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Poecillastra macquariensis Kelly & Cárdenas sp. nov. urn:lsid:zoobank.org:act: 78B12A37-93E0-4CA9-9317-81F835B112FE Fig. 5 Etymology Named for the type location of this species, the Macquarie Ridge. Type material Holotype NEW ZEALAND • Subantarctic region of New Zealand, Seamount 5, Macquarie Ridge, NIWA Station TAN0803/48; 51.096° S, 161.976° E; depth 462–524 m; 4 Apr. 2008; NIWA 52640 leg.; epibenthic sled; NIWA. Description Solid stalk of sponge of unknown morphology, 15 mm in diameter, 20 mm high, expanding on the broken, upper surface, sides of stalk sculpted, attachment base contains patches of substrate (Fig. 5A). Surface hispid and scratchy to the touch; texture firm, incompressible. Colour in preservative tan. Skeleton Stalk composed of huge swathes of contort oxeas and triaenes between which are abundant microscleres. Spicules MEGASCLERES (Fig. 5 D–E) Abundant contort to sinuous oxeas (Fig. 5D) with slightly rounded tips, 3725(2125‾5750) × 53(30‾70) µm; medium-shafted triaenes (Fig. 5E), rhabd slightly curved, tapering to a sharp tip, 852(550‾1225) µm, clads of slightly uneven length, slightly curved downwards, 578(450‾680) µm, overall cladome width, about 900‾1360 µm long, ranging to pseudocalthrops. Broken true oxeas are evident but unmeasurable. MICROSCLERES (Fig. 5 B–C, F–G) Microxeas (Fig. 5 B–C), straight to slightly curved, roughened, abundant, 332(260‾420) × 7(5‾8) µm, n = 20; plesiasters (Fig. 5F), with 3‾5 microspined blunt-tipped rays, overall 67(50‾100), ray length 37(25‾60) µm, n = 10; metaster- to amphiaster- to spiraster-like streptasters (Fig. 5G), with long, microspined rays, abundant, 19(15‾20) µm long. Distribution Macquarie Ridge. Substrate, depth range and ecology Attached to rock substrate; depth 462– 524 m. Remarks The specimen is the attachment base of a sponge of unknown morphology, but it clearly differs from the holotype of Poecillastra ducitriaena sp. nov. in having a very hispid, crisp, scratchy surface, indicating a reduction of the ectosomal crust of microscleres, and the abundance of large megascleres. It is similar to Poecillastra ducitriaena sp. nov. in the possession of abundant contort oxeas in the stalk, but differs in the lack of straight oxeas in the stalk and the much larger dimensions of all the spicules: the contort oxeas are up to 2000 µm longer, on average, in Poecillastra macquariensis sp. nov., and the triaenes are about double the size of those in Poecillastra ducitriaena sp. nov., and much more abundant, the microxeas are about ten times larger, and the sponge contains plesiasters, absent in Poecillastra ducitriaena sp. nov. Because our knowledge of Poecillastra in the New Zealand region is reasonable (see above), we have made the decision to record and name this second Poecillastra species, despite our lack of information on the body shape, and because surface texture, spicule types and dimensions are so different from those of Poecillastra ducitriaena sp. nov.

