taxonID	type	description	language	source
03B50F5C7106FF83800AFA4BFC50D6DE.taxon	materials_examined	Type species	en	Ruiz, César, Muricy, Guilherme, Lage, Anaíra, Domingos, Celso, Chenesseau, Sandrine, Pérez, Thierry (2017): Descriptions of new sponge species and genus, including aspiculate Plakinidae, overturn the Homoscleromorpha classification. Zoological Journal of the Linnean Society 179 (4): 707-724, DOI: 10.1111/zoj.12480, URL: https://www.mendeley.com/catalogue/813c5d34-fdc1-3687-80eb-affa77eab333/
03B50F5C7106FF83800AFA4BFC50D6DE.taxon	description	Plakinidae with spicules diods, triods and calthrops in a single size class, and with homolophose calthrops with one to four lophate rays (Muricy & Dıaz, 2002).	en	Ruiz, César, Muricy, Guilherme, Lage, Anaíra, Domingos, Celso, Chenesseau, Sandrine, Pérez, Thierry (2017): Descriptions of new sponge species and genus, including aspiculate Plakinidae, overturn the Homoscleromorpha classification. Zoological Journal of the Linnean Society 179 (4): 707-724, DOI: 10.1111/zoj.12480, URL: https://www.mendeley.com/catalogue/813c5d34-fdc1-3687-80eb-affa77eab333/
03B50F5C7106FF8082AFFE81FD12D201.taxon	description	FIGS 2, 3 Material examined Holotype: MNHN DJV 177, Grotte Chauve-Souris at 12 m depth, Anse Noire, La Martinique (14 ° 32.024 0 N, 61 ° 05.278 0 W). Collector: T. Perez, 10 December 2013. Slides and a fragment of the holotype were deposited in the sponge collection of the Museu Nacional of Universidade Federal do Rio de Janeiro, Brazil (MNRJ 18460). Paratype: MNHN DJV 178, Grotte Chauve-Souris at 11 m depth, Anse Noire, La Martinique. Collector: T. Perez, 6 March 2014. Other specimens examined 150530 - GU 6 - TP 04, Grotte Amedien at 12 m depth, La Guadeloupe (16 ° 30.033 0 N, 061 ° 28.774 0 W). Collector: T. Perez, 30 May 2015. 150516 - MT 8 - CR 03, Anse Fortune at 7 m depth, La Martinique (14 ° 30.377 0 N, 61 ° 05.850 0 W). Collector: C. Ruiz, 16 May 2015. Comparative material examined Plakina jamaicensis Lenhert & van Soest, 1998. RBINS POR 70, Chalet Caribe caves, West of Montego Bay, Jamaica (18 ° 27.246 0 N, 77 ° 58.287 0 W). Collector P. Willenz and A. Ereskovsky. Etymology: Plakina arletensis refers to the little village ‘ Les Anses d’Arlet’, close to the cave where this sponge was first found, which is one of the most beautiful places in La Martinique. This is also a special dedication to the kind and warm inhabitants of this village. Diagnosis: Thin encrusting and white Plakina, with a rugose folded surface, a skeleton made of monolophose, trilophose and tetralophose calthrops, actines with and without terminal spines. Diods and triods are absent. Well-developed mesohyl with a high abundance of prokaryotic symbionts. Description Small crust, about 2 – 15 cm in size, 0.2 – 0.5 cm thick. White colour in vivo, becoming slightly cream or light brown in alcohol (Fig. 2 A, B). The consistency is cartilaginous and the surface is rugose and irregularly folded. Oscules are circular, 1 – 2 mm in diameter, with a slightly elevated rim. Skeleton: The skeleton is dense, especially at the surface which is pierced principally by trilophose calthrops. The choanosomal skeleton has an alveolar arrangement formed by all spicule types (Fig. 2 C). Spicules: Diods and triods are absent. Calthrops are irregular, abundant, their actines (20 – 30 lm long) being with or without spines. Spines (1 – 4) are short and conical, usually at the basis of each actine (Fig. 2 D, E). Monolophose calthrops are abundant (13 – 28 lm actines long). The lophose actine is distally ramified by 2 – 4 short conical rays with sharp endings (Fig. 2 F). Trilophose calthrops are common; lophose actines ramify close the base in 2 – 3 rays, distally in 2 – 4 smaller rays with terminal spines (ramification pattern 1 proximal, 2 distal, terminal spines; for the terminology of ramification patterns, see Muricy & Dıaz, 2002). The non-lophose actines are 10 – 30 lm long (Fig. 2 G). Tetralophose calthrops are heterolophose and the less abundant spicule type. The three apical actines ramify distally in 3 – 4 rays, then distally in 2 – 5 short rays with terminal spines. The basal actine ramifies distally in two rays, and then distally again in 2 – 3 short rays with terminal spines (ramification pattern 1 distal, 2 distal, terminal spines). The actine length is 5 lm on