identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
6864879DFFCDA777FEDAFCDFFD66FC73.text	6864879DFFCDA777FEDAFCDFFD66FC73.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Spongia Linnaeus 1759	<div><p>Genus Spongia Linnaeus, 1759</p><p>Type species: Spongia officinalis Linnaeus, 1759 .</p><p>Definition: Unarmoured spongiids, not heavily lacunose, with simple primary fibres (from Cook and Bergquist 2002).</p></div>	https://treatment.plazi.org/id/6864879DFFCDA777FEDAFCDFFD66FC73	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grenier, Marie;Simmler, Charlotte;Chevaldonné, Pierre;Callizot, Noëlle;Pérez, Thierry	Grenier, Marie, Simmler, Charlotte, Chevaldonné, Pierre, Callizot, Noëlle, Pérez, Thierry (2024): Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach. Zoological Journal of the Linnean Society 202: 1-23, DOI: 10.1093/zoolinnean/zlad166
6864879DFFCDA777FEDEFDE5FE05FCA0.text	6864879DFFCDA777FEDEFDE5FE05FCA0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Spongiidae Gray 1867	<div><p>Family Spongiidae Gray, 1867</p><p>Definition: Dictyoceratida with homogeneous skeletal fibres, without distinct laminations, typically dominated by subprimary skeletal fibres, and with diplodal choanocyte chambers (from Cook and Bergquist 2002).</p></div>	https://treatment.plazi.org/id/6864879DFFCDA777FEDEFDE5FE05FCA0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grenier, Marie;Simmler, Charlotte;Chevaldonné, Pierre;Callizot, Noëlle;Pérez, Thierry	Grenier, Marie, Simmler, Charlotte, Chevaldonné, Pierre, Callizot, Noëlle, Pérez, Thierry (2024): Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach. Zoological Journal of the Linnean Society 202: 1-23, DOI: 10.1093/zoolinnean/zlad166
6864879DFFCDA77BFF4FFB8FFC59FDB6.text	6864879DFFCDA77BFF4FFB8FFC59FDB6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Spongia maitasuna Grenier & Perez	<div><p>Spongia maitasuna Grenier &amp; Pérez, 2 0 23 sp. nov.</p><p>(Fig. 2A–D)</p><p>Material examined</p><p>Holotype: MNHN-IP-2018-419, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59″N, 1°39 ʹ 52″W). Collector: T. Pérez, 26 July 2021. Voucher code: 20210726-CIB-BTP6. GenBank accession numbers OR297953 and OR291312 (CO1 and 28S, respectively) .</p><p>Paratype 1: MNHN-IP-2018-420, Western part of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-3.7561111&amp;materialsCitation.latitude=43.47278" title="Search Plazi for locations around (long -3.7561111/lat 43.47278)">Mouro island</a>, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021. Voucher code: 20210729-ESP-MTP1. GenBank accession number OR291316 (28S) .</p><p>Paratype 2: MNHN-IP-2018-421, Eastern part of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-3.755&amp;materialsCitation.latitude=43.4725" title="Search Plazi for locations around (long -3.755/lat 43.4725)">Mouro island</a>, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021. Voucher code: 20210730-ESP-M2TP1. GenBank accession numbers OR297955 and OR291323 (CO1 and 28S, respectively) .</p><p>Other specimens examined</p><p>20210630-CIB-BPC2, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59″N, 1°39 ʹ 52″W). Collector: P. Chevaldonné, 30 June 2021.</p><p>20210726-CIB-BTP11, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59″N, 1°39 ʹ 52″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP12, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59″N, 1°39 ʹ 52″W). Collector: T. Pérez, 26 July 2021.</p><p>20210729-ESP-MTP2, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210729-ESP-MTP3, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210729-ESP-MTP4, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210729-ESP-MTP6, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210729-ESP-MTP7, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210729-ESP-MTP8, Western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22″N, 3°45 ʹ 22″W). Collector: T. Pérez, 29 July 2021.</p><p>20210730-ESP-M2TP7, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP8, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP9, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP10, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP11, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP13, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP14, Eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21″N, 3°45 ʹ 18″W). Collector: T. Pérez, 30 July 2021.