Family Vulcanellidae C��rdenas, Xavier, Reveillaud, Schander & Rapp, 2011 Diagnosis (modified from C��rdenas et al. 2011) Astrophorina with calthrops, short-shafted triaenes or long-shafted triaenes, in addition to large oxeas and contort or sinuous strongyloxeas.Aster microscleres include several categories of streptasters (spirasters, metasters, amphiasters, and plesiasters). Monaxonic spicules consist of one to three categories of spiny microxeas or microstrongyles., Published as part of Kelly, Michelle, C��rdenas, Paco, Rush, Nicola, Sim-Smith, Carina, Macpherson, Diana, Page, Mike & Bell, Lori J., 2019, Molecular study supports the position of the New Zealand endemic genus Lamellomorpha in the family Vulcanellidae (Porifera, Demospongiae, Tetractinellida), with the description of three new species, pp. 1-25 in European Journal of Taxonomy 506 on page 5, DOI: 10.5852/ejt.2019.506, http://zenodo.org/record/2612959, {"references":["Cardenas P., Xavier J. R., Reveillaud J., Schander C. & Rapp H. T. 2011. Molecular Phylogeny of the Astrophorida (Porifera, Demospongiae) Reveals an unexpected high level of spicule homoplasy. PLoS ONE 6 (4): e 18318. https: // doi. org / 10.1371 / journal. pone. 0018318"]}

Poecillastra ducitriaena Kelly & Cárdenas sp. nov. urn:lsid:zoobank.org:act: E3C570FA-832A-4FDB-97D3-5CDA250A3797 Figs 1, 4, 6 Etymology Named for the possession of triaenes in addition to the apparent spiculation of Lamellomorpha, and their guide to the phylogenetic origins of this species (‘ duci -’ in the sense of a guide). Type material Holotype NEW ZEALAND • Subantarctic region of New Zealand, east of Snares Island Platform, NIWA Station TRIP3072 /8; 48.5° S, 168.0° E; depth 125–213 m; 21 Oct. 2010; NIWA 61944 leg.; fish bottom trawl; UPSZTY 178604 (a small piece of the holotype preserved in 70% ethanol, as well as a spicule preparation), NIWA. Type locality Subantarctic region of New Zealand, Snares Island Platform, depth 125– 213 m. Description Multilamellate, foliose, fan sponge (Fig. 4A), 160 mm high, 104 mm wide, with a short thick stalk about 2 cm thick. Lamella up to 2 cm thick in places, attenuating to curled margins. Oscules were not visible on the holotype. Cribriporal pore areas are widespread between parallel tangential tracts of oxeas; individual pores 80–160 µm in diameter, on both sides of the lamella. Texture firm, compressible, flexible, granular and smooth to the touch. Colour in preservative tan. Skeleton Choanosome composed of huge swathes of long straight oxeas, radiating through the lamella, terminating below the surface (Fig. 6E). Contort oxeas are found in the stalk. Ectosome, relatively thick, packed with microxeas and perforated by ostia. Spicules MEGASCLERES (Fig. 4 B–C) Oxeas with long fine attenuated tips, 1725(1150‾2271) × 16(8‾22) µm; contort oxeas in the stalk, 1801(1095‾2671) × 13(5‾17) µm; short-shafted triaenes, relatively uncommon, rhabd straight, attenuating, 290(260‾300) µm, clads curved or acutely bent, occasionally with bifurcating tips, 232(200‾250) µm, cladome width 400‾500 µm long. MICROSCLERES (Fig. 4 D–F) Microxeas, heavily microspined, sometimes faintly centrotylote and acutely centrally bent, sharp ended, abundant, 38(28‾46) × 3(2‾4) µm; metaster- to amphiaster-like streptasters with long, microspined rays, rare, 8(5‾12) µm long; spirasters with dense, short rays, rare, 5‾7 µm long. Distribution East of Snares Island Platform. Substrate, depth range and ecology Attached by a thick stalk to sediment covered rocky substrate, depth 125‾ 213 m. DNA barcodes COI. Holotype (MK033626); 28 bp difference with the COI of L. strongylata; 24 bp difference with the COI of Poecillastra compressa (Bowerbank, 1866) (HM592675). 28S (C1-C2). Holotype (MK 033144); 5 bp difference with the 28S (C1-C2) of L. strongylata; 3 bp difference with the 28S (C1-C2) of Poecillastra compressa (HM592757). Remarks This remarkable sponge was first identified as a third species of Lamellomorpha, as it appeared to have an almost identical form (stalked, multilamellar fan), a megasclere complement of straight and contort oxeas (more or less restricted to the stalk), small centrotylote microxeas, and metasters (albeit rare). Because the short-shafted triaenes were relatively uncommon, it was initially hypothesised that this was a species of Lamellomorpha with rudimentary triaenes that ‘showed the way’ to the true affinity of the genus with other triaene-bearing Tetractinellida. However, molecular sequencing consistently linked Poecillastra ducitriaena sp. nov. with other Poecillastra species (Fig. 7). Despite its consistency with two independent markers, we note that this grouping is not supported (bootstrap of 60 for COI, of 10 with 28S). This may be due to the absence of other subantarctic Poecillastra species in our sampling which Poecillastra ducitriaena sp. nov. may be closer to (P. Cárdenas, unpublished results). Although not fully documented (Kelly et al. 2009), our knowledge of Poecillastra in the New Zealand region is reasonable and includes what we consider to be Poecillastra laminaris (Sollas, 1886) (Zeng et al. 2016) and Poecillastra schulzei (Sollas, 1886). While several undescribed species are known from the New Zealand EEZ, no specimens are known that contain the characteristic contort oxeas of Poecillastra ducitriaena sp. nov.