average (Fig. 2 H; Table 1). Tissue general organization: The ectosome is 15 – 30 lm thick, separated from the choanosome by subectosomal cavities (15 – 40 lm) with a well-developed system of inhalant / exhalant canals (25 – 140 lm wide). The aquiferous system is leuconoid (Fig. 3 A, B). Choanocyte chambers, 47 – 78 lm in diameter, are spherical and diplodal (Fig. 3 B, C). Cytology: Choanocytes are 3 – 7 lm wide and 4 – 6 lm high, their collar, 40 – 50 lm in diameter, is composed of about 50 microvilli. The choanocyte nucleus is spherical and located on the apical area of the cell (4 – 5 lm in diameter). Their cytoplasm is dense, often with one to five phagosomes of about 1.0 – 1.5 lm (Fig. 3 B, C). The exo- and endopinacocytes are flat or ovoid (15 – 27 lm long and 7 – 11 lm large), with several vacuoles observed in their cytoplasm (Fig. 3 B – D). Their nucleus is also flattened (4 – 5 lm in diameter). Choanocytes and pinacocytes are flagellated, underlined by a basement membrane (Fig. 3 C, D). Few archaeocytes were observed. They have an irregular form with a spherical nucleus 2 lm in diameter (Fig. 3 D, E). Symbiotic prokaryotes: Plakina arletensis sp. nov. can be considered as a high microbial abundance (HMA) sponge. Three main morphotypes were easily detected, all three being extracellular and randomly dispersed in most of the mesohyl (Fig. 3 E, F). The first morphotype corresponds to an abundant ovoid cell (1.5 – 1.8 lm long; 0.4 – 1 lm high), with a more or less dense periplasm. We consider most of the small spherical forms as a perpendicular view of morphotype 1 (Fig. 3 E, F). The second morphotype, less abundant, has a rod-like to irregular shape (2 – 3 9 0.3 – 0.8 lm) with several translucent vacuoles. The third morphotype, less abundant, has the same ovoid form and dense periplasm as the first morphotype, but is bigger (3 – 5 9 1 – 1.7 lm) (Fig. 3 E, F). Ecology: Plakina arletensis sp. nov. is found only in shallow water caves, where it forms a white crust with a patchy distribution on vertical walls. No indications of epibiosis or predation were observed. Taxonomic remarks	en	Ruiz, César, Muricy, Guilherme, Lage, Anaíra, Domingos, Celso, Chenesseau, Sandrine, Pérez, Thierry (2017): Descriptions of new sponge species and genus, including aspiculate Plakinidae, overturn the Homoscleromorpha classification. Zoological Journal of the Linnean Society 179 (4): 707-724, DOI: 10.1111/zoj.12480, URL: https://www.mendeley.com/catalogue/813c5d34-fdc1-3687-80eb-affa77eab333/
03B50F5C7106FF8082AFFE81FD12D201.taxon	description	PLAKINA NATHALIAE (ERESKOVSKY, WILLENZ &	en	Ruiz, César, Muricy, Guilherme, Lage, Anaíra, Domingos, Celso, Chenesseau, Sandrine, Pérez, Thierry (2017): Descriptions of new sponge species and genus, including aspiculate Plakinidae, overturn the Homoscleromorpha classification. Zoological Journal of the Linnean Society 179 (4): 707-724, DOI: 10.1111/zoj.12480, URL: https://www.mendeley.com/catalogue/813c5d34-fdc1-3687-80eb-affa77eab333/
03B50F5C7105FF8A8294FA0BFEF8D05C.taxon	description	FIGS 5, 6 Diagnosis: Plakinidae without spicules, well-developed mesohyl with a thick collagen layer and without subectosomal cavities. Generally brown, but also cream to white. Smooth and ‘ grooved’ surface. Jelly-like consistency. Leuconoid aquiferous system and aphodal choanocyte chambers. High abundance of prokaryotic symbionts. Holotype: MNHN DJV 180, Rocher du Diamant at 12 m depth, La Martinique (14 ° 26.5 0 N, 61 ° 03.083 0 W). Collector: T. Perez, 13 June 2011. +, Presence;, absence; HMA, high microbial abundance; LMA, low microbial abundance; NA, data not available (modified from Muricy et al., 1999). Paratype 1: MNHN DJV 181, Tintamare, Les Arches at 10 m depth, Saint Martin (18 ° 07.588 0 N, 62 ° 58.248 0 W). Collector: C. Ruiz, 26 May 2015. Paratype 2: MNHN DJV 182, Grotte Cathedrale at 16 m depth, Anse Bertrand, La Guadeloupe (16 ° 27.740 0 N, 061 ° 31.837 0 W). Collector: C. Ruiz, 29 May 2015. Other specimens examined 150514 - MT 6 - TP 1, Grotte Chauve-Souris at 7 m depth, Anse Noire, La Martinique (14 ° 32.024 0 N, 61 ° 05.278 0 W). Collector: T. Perez, 14 May 2015. 150517 - SV 1 - CR 14, Bat Cave at 8 m depth, Bucament Bay, Saint Vincent (13 ° 11.275 0 N, 61 ° 16.174 0 W). Collector C. Ruiz, 17 May 2015. 