of N. galapagoensis sp. nov.

In the original description of tubular N. microstylifera from New Caledonia, Lévi & Lévi (1983) documented microstyles of 110–170 µ m length, which are absent in our new species, and the dichotriaenes are more robust than those of N. galapagoensis sp. nov.