</p><p>Comparative material examined</p><p>Spongia officinalis Linnaeus, 1759</p><p>(Fig.2E)</p><p>20200519-MRS-ITP3, Impérial de Terre, semi-dark cave, between 10 and 20 m depth, Marseille, France (43°10 ʹ 22″N, 5°23 ʹ 35″E). Collector: T. Pérez, 19 May 2020.</p><p>20200519-MRS-ITP4, Impérial de Terre, semi-dark cave, between 10 and 20 m depth, Marseille, France (43°10 ʹ 22″N, 5°23 ʹ 35″E). Collector: T. Pérez, 19 May 2020.</p><p>20200519-MRS-ITP5, Impérial de Terre, semi-dark cave, between 10 and 20 m depth, Marseille, France (43°10 ʹ 22″N, 5°23 ʹ 35″E). Collector: T. Pérez, 19 May 2020.</p><p>20200522-MRS-FTP2, Fauconnière, semi-dark tunnel, between 5 and 10 m depth, Saint-Cyr Les Lecques, France (43°09 ʹ 16″N, 5°40 ʹ 59″E). Collector: T. Pérez, 22 May 2020.</p><p>20200522-MRS-FTP3, Fauconnière, semi-dark tunnel, between 5 and 10 m depth, Saint-Cyr Les Lecques, France (43°09 ʹ 16″N, 5°40 ʹ 59″E). Collector: T. Pérez, 22 May 2020.</p><p>20200602-MRS-MTP9, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200617-MRS-NTP1, Niolon, semi-dark cave, between 8 and 10 m depth, Marseille, France (43°20 ʹ 15″N, 5°15 ʹ 22″E). Collector: T. Pérez, 17 June 2020.</p><p>20220909-MRS2-RTP4, Riou island, overhang at 28 m depth, Marseille, France (43°10 ʹ 23″N, 5°23 ʹ 15″E). Collector: T. Pérez, 9 September 2022.</p><p>20220909-MRS2-RTP5, Riou island, overhang at 28 m depth, Marseille, France (43°10 ʹ 23″N, 5°23 ʹ 15″E). Collector: T. Pérez, 9 September 2022.</p><p>20220909-MRS2-RTP6, Riou island, overhang at 28 m depth, Marseille, France (43°10 ʹ 23″N, 5°23 ʹ 15″E). Collector: T. Pérez, 9 September 2022.</p><p>20220923-MRS2-ITP25, Impérial de Terre, semi-dark cave at 20 m depth, Marseille, France (43°10 ʹ 22″N, 5°23 ʹ 35″E). Collector: T. Pérez, 23 September 2022.</p><p>20220923-MRS2-ITP26, Impérial de Terre, semi-dark cave at 20 m depth, Marseille, France (43°10 ʹ 22″N, 5°23 ʹ 35″E). Collector: T. Pérez, 23 September 2022.</p><p>20220923-MRS2-RTP27, Riou island, overhang at 25 m depth, Marseille, France (43°10 ʹ 23″N, 5°23 ʹ 15″E). Collector: T. Pérez, 23 September 2022.</p><p>Spongia cf.officinalis</p><p>(Fig.2F)</p><p>20090617-CEU-ECITP1, ECIMAR campaign, between 3 and 30 m depth, Ceuta, Spain (35°54 ʹ 25″N, 5°18 ʹ 00″W). Collector: T. Pérez, 17 June 2009.</p><p>20070701-CEU-ECITP2, ECIMAR campaign, between 3 and 30 m depth, Ceuta, Spain (35°52 ʹ 38″N, 5°18 ʹ 20″W). Collector: T. Pérez, 1 July 2007.</p><p>20070705-CEU-ECITP3, ECIMAR campaign, between 3 and 30 m depth, Ceuta, Spain ((35°52 ʹ 49″N, 5°18 ʹ 45″W). Collector: T. Pérez, 5 July 2007.</p><p>Spongia mollissima Schmidt, 1862 20200809-IT-NTP1, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP2, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP3, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP4, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP5, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP6, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200809-IT-NTP8, Neuralia wreck, between 30 and 35 m depth, Porto Cesareo, Italy (40°10 ʹ 27″N, 17°54 ʹ 19″E). Collector: T. Pérez, 9 August 2020.</p><p>20200810-IT-CTP1, El Camino, photophilic rocky bottoms at 17 m depth, Porto Cesareo, Italy (40°16 ʹ 12″N, 17°51 ʹ 15″E). Collector: T. Pérez, 10 August 2020.</p><p>20200810-IT-StTP2, La Strea, photophilic rocky bottoms at 17 m depth, Porto Cesareo, Italy (40°14 ʹ 54″N, 17°53 ʹ 38″E). Collector: T. Pérez, 10 August 2020.</p><p>Spongia nitens (Schmidt, 1862)</p><p>Po.25493, Tel Aviv, Israel (Idan et al. 2018).</p><p>Po.25665 Tel Aviv, Israel (Idan et al. 2018).</p><p>Spongia zimocca Schmidt, 1862</p><p>Po.25742 Tel Aviv, Israel (Idan et al. 2018)</p><p>Hippospongia communis (Lamarck, 1814)</p><p>20200519-MRS-JTP1, Jarre, cave in the semi-dark part at 15 m depth, Marseille, France (43°11 ʹ 46″N, 5°21 ʹ 55″E). Collector: T. Pérez, 19 May 2020.</p><p>20200522-MRS-FTP4, Fauconnière, semi-dark tunnel, between 5 and 10 m depth, Saint-Cyr Les Lecques, France (43°09 ʹ 16″N, 5°40 ʹ 59″E). Collector: T. Pérez, 22 May 2020.