150523 - GU 1 - CR 7, Cave north-west of Les Saintes at 8 m depth, Les Saintes, La Guadeloupe (15 ° 52.984 0 N, 61 ° 34.25 0 W). Collector: C. Ruiz & T. Perez, 23 May 2015. 131206 - MT 3 - AE 2, Grotte Couleur, Pointe Burgos at 12 m depth, Anse d’Arlet, La Martinique (14 ° 29.752 0 N, 61 ° 05.407 0 W). Collector: A. Ereskovsky, 6 December 2013. 150527 - SN 5 - TP 5, Basses Espagnoles at 10 m depth, Saint Martin (18 ° 07.821 0 N, 63 ° 00.270 0 W). Collector: T. Perez, 27 May 2015. 150528 - AG 3 - CR 1, Little Scrub at 15 m depth, Anguilla (18 ° 17.903 0 N, 62 ° 57.294 0 W). Collector C. Ruiz, 28 May 2015. 150530 - GU 6 - TP 7, Grotte Amedien at 12 m depth, La Guadeloupe (16 ° 30.033 0 N, 061 ° 28.774 0 W). Collector: T. Perez, 30 May 2015. Etymology: The genus name reflects the absence of skeleton: from Latin a (= without) spiculum (= spearhead, arrowhead) phora (= bearing). The species name, madinina, refers to the amerindian name, ‘ Flower Island’, of La Martinique Island, which is the first island of the Caribbean Sea where this aspiculate Plakinidae was found. Description: Aspiculophora madinina has mainly a cushion shape, measuring 2 – 15 cm in diameter by 2 – 5 cm high, but sometimes it can be found hanging down, with prominent oscules surrounded by a thin membranous collar (Fig. 5 A). The colour in vivo is mainly brown, occasionally yellow to cream (Fig. 5 A). Brown specimens produce a dark exudate in contact with alcohol. The consistency is soft, almost gelatinous. The surface is smooth and grooved. Oscules are circular, 1 – 2 mm in diameter with an elevated rim. Soft tissue organization: The ectosome is 6 – 15 lm. The aquiferous system is leuconoid with inhalant / exhalant canals about 30 – 100 lm wide. A dense collagen fibril layer, about 8 lm in thickness, surrounds the exhalant canals. Some specimens also exhibit a thicker collagenous layer, about 1 mm, between the ectosome and the basal part of the sponge, where very few cells or prokaryotes are found (Fig. 5 B). Choanocyte chambers (18 – 36 lm in diameter) are spherical and aphodal. Cytology: The choanocytes are cylindrical to spherical, 4 – 7 lm wide and 6 – 7 lm high. Their nucleus, 2 lm in diameter, is spherical and in apical position (Fig. 6 A, B). Their cytoplasm is not dense, often with one to three phagosomes and microgranular inclusions of about 1.5 lm in diameter. These inclusions are also observed in endopinacocytes (Fig. 6 C). The exo- and endopinacocytes are flattened, 15 – 20 lm long and 7 – 11 lm wide. Only one type of vacuolar cell is present and restricted to the ectosomal region, sometimes in aggregates, always close to exhalant canals (Fig. 6 C). This spherical cell, 9 – 13 lm in diameter, has a nucleus of 3 – 4 lm and harbours between two and ten vacuoles. Few archaeocytes were observed, with an irregular form and a nucleus of about 2 lm in diameter. Symbiotic prokaryotes: This sponge can be considered as an HMA sponge. Five morphotypes were distinguished based on their morphological traits. All prokaryotes are randomly dispersed and occupy most of the mesohyl (Fig. 6 B – D). The first morphotype corresponds to an abundant ovoid cell (2 – 3 lm long; 1 – 1.5 lm wide), with a clear membrane about 0.1 lm wide. The second morphotype, also abundant in the sponge mesohyl, has a rod form (1.5 lm long; 0.5 lm wide). The third morphotype has an irregular ovoid form (1.5 – 2.0 lm long; 1 lm wide), with a dense cytoplasm and one to three external vacuoles in contact with the cells’ outer membrane. The fourth morphotype is a spherical cell, with a dense periplasm, of about 1 lm in diameter. The fifth morphotype is a small spherical cell of about 0.1 lm in diameter. This morphotype was sometimes observed in groups of three to five cells in a row (Fig. 6 D). Ecology: Aspiculophora madinina sp. nov. is a sciaphilous species, living on vertical walls of semi-dark and dark caves or under overhangs. It was recorded between 5 and 50 m depth. Taxonomic remarks Aspiculophora madinina sp. nov. is a homoscleromorph sponge without skeleton and with a high content of collagen, providing a jelly-like consistency. As yet, massive forms such as this have not been reported in the aspiculate Oscarellidae. On the other hand, this species could be a Plakortis without spicules because most of its representatives occur as massive or globular forms. There is one report of