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Aspiculortis garifuna sp. nov. Etymology: The genus name Aspiculortis refers to the absence of skeleton and to the genetic proximity of Plakortis. The species name garifuna honors the ethnical and cultural diversity of the Garifuna people, descendants of the first Caribs, Arawak Amerindians and the African people who inhabited the Lesser Antilles. Diagnosis: Thick crust (about 5 mm thick) with irregular form. No spicules, nor fibrous skeleton. The surface is covered by ostia with a series of translucent canals ending in prominent oscula of about 2 mm diameter. One type of vacuolar cell abounds in the ectosome and is also present in the mesohyl. Examined material: Holotype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV202 Martinique, “Grotte Fer à Cheval” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark section of the cave, date 25/03/2012, sample code: 120325 MT4 b-TP6, collector: T. Pérez. Genbank accession number: MK 949483 Paratype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV203 Guadeloupe, “Grotte aux Barracudas” (16°27.301’ N, 61°32.253’ W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 30/05/2015, sample code: 150530GU5-CR6, collector: C. Ruiz. Genbank accession number: MK 949485 Other material examined: Bequia, The Grenadines “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 19/05/2015, sample code: 150519LG1-CR14, collector: C. Ruiz. Genbank accession number: MK 949484 Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark section of the cave, date 11/06/2011, sample code: 110613 MT4 -TP3, collector: T. Pérez. Genbank accession number: MK 949482 Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 11/06/2011, sample code: 110613 MT4 -TP8, collector: T. Pérez. Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 15/05/2015, sample code: 150515 MT4 -CR1, collector: C. Ruiz. Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 16/05/2015, sample code: 160515 MT4 -CR9, collector: C. Ruiz. Comparative material examined: Aspiculophora madinina Ruiz, Muricy, Lage, Domingos, Chenesseau & Pérez 2017, Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark section of the cave, date 16/05/2015, sample code: 160515 MT4 -CR10, collector: C. Ruiz. Plakina nathaliae (Ereskovsky et al. 2014), Saint Martin, “Basses Espagnoles” (18°07.821’N, 63°00.270’W), 15 m depth, beneath an overhang, date 27/05/2015, sample code: 270515SN5-TP01, collector: T. Pérez. Oscarella zoranja Pérez & Ruiz, 2018, Martinique, Rocher du Diamant, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark section of the cave, date 16/05/2015, sample code: 150516 MT4 -CR05, collector: C. Ruiz. Oscarella filipoi Pérez & Ruiz, 2018, Saint Lucie, “Anse Chastenet reef” (13°51.848’ N, 61°04.729’ W), 15 m depth, beneath an overhang, date 22/05/2015, sample code: 150522SL1-CR05, collector: C. Ruiz. Description: External morphology: Aspiculortis garifuna sp. nov. has an encrusting irregular form and a brown color in vivo and after fixation in ethanol. The largest specimen recorded measured 10 cm in diameter. This sponge is quite fragile. The smooth surface is pseudo-criblate, that it is to say covered by numerous ostia diverging soon under the epidermal surface in various inhalant canals. The sponge surface also presents a series of larger translucent exhalant canals ending in prominent oscula of about 2 mm in diameter. This dense network can appear contracted in some specimens (Figure 2 A-B). Internal Anatomy: No spicule nor fibrous skeleton. The ectosome is 40-60 µm thick (Figure 2 C-D). The aquiferous system is leuconoid. Inhalant canals are 10-15 µm wide, running perpendicular to the surface. Choanocyte chambers (15-33 µm in diameter) are diplodal, spherical to ovoid (Figure 2E). The proportion between mesohyl and choanocyte chambers is greater than 1. Cytology: Choanocytes are cylindrical, 5 µm wide and 7 µm high. Their nucleus is spherical (2 µm diameter) in apical position, their cytoplasm usually contains osmiophilic inclusions and vacuoles of about 2-3 µm in diameter that often harbor phagocytized bacteria (Figure 3A). The exo- and endopinacocytes are flagellated and flattened (12- 19 µm long; 3-4 µm wide) with several vacuoles and osmiophilic inclusions. The external surfaces of the exo- and