</p><p>20200522-MRS-FTP6, Fauconnière, semi-dark tunnel, between 5 and 10 m depth, Saint-Cyr Les Lecques, France (43°09 ʹ 16″N, 5°40 ʹ 59″E). Collector: T. Pérez, 22 May 2020.</p><p>20201225-TUN-Ep8, harvested by a fisherman in a seagrass meadow, Djerba, Tunisia, 25 December 2020.</p><p>Etymology</p><p>The species name ‘maitasuna’ comes from ‘maitasun’, which means ‘love’ in Euskara, the Basque Country language. Collected first in a site of the French Pays Basque, the new sponge is thus dedicated to all beloved members of our families, with special thoughts to those recently departed, especially to Marguerite Grenier, century-old grandmother, who passed away while this manuscript was in preparation.</p><p>Diagnosis</p><p>Massive sponge with unarmoured and non-lacunose surface. Surface covered with evenly disposed conules, sometimes with traces of epibiosis or foreign bodies. Soft and compressible consistency. Skeleton made of primary, secondary, and tertiary fibres. Primary fibres are anastomosed and denser at the surface, piercing the surface, forming conules, and presenting an open space variable in size between each anastomosed primary fibre. Primary fibres harbouring inclusions such as foreign spicules and/or a pith. Secondary fibres containing a pith inclusion and forming a clearly and regular honeycomb network. Tertiary fibres present both at the surface and within the sponge body.</p><p>Description</p><p>The new sponge is massive and irregular, sometimes slightly branched, with small lobes or protuberances (Fig. 2A–D). The sponge can measure ≤ 25 cm in its largest diameter. Its surface is white, beige, light to dark grey, whereas the internal tissue is usually tawny yellow. No change in colour was detected after fixation in 95% ethanol. The sponge surface is sometimes covered by dirty foreign bodies or epibiotic organisms. The surface is very conulose. Oscules can be grouped or not and are often found on the top of each lobe or protuberance. They are bordered by a thin and rather translucid membrane (Fig. 2A–D). The sponge consistency is rather soft and tearable.</p><p>The ectosomal skeleton presents a very thin epidermal skin, which has a star-like appearance, is easily detachable, and is made of collagen fibrils, from 60 to 350 µm in thickness (Fig. 3A) Scattered foreign spicules, debris, or remains of associated invertebrates, such as cirripeds, can be observed.</p><p>The choanosomal skeleton presents primary fibres, which are irregular and sinuous, sometimes difficult to observe and to distinguish because they appear anastomosed with secondary fibres, and in some places, they can also divide and join again (Fig. 3B–D). They measure 30–70 µm in diameter and are cored with inclusions of foreign spicules and/or debris of various abundance. They also include a black or translucid pith. These primary fibres with a pith can be observed near the surface into conule formation and/or across the sponge body (Fig. 3A, C). However, when foreign debris and spicules are widely present, the pith in the primary fibres can be difficult to observe. Near the surface, close to the conule formations, the network of primary fibres is denser in comparison to the rest of the sponge body. The space between two primary fibres, thus between conules, seems to be empty, with very few tertiary fibres and foreign spicules and debris. Perforated plates are inconsistently present close to the primary fibres (Fig. 3D). The secondary fibres form a dense network, clearly showing a honeycomb shape of variable mesh size (Fig. 3B), from 70 µm × 120 µm to 300 µm × 500 µm. These fibres measure 20–50 µm in diameter and they can contain an irregular pith, black or translucid (Fig. 3E). Tertiary fibres, ~10 µm in diameter, are found throughout the sponge body (Fig. 3F, G). Sometimes they form a thinner network that also has a clear honeycomb shape with variable mesh size (50–130 µm in diameter). In some places, these fibres appear to be linked to the thicker secondary fibres (Fig. 3H), but in some places they look rather disconnected from the rest of the skeletal architecture.</p><p>Distribution</p><p>South European Atlantic Shelf (Spalding et al. 2007), Bay of Biscay, French Pays Basque, and Spanish Cantabria (this work).