such a Plakortis without spicules in the Caribbean Sponge Guide (Zea, Henkel & Pawlik, 2014), but the taxonomic affiliation to Plakortis was based only on the external morphology and the pungent smell of some specimens. It is difficult to compare the cytological characteristics of a putative new species with other Plakinidae, because of the lack of data at this level of biological organization. Indeed, traditional taxonomy has placed most attention on the skeletal description of this family, even if skeletal differences are poorly detectable in many species. Some other characters such as a leuconoid aquiferous system, eurypylous choanocyte chambers, well-developed mesohyl and subectosomal cavities are not sufficiently diagnostic, as they can be present or absent among Plakina, Plakortis and Plakinastrella species. We thus believe that more thorough cytological investigations of Plakortis, Plakinastrella and Corticium might help in resolving a good number of taxonomic questions in this family. The new genus is proposed because of the singularity of its CO 1 sequences positioning the new species among the Plakinidae, near the base of this clade, but outside all previously known genera. Moreover, we believe that its internal organization, with a thick collagen layer surrounding the inhalant canals and within the mesohyl, is unique among Homoscleromorpha. DNA ANALYSIS OF STUDIED SPECIES GenBank accession numbers for all sequences in this study are presented in Table 3. The two families of Homoscleromorpha are well supported in the phylogenetic reconstructions (Figs 7 – 9). With the Folmer partition (Fig. 7), Corticium appears as a monophyletic taxon of Plakinidae. Plakortis seems to be paraphyletic, a first clade being composed of Plakortis angulospiculatus (Carter, 1879) and the type species P. simplex Schulze, 1880, while a second clade clusters P. albicans Cruz-Barraza & Carballo, 2005 with two Plakinastrella species. In this representation lacking a sequence of the type species, Plakina appears as a poorly supported monophyletic group (Fig. 7), with two clades (bootstrap values of 312 / 47), one with P. crypta Muricy, Boury-Esnault, Bezac & Vacelet, 1998, P. monolopha and P. muricyae Cruz-Barraza, Vega & Carballo, 2014 closely related, and the other comprising three specimens of P. nathaliae, P. trilopha, P. jani and P. kanaky. The second phylogenetic tree using the I 3 - M 11 CO 1 fragment (Fig. 8) supports three monophyletic genera among Plakinidae, Aspiculophora, Plakinastrella and Plakortis. In this analysis Plakina is paraphyletic, P. monolopha (type species) forming a highly supported clade with C. candelabrum Schmidt, 1862, a second group containing P. arletensis sp. nov. and P. jani, and a third with P. crypta and P. trilopha. The third phylogenetic tree was made by concatenating all available sequences (Fig. 9). Because of missing data for one of the two fragments, the alignment contains several gaps and the concatenated tree appears less robust than the two-first representations of the inter-specific relationships among Homoscleromorpha. However, the topology of this third tree is quite congruent with the previous ones. In this representation, Aspiculophora forms a separate clade within the Plakinidae. Also, Plakina seems paraphyletic with two different clades, the first one grouping P. arletensis sp. nov. with P. kanaky, P. jani, P. trilopha and P. nathaliae, whereas the second one, containing the type species P. monolopha, P. muryciae and P. crypta, is related to Corticium, which seems monophyletic. Plakortis also appears paraphyletic, a first clade grouping the type species P. simplex, P. halichondroides and P. angulospiculatus, and a second clade containing P. albicans, Plakinastrella onkodes and an undetermined Plakinastrella. Finally, a separate clade containing five specimens of Plakinastrella also makes this genus paraphyletic in this phylogenetic representation.	en	Ruiz, César, Muricy, Guilherme, Lage, Anaíra, Domingos, Celso, Chenesseau, Sandrine, Pérez, Thierry (2017): Descriptions of new sponge species and genus, including aspiculate Plakinidae, overturn the Homoscleromorpha classification. Zoological Journal of the Linnean Society 179 (4): 707-724, DOI: 10.1111/zoj.12480, URL: https://www.mendeley.com/catalogue/813c5d34-fdc1-3687-80eb-affa77eab333/