endopinacocytes are covered with a double-layer of 0.5 µm thick glycocalyx (Figure 3B). One type of vacuolar cell is found in abundance in the ectosome, and to a lesser extent in the mesohyl. This irregular shaped-cell, 10 x 20 µm in dimension, harbors 2-4 translucent vacuoles which are 3 to 8 µm in diameter (Figure 3C) and which often contain dense or loose heterogeneous material (Figure 3D). No archaeocyte was observed. There is a high abundance of prokaryotic symbionts in the mesohyl, with at least five dominant morphotypes randomly distributed throughout the mesohyl (Figure 3E). The first morphotype is an ovoid cell 2 - 4 µm long, 2.5 – 3 µm wide with a clear cytoplasm. The second morphotype has the same ovoid shape and size but has a denser cytoplasm. The third morphotype is an irregular cell shape which is 2 – 4 µm long and about 1 – 2.5 µm wide. The fourth morphotype is bean-shaped, 2.5 - 3 µm long and about 0.5 wide. The fifth morphotype is circular with a diameter of about 0.5 µm. Ecology: This new sponge has been collected in the Caribbean islands (Lesser Antilles) of Guadeloupe, Martinique and Bequia. In all sites, this sponge inhabits vertical walls and ceilings of dark marine caves, located at depths between 10 and 25 m, in syntopy with several other Homoscleromorpha species. No sign of predation was observed. Serpulidae worms live in close association and are often observed living among the new sponge species (Figure 2B). Taxonomic remarks: Aspiculortis garifuna gen. nov. sp. nov. has an encrusting form, as do most of the cave-dwelling Homoscleromorpha. It clearly belongs to a well-supported clade gathering Plakortis species, but its pseudo-criblate surface is unique, and thus has never been observed in Plakortis. The lack of cytological description of Caribbean Plakortis did not allow any comparison with the cells content of the new sponge. The absence of skeleton and this original pseudo-criblate surface justify the creation of a new genus. The new sponge also clearly differs from the other skeleton-less Plakinidae by its external morphology and internal organization, particularly the proportion between mesohyl and choanocyte chambers and the low abundance of symbiotic microorganisms in the thinly encrusting Plakina nathaliae, and the thick collagen layer found in the ectosome of the massive Aspiculophora madinina, as well as its aphodal choanocyte chambers.

Oscarella minka sp. nov. Etymology: From the Quechuan mink’a (minga), this species name honors the pre-Columbian tradition of communal work for social purposes. Diagnosis: Thin crust less than 3 mm thick. Smooth surface. Ectosome 25 to 50 µm thick, well-developed mesohyl. Two types of vacuolar cells, leuconoid aquiferous system with diplodal choanocyte chambers. High abundance of prokaryotic symbionts in the mesohyl. Examined material: Holotype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV206 La Martinique, “Grotte Fer à Cheval” (14°27.882’ N, 61°01.162 W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 25/03/2012, sample code: 120325 MT4 b-TP8, collector: T. Pérez. Genbank accession number: MK 949481 Paratype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV207 Bequia, The Grenadines, “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 19/05/2015, sample code: 150519LG1-CR9, collector: C. Ruiz. Genbank accession number: MK 949480 Other material examined: Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 13/06/2011, sample code: 110613 MT4 -TP4, collector: T. Pérez. GenBank accession number: MK 949478 Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 15/05/2015, sample code: 150515 MT4 - CR2, collector: C. Ruiz. Genbank accession number: MK 949479 Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 13/06/2011, sample code: 110613 MT4 -TP6, collector: T. Pérez. Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 16/05/2015, sample code: 150516 MT4 -CR11, collector: C. Ruiz. Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 16/05/2015, sample code: 150516 MT4 -CR16, collector: C. Ruiz. Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 16/05/2015, sample code: 150516 MT4 -CR17, collector: C. Ruiz. Bequia, The Grenadines, “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 28/04/2015, sample code: 150428LG1-AE55, collector: A. Ereskovsky. Bequia, The Grenadines, “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 19/05/2015, sample code: 150519LG1-TP5, collector: T. Pérez. Guadeloupe, “Grotte Cathédrale” (16°27.740’ N, 61°31.837’ W), 18 m depth, on a vertical wall in the dark part of the cave, date 29/05/2015, sample code: 150529GU3-TP2, collector: T. Pérez. Guadeloupe, “Grotte aux Barracudas” (16°27.301’ N, 61°32.253’ W), 19 m depth, beneath an overhang, date 30/05/2015, sample code: 150530GU5-CR4, collector: C. Ruiz. Guadeloupe, “Grotte aux Barracudas” (16°27.301’ N, 61°32.253’ W), 19 m depth, beneath an overhang, date 30/05/2015, sample code: 150530GU5-CR10, collector: C. Ruiz. Comparative material examined: Oscarella filipoi, Martinique, Anse d’Arlet, Grande Anse, Pointe Burgos - Grotte Couleur (14°2.787’N, 61°5.351’W), 10 m depth, on a vertical wall in semi-dark conditions, date 11/06/2011, sample code: 110611 MT3 -TP05, collector: T. Pérez. Oscarella zoranja, “ Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, date 13/06/2011, sample code: 110613 MT4 -TP1, collector: T. Pérez

Genus Oscarella Vosmaer, 1884 Diagnosis: Homosclerophorida without skeleton, with a variable development of the ectosome with a collagen layer between ecto- and choanosome. The aquiferous system has a sylleibid-like or leuconoid organization, with spherical to ovoid eurypilous or diplodal choanocyte chambers arranged around large exhalants canals (Muricy & Díaz 2002). Mitochondrial genomes that have been sequenced so far encode a gene absent in other animal mitochondrial genomes: tatC (Ereskovsky et al. 2017).

Aspiculophora papillata sp. nov. Etymology: the species name refers to the papillate surface of this Plakinidae, papilla = nipple in Latin. Diagnosis: encrusting Aspiculophora with papillate surface. Collagen layer ( Examined material: Holotype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV204 Bequia, The Grenadines, “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 19/05/2015, sample code: 150519LG1-CR13, collector: C. Ruiz. Genbank accession number: MK 949486 Paratype: Muséum National d’Histoire Naturelle de Paris, France MNHN DJV205 Guadeloupe, “Grotte Cathédrale” (16°27.740’ N, 61°31.837’ W), 18 m depth, on a vertical wall in the dark part of the cave, date 29/05/2015, sample code: 150529GU3-CR1, collector: C. Ruiz. Other material examined: Martinique, “Grotte Zeb” (14°27.832’N, 61°01.065’W), 19 m depth, on a vertical wall in the semi-dark part of the cave, dates 15 and 16/05/2015, sample codes: 150515 MT4 -CR3, 150515 MT4 - CR6, 160515 MT4 -CR14, collector: C. Ruiz. Bequia, The Grenadines “Full Moon Cave” (12°59.525’ N, 61°16.731’ W), 10-12 m depth, beneath an overhang, date 19/05/2015, sample codes: 150519LG1-CR10, 150519LG1-CR13, 150519LG1-TP11, collectors: C. Ruiz & T. Pérez Comparative material examined: Aspiculophora madinina MNHN DJV 180, Rocher du Diamant “Grotte Zeb”, Martinique (14°26.5´N, 61°03.083´W), 12 m depth, date 13/06/2011, sample code: 110613 MT4 -TP11 collector: T. Pérez, Aspiculophora madinina, “ Grotte Amédien”, Guadeloupe (16°30.033’ N, 061°28.774’ W), 12 m depth, date 30/05/2015, sample code: 150530-GU6-TP07, collector: T. Pérez. Plakina nathaliae MNHN DJV 179, Rocher du Diamant “Grotte Zeb”, La Martinique (14°26.5´N, 61°03.083´W), 17 m depth, date 13/06/2011, sample code: 110613 MT4 -TP2, collector: T. Pérez. Description: External morphology: Aspiculophora papillata sp. nov. is a thinly encrusting irregular sponge which is quite hard to collect because of its fragile consistency. Collected specimens measured up to 10 cm wide and 1 mm thick. The sponge has a papillate surface and its color varies from yellow to light brown in vivo, whereas it is light brown in ethanol. Papillae are cylindrical, up to 3 mm high and about 1 mm in diameter. The top of the papillae is often larger with an umbrella-like form of 2-3 mm in diameter. Oscula of about 1 mm in diameter are sometimes visible in the top of the papillae (Figure 4A, B). Internal Anatomy: No spicule or fibrous skeleton. A collagen lager Cytology: Choanocytes are cylindrical, 5 µm wide and 8 µm high. Their nuclei are spherical, about 2 µm in diameter and are located in apical position. Their cytoplasm usually contains osmiophilic inclusions of about 0.1 µm in diameter in basal position (Figure 5A, B). Apopylar