</p><p>Ecology</p><p>Spongia maitasuna is found on rocky substrate in shallow water cliffs, boulders, or semi-dark and dark cavities, habitats usually with high wave energy. This sponge presents a patchy distribution and lives in syntopy with S. lamella . In some cases, both Spongiidae present contact interactions.</p><p>Some specimens may have epibiotic organisms, mostly encrusting sponges and zoanthids. No sign of predation was observed. In the specimens collected between late June and late July, various stages of embryogenesis were observed, including early fertilized eggs, various morula stages, and pre-larvae. No spermatic cysts were observed.</p><p>Taxonomic remarks</p><p>The new species is assigned to Spongia because its body is not lacunose, as in Hippospongia or Hyatella, and it does not present an armoured ectosomal skeleton, as in Coscinoderma or Leiosella . Unlike Rhopaloeides, which presents long and simple fascicles of primary fibres, the primary fibres of the new species are simple and sometimes anastomosed. In Spongia, the new species should belong to the subgenus Spongia, because it presents a simple primary skeleton with regular and polygonal meshes. However, considering the present status of the genus, we prefer not to use this infrageneric, lower rank classification (see the Discussion section).</p><p>Spongia maitasuna shares with the type species of Spongia the very thin collagenic epidermal skin and the fact that primary fibres are never forming fascicles, in other words they are never fused. In S. officinalis, the primary fibres are thicker than in the new species, ranging from 50 to 100 µm. The network of secondary fibres is much denser in S. officinalis, and the meshes do not have the typical honeycomb shape of the new species. This structure of the secondary skeleton is found in S. cf. officinalis from Ceuta, Strait of Gibraltar. The main difference in this case lies in the primary fibres, which are slightly larger than in the new species (70–110 µm), and in the overall growth form (Fig.2). Spongia maitasuna has a morphology similar to S. nitens, and both species display a pith within their primary fibres. But S. nitens has a denser secondary skeleton, and the secondary fibres are devoid of pith. Moreover, tertiary fibres were present in the two studied specimens of S. nitens, but according to the literature they can be absent (Vacelet 1959, Manconi et al. 2013). They were recorded in all specimens of the new species. Finally, the new species shares with the studied specimen of S. zimocca the anastomosed primary fibres and the thickness of the secondary fibres, although they can be reported thicker (50–80 µm) in the literature (Vacelet 1959, Manconi et al. 2013). In the studied specimen of S. zimocca, the primary fibres are much thicker, 50–100 µm (100–200 µm in the literature) than in the new species. These fibres also present a pith, although this trait has not been reported previously (Vacelet 1959, Castritsi-Catharios et al. 2011, Manconi et al. 2013), and rare foreign spicules. Finally, the spongin network is overall irregular and denser in S. zimocca than in the new species.</p></div>	https://treatment.plazi.org/id/6864879DFFCDA77BFF4FFB8FFC59FDB6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grenier, Marie;Simmler, Charlotte;Chevaldonné, Pierre;Callizot, Noëlle;Pérez, Thierry	Grenier, Marie, Simmler, Charlotte, Chevaldonné, Pierre, Callizot, Noëlle, Pérez, Thierry (2024): Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach. Zoological Journal of the Linnean Society 202: 1-23, DOI: 10.1093/zoolinnean/zlad166
6864879DFFC1A77FFC2FFD13FB45F848.text	6864879DFFC1A77FFC2FFD13FB45F848.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Spongia lamella (Schulze 1879)	<div><p>Spongia lamella (Schulze, 1879)</p><p>(Fig. 4)</p><p>Material examined</p><p>20200602-MRS-MTP3, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200602-MRS-MTP4, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200602-MRS-MTP5, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200602-MRS-MTP6, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200602-MRS-MTP7, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20200602-MRS-MTP8, Pharillons de Maïre, cliff at 20 m depth, Marseille, France (43°12 ʹ 27″N, 5°20 ʹ 17″E). Collector: T. Pérez, 2 June 2020.</p><p>20160607-GIJ-JC1, Las Gemelas island, rock substrate at 12 m depth, Gijon, Spain (43°33 ʹ 22″N, 5°36 ʹ 31″W). Collector: J. Cristobo, 7 June 2016.</p><p>20200624-HEN-BMNC2, Belhara, on reef flat at 22 m depth, Hendaye, France (43°24 ʹ 00″N, 1°43 ʹ 03″W). Collector: M. N. de Casamajor, 24 June 2020.</p><p>20200814-HEN-AMNC3, Abbadia, on reef flat at 13 m depth, Hendaye, France (43°23 ʹ 56″N, 1°45 ʹ 28″W). Collector: M. N. de Casamajor, 14 August 2020.</p><p>20210630-CIB-BPC1, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: P. Chevaldonné, 30 June 2021.</p><p>20210630-CIB-BPC3, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: P. Chevaldonné, 30 June 2021.</p><p>20210726-CIB-BTP1, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP2, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP3, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP4, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP7, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210726-CIB-BTP10, Baloo tunnel at 14 m depth, Ciboure, France (43°24 ʹ 59.0″N, 1°39 ʹ 52.2″W). Collector: T. Pérez, 26 July 2021.</p><p>20210729-ESP-MTP5, western part of Mouro island, semi-dark to dark cave, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 22.0″N, 3°45 ʹ 21.7″W). Collector: T. Pérez, 29 July 2021.</p><p>20210730-ESP-M2TP2, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21.1″N, 3°45 ʹ 18.3″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP3, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21.1″N, 3°45 ʹ 18.3″W). Collector: T. Pérez, 30 July 2021.</p><p>20210730-ESP-M2TP12, eastern part of Mouro island, shallow water cliffs and chaos of rocks and crevices, between 3 and 12 m depth, Santander, Spain (43°28 ʹ 21.1″N, 3°45 ʹ 18.3″W). Collector: T. Pérez, 30 July 2021.</p><p>Comparative material examined</p><p>Spongia virgultosa (Schmidt, 1868)</p><p>20070718-MRS-ECITP5, Jarre, cave in the semi-dark part at 15 m depth, Marseille, France (43°11 ʹ 46″N, 5°21 ʹ 55″E). Collector: T. Pérez, 18 July 2007.</p><p>Coscinoderma sporadense Voultsiadou-Koukoura, van Soest &amp; Koukouras, 1991</p><p>Po.25932, Tel-Aviv, Israel (Idan et al. 2018).</p><p>Description</p><p>This sponge can be massive and irregular, with lobes or protuberances, fan- or vase-shaped when it becomes very big (Fig. 4A–D). The largest studied specimens measured 50 cm in their largest diameter, but this dimension can be&gt; 1 m in the Mediterranean Sea (Fig. 4D). No true vase- or fan-shaped specimen was observed among our Atlantic specimens. This sponge presents a light to dark grey external colour, and the internal tissue is usually light or tawny yellow. The colour does not change upon fixation in 95% ethanol. The sponge consistency is flexible and hardly tearable. The surface is finely covered with small, evenly distributed conules. Oscules can be grouped in large individuals, or more randomly distributed in smaller ones, always on the outer sides in large individuals or on the top of each lobe or protuberance in smaller ones. They measure 1–3 mm in diameter and are slightly raised, with a small whitish membrane delimiting the opening.</p><p>The ectosomal skeleton harbours an epidermal skin, 150– 600 µm thick, easily detachable, with a star-like appearance. It is made of a network of abundant foreign spicules and debris (Fig. 5A–C). Small canals can be observed under the surface. In the choanosome, debris and foreign spicules are still abundant at the periphery of canals, and thus throughout the sponge body (Fig. 5D). The primary fibres, 30–120 µm in diameter, are common, simple, irregular, and cored with numerous inclusions of foreign spicules and debris (Fig. 5A, B). Near the surface, they can be ramified. Secondary fibres, 20–40 µm in diameter, without inclusions or pith, form a dense ramified and irregular network (Fig. 5B, E). Tertiary fibres are present or absent. When they exist, their thickness is ≤10 µm in diameter.