cells have a trapezoid shape (Figure 5A). Exo- and endopinacocytes are flat and flagellated (5-15 µm long; 1-2 µm wide) with a nucleus of about 2.5 µm in diameter (Figure 5C). One type of abundant vacuolar cell was found randomly distributed in the mesohyl. This cell has an ovoid to spherical shape, is 12-20 µm in diameter, and generally harbors one to five vacuoles occupying most of the volume of the cytoplasm. Some heterogeneous to filamentous materials can be seen inside the vacuoles, and microorganisms can be found in their cytoplasm (see arrow in Figure 5C). No archaeocyte was observed. The new species can be considered as an HMA sponge. At least four prokaryotic morphotypes seem to be dominant in the mesohyl (Figure 5D). The first morphotype is peanut-shaped, 2.5 µm long 0.5 µm wide, with a dense cytoplasm and a double membrane. The second morphotype is of similar size, is cylindrical in shape and has a translucent external membrane. The third morphotype has an irregular shape and is 2-4 µm long and 1 µm wide. The fourth morphotype is bean-shaped with a dense membrane and translucent cytoplasm. Ecology: The new sponge species was found in the Caribbean islands of Guadeloupe, Martinique and Bequia (Lesser Antilles). It lives in sympatry (and sometimes in syntopy) with A. madinina, on walls and ceilings of dark marine caves, where it is always found between 10 and 20 m depth. No sign of predation was observed, but the new species usually lives in close association with other sponges and invertebrates such as Serpulidae and Sabellidae worms (Figure 4A). Taxonomic remarks: Aspiculophora papillata sp. nov. is a skeleton-less Plakinidae like A. madinina. The new species mainly differs by its external morphology, the papillate surface being a trait observed for the first time among Homoscleromorpha. In contrast with A madinina, no dark exudate was recorded in contact with alcohol for A. papillata. The collagen layer between the exopinacoderm and the ectosome region in A. papillata sp. nov. is very thin in comparison with A. madinina. Another difference between these species is the presence of sub-ectosomal cavities in A. papillata sp. nov. that do not exist in A. madinina. Both species share the leuconoid aquiferous system with aphodal choanocyte chambers, only one type of vacuolar cell which is mostly concentrated in the ectosome and similar microbial compositions.

Aspiculortis gen. nov. Diagnosis: Plakinidae without spicules. Smooth pseudo-criblate surface covered by numerous ostia diverging soon under the epidermal surface in various inhalant canals. A network of superficial translucent much larger exhalant canals leading to prominent oscula. Well-developed mesohyl with several layers of vacuolar cells in the ectosome. No sub-ectosomal cavities. Leuconoid aquiferous system with diplodal choanocyte chambers. High abundance of prokaryotic symbionts in the mesohyl.

Genus Aspiculophora Ruiz et al. 2017 Diagnosis: Plakinidae without spicules, well-developed mesohyl with a collagen layer between ecto- and choanosome, with or without sub-ectosomal cavities. Leuconoid aquiferous system and aphodal choanocyte chambers. High abundance of prokaryotic symbionts.

International audience; Potential for, and limits to, adaptation to environmental changes are critical for resilience and risk mitigation. The Mediterranean basin is a mosaic of biodiversity-rich ecosystems long affected by human influence, whose resilience is now questioned by climate change. After reviewing the different components of biological adaptation, we present the main characteristics of marine and terrestrial biodiversity in the Mediterranean basin and of the pressures they face. Taking climatic trends into consideration, we discuss the adaptive potential of a range of ecosystems dominated by species without active dispersal. We argue that the high heterogeneity of Mediterranean landscapes and seascapes constitutes a laboratory for the study of adaptation when environmental conditions change rapidly and may provide opportunities for adaptation and adaptability of species and ecosystems. Adaptive management in the Mediterranean can and should harness the nature-based solutions offered by both ecological and evolutionary processes for increasing the resilience of ecosystems to climate change