</p><p>Distribution</p><p>Mediterranean Sea and South European Atlantic Shelf (e.g. Noyer and Becerro 2012). This study expands the distribution of S. lamella northwards.</p><p>Ecology</p><p>In the Mediterranean Sea, S. lamella is well known to be distributed mainly along coralligenous cliffs or on deep mesophotic bottoms (Vacelet 1959, Pronzato and Manconi 2008). In the Northeast Atlantic, S. lamella has mostly been found in semi-dark and dark cavities, sometimes in crevices found along shallow water cliffs or under rocks. All specimens reported here were found in habitats with very high wave energy, which might explain why no large or fan-shaped individuals were observed. This sponge has a patchy distribution, and it lives in syntopy with S. maitasuna . In caves, this species is commonly associated with numerous other sponges, solitary scleractinians, such as Leptopsammia pruvoti Lacaze-Duthiers, 1897 ?, or zoanthids, such as Parazoanthus axinellae (Schmidt, 1862) . No sign of epibiosis or predation was observed. No reproductive element was recorded in the processed specimens.</p><p>Taxonomic remarks</p><p>The 15 specimens examined from the Northeast Atlantic differ slightly in their growth form from S. lamella in the Mediterranean Sea, but their skeletons are identical. Openings of the oscula are also twice as large in the Atlantic specimens. They all present in their ectosome a rather thick layer of foreign spicules and debris, which is an unusual structure among Spongia that was not observed in S. maitasuna . Moreover, the overall network of fibres cannot be confounded with the specific honeycomb-shaped skeleton of the new species. A rather organized ectosomal skeleton made of foreign spicules and debris is known only in Coscinoderma and Leiosella, but in these cases, this structure is much thicker and the organization of the choanosome skeleton very different (e.g. intertwined secondary fibres in the Mediterranean C. sporadense). Atlantic and Mediterranean specimens of S. lamella share the same common simple and irregular primary fibres, with inclusions of foreign spicules and debris, easily observable from the inner part of the sponge to the surface. They remain simple until the formation of conules, whereas in other Spongia species they become anastomosed at this level. The size of the primary fibres appears a little more variable in the Atlantic specimens (30–120 µm in diameter) than in the Mediterranean specimens (50–100 µm in diameter). Foreign spicules and debris seem to be more abundant in Atlantic specimens. In comparison, the primary fibres of S. virgultosa, which has an unusual encrusting papillate growth form for Spongia, are also full of mineral debris, but they are rare and significantly thinner (40–50 µm in diameter) than in S. lamella .</p><p>Therefore, the slight morphological differences recorded between Atlantic and Mediterranean specimens (growth form, abundance of debris, and variation in the size of primary fibres) do not seem to us to be sufficient to consider the Atlantic specimens as a different species and are thus interpreted as variations attributable to the different environmental contexts.</p><p>DNA analysis of studied species</p><p>GenBank accession numbers for all sequences generated or downloaded in this study are presented in Table 1. For the 28S (C2–D2 fragment) and CO1 (dgLCO1490–dgHCO2198 fragment) molecular markers, we obtained, respectively, 18 and 3 sequences of S. maitasuna, 21 and 14 sequences of S. lamella, 14 and 5 sequences for S. officinalis, 8 and 2 sequences of S. mollissima, 4 and 3 sequences of H. communis, 1 and 1 sequence of S. virgultosa, and only 1 sequence of S. zimocca (28S) and 1 sequence of C. sporadense (28S).</p><p>Molecular markers allowed us to build highly congruent trees, although the phylogenetic reconstruction with 28S gave a better resolution (Fig. 6) than CO1 (Supporting Information, Fig. S1). Both methods (NJ and ML) of phylogenetic reconstruction using 28S produced the same topology: 70 Spongiidae specimens divided in two, with only one sequence of C. sporadense on one side and 69 specimens of Spongia and Hippospongia on the other side (Fig. 6). This set of specimens can be divided into three different groups. The first group is well supported by both methods of phylogenetic constructions (NJ and ML). It is composed by the proposed new species S. maitasuna, individuals from the two sampled localities presenting the same sequence of 28S, and by a well-supported group made of one sequence of S. zimocca, two downloaded sequences of S. nitens, and three sequences obtained in the present study of S. cf. officinalis from Ceuta. Spongia maitasuna thus appears closely related to this group, with the p-distance being very low between the proposed new species and S. zimocca (1.05%, four substitutions), S. nitens, and S. cf. officinalis (0.79%, the substitutions). The second group is weakly supported, whatever the method of construction. It includes S. virgultosa and a well-supported group of all S. lamella from the Mediterranean and Northeast Atlantic. With only one exception,thesequencedindividualsaregroupedbygeographical origin. The Mediterranean sequence (20200602-MRS-MTP8) mixed with the Atlantic presents a p-distance of 0.26% (one substitution) from the rest of the Mediterranean S. lamella . In comparison, S. virgultosa presents a p-distance of 13.4% (51 substitutions) from all S. lamella . The third group is composed by a weakly supported set of sponges, including all S. mollissima, S. officinalis, and H. communis . The genetic distances between these three species are low, with a minimal p-distance of 0.26% between S. mollissima and H. communis from Tunisia, and 0.79– 1.57% (three to six substitutions) between S. mollissima and S. officinalis or H. communis .</p><p>A phylogenetic tree was made by concatenating all sequences available for both 28S and CO1 (Fig. 7), with the exception of two samples of S. zimocca and H. communis for which only 28S could be amplified. The topology of the concatenated tree is congruent with the 28S tree. Both phylogenetic reconstruction methods used here (BI and ML) presented the same pattern. Thirty specimens are grouped into two different groups representing the Spongiidae family. Group 1 is composed by one specimen of the genus Coscinoderma, whereas group 2 is composed by 29 specimens of Spongia and Hippospongia . The proposed new species, S. maitasuna, forms a well-supported group in both phylogenetic analyses and still separate but relatively close to S. zimocca, S. nitens, and S. cf. officinalis from Ceuta. All S. lamella form a well-supported group, which still appears separate by geographical origin. In this representation, S. officinalis and H. communis appear mixed in a subgroup also containing S. mollissima .</p><p>Chemical fingerprints of the studied species</p><p>We evaluated whether Spongiidae could be organized and grouped based on their chemical fingerprints and whether this chemical-based classification was congruent with the molecular phylogeny. Using the first aligned feature list, containing 915 chemical signals, we performed hierarchical clustering analysis to group sponge extracts according to their chemical similarities.</p><p>The dendrogram (Fig. 8) shows distinct groups of extracts all attributed to three distinct groups of species. The first group (group 1) includes all specimens of the proposed new species, S. maitasuna, in a subgroup, and S. nitens and S. zimocca in another subgroup. No sample of S. cf. officinalis from Ceuta (Strait of Gibraltar) was available for this analysis. The second group (group 2) includes all specimens of S. lamella, still separated into two subgroups according to their geographical origin. The Mediterranean exception (20200602-MRS-MTP8) that was grouping in the 28S phylogenetic with Atlantic specimens is grouped here with all other Mediterranean samples of the same species. The third main group (group 3) includes S. officinalis, S. mollissima, and H. communis, but in the case of this metabolomic analysis, all species formed well-separated subgroups. Thus, although its typology cannot be compared with those of the phylogenetic trees, the groupings presented in the dendrogram resulting from the metabolomic analysis are well congruent with those permitted by the morphological and genetic analyses.</p><p>The heatmap (Fig. 9) complements these results by depicting the whole set of chemical features that participated in the sample classification. This representation illustrates clearly that extracts from S. maitasuna contain a distinct set of chemical features compared with other Spongiidae samples. Next, we assessed the distribution of features within the three identified groups of species to evaluate the chemical diversity and number of unique chemical signals characterizing each species.</p><p>To that end, a second aligned feature list was generated from pooled extracts from each identified taxonomic unit and contained 240 features. Those features represent the most abundant and redundant chemical signals in each taxonomic unit. Venn diagrams were constructed to depict the number of chemical features shared between species as opposed to those that were detected in one or two species (Fig. 10).</p><p>The lowest number of features was detected for group 1, for which 45 chemical signals were shared between all three species ( S. maitasuna, S. nitens, and S. zimocca). A total of 16 unique signals were detected for S. maitasuna, illustrating a much higher chemical diversity and uniqueness in extracts than in S. nitens (2) and S. zimocca (1). At this stage, none of these features has been identified formally, because the purpose of this analysis was not to assign the associated molecular structure to each feature. However, it is certain that none of the well-known and referenced compounds from groups 2 and 3 (as shown later) were detected among the signals.</p><p>The highest number of features was associated with group 2, represented by S. lamella from different locations (Mediterranean and Atlantic). There were between 80% and 85% similarities of detected signals (111 in total) in all extracts. Among the detected chemical features, nitenin is one of the most abundant and well-known furanoterpenoids from this species (Noyer et al. 2011). Extracts from the Mediterranean specimens present a greater number of unique detected signals (13) than those from the Atlantic specimens (5 and 4).</p><p>Group 3, represented by S. officinalis, S. mollissima, and H. communis, also contains a relatively high number of detected features compared with the proposed new species. Extracts from these three species have 98 chemical signals in common. Some other well-known and characteristic furanoterpenoid compounds could be identified among these signals, such as furospongin-1, furospongin-4, and demethylfurospongin-4 (Bauvais et al. 2017). Extracts from S. mollissima have the highest number of unique detected signals (23), compared with S. officinalis (3) and H. communis (9). Among all Spongiidae, S. lamella is the species presenting the highest chemical diversity, with a total of 170 chemical signals (including its unique chemical signals and others shared with other Spongiidae). In comparison, extracts from the proposed new species contain 151 chemical features, and&gt;10% of them are unique and thus have unknown molecular structures. After undergoing purification and determination of their chemical/molecular structure, they have the potential to become chemotaxonomic markers.</p><p>Therefore, the hierarchical classification of species, based on their chemical fingerprints, is congruent with the phylogenetic trees previously presented. Each species can be differentiated by its chemical fingerprints associated with a unique set of chemical features that might serve as chemotaxonomic markers. These putative new phenotypic traits would need to be annotated or identified after isolation, purification, and structural analysis by means of additional spectroscopic (e.g. nuclear magnetic resonance) and spectrometric methods.</p></div>	https://treatment.plazi.org/id/6864879DFFC1A77FFC2FFD13FB45F848	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grenier, Marie;Simmler, Charlotte;Chevaldonné, Pierre;Callizot, Noëlle;Pérez, Thierry	Grenier, Marie, Simmler, Charlotte, Chevaldonné, Pierre, Callizot, Noëlle, Pérez, Thierry (2024): Searching for Mediterranean bath sponges (Demospongiae: Dictyoceratida: Spongiidae) in the Northeast Atlantic reveals a new species: an integrative taxonomic approach. Zoological Journal of the Linnean Society 202: 1-23, DOI: 10.1093/zoolinnean/zlad166