This revision of the classification of eukaryotes follows that of Adl et al., 2012 [J. Euk. Microbiol. 59(5)] and retains an emphasis on protists. Changes since have improved the resolution of many nodes in phylogenetic analyses. For some clades even families are being clearly resolved. As we had predicted, environmental sampling in the intervening years has massively increased the genetic information at hand. Consequently, we have discovered novel clades, exciting new genera and uncovered a massive species level diversity beyond the morphological species descriptions. Several clades known from environmental samples only have now found their home. Sampling soils, deeper marine waters and the deep sea will continue to fill us with surprises. The main changes in this revision are the confirmation that eukaryotes form at least two domains, the loss of monophyly in the Excavata, robust support for the Haptista and Cryptista. We provide suggested primer sets for DNA sequences from environmental samples that are effective for each clade. We have provided a guide to trophic functional guilds in an appendix, to facilitate the interpretation of environmental samples, and a standardized taxonomic guide for East Asian users. (VLID)3393271

Discorhabdella pseudaster n. sp. Holotype. MNHN-IP-2015-1083, station DW 4788, Warén dredge, 22/01/2017, Banc du Geyser, 12°22’ S, 46°25’ E, 346–349 m depth, rocks and pebbles. Field# PMG 0 5, coll. C. Debitus, ethanol 95%. Description. A minute, incrusting sponge, 4 x 2.5 mm and 200 µm thick, hispid (Fig. 2 A). Color grayish white alive and in alcohol. No visible aperture. A stoloniferous calcified octocoral, possibly Scleranthellia (Gary Williams, pers. com.), is growing on the same rock. Skeleton. The skeleton is made of long styles with a barely inflated head bearing low, round tubercles. These styles are disposed vertically on the substratum and give the long hispidation. They are surrounded by smaller tylostyles irregularly arranged, sometimes forming poorly defined bouquets. Pseudoastrose acanthostyles are arranged vertically, with the head on the substratum. Pseudoasters and isochelae are dispersed in the whole choanosome. Spicules. (1) Subtylostyles (Fig. 2 B), straight or slightly curved, with a barely tuberculated head, all broken in the spicule or skeleton slides. Axial canal ending in a small vesicle in the head (not divided), although faint, radiating darker lines could be seen (Fig. 3 A). Maximum length of the broken spicules: 600 µm, diameter 40–56 µm near the base.(2) Tylostyles (Fig. 2 C), straight or slightly curved, with a well-marked oval head, 240– 370 x 9–10 µm.(3) Pseudoastrose acanthostyles (Fig. 2 D–F). Spine-like rays of the head obtuse, those of the end of the shaft generally more acute. Total length 35–45 µm, diameter of the head 35–40 µm, shaft 20-25 x 12 µm with spines at the extremity. Axial canal barely visible by transparency (not divided in the head), although faint, radiating darker lines could be seen in the head (Fig. 3 B). Some rare spicules, likely juvenile, are a little smaller (e.g. 28 µm in length), and have more acute spines.(4) Pseudoasters (Fig. 2 G–I), similar to spherasters, very abundant, 12.5–18 µm in diameter, with conical, sharp spine-like rays, 2.5–4 µm long. No visible axial canals in the spines.(5) Unguiferous cleistochelae (Fig. 2 J), small, 12–15 µm. Shaft straight, 4 or 5 teeth nearly in contact, often with the end on one side blunt and the other slightly indented. The shaft bears a conspicuous lamella, attached by a part of its length, similar to a hatchet. The free part of the hatchet is most often directed towards the nonindented teeth, but the opposite case has also been observed. Etymology. From Latin pseudo (meaning false) and aster. Remarks. The presence of chelae makes this sponge a Poecilosclerida. This minute sponge clearly belongs to the genus Discorhabdella by its shape, skeleton and especially the diagnostic club-shaped pseudoastrose acanthostyles. Unfortunately, the unique specimen is too small to allow obtaining sequences, so the affinities between Discorhabdella and Crambe remain likely (Maldonado & Uriz 1996), but not formerly demonstrated. It is rather puzzling to find “aster” spicules in a poecilosclerid, which moreover have the same position in the skeleton as true asters in non-poecilosclerid encrusting sponges (e.g. Spirastrellidae, Chondrillida or Tethyida). The true nature of these asters is questionable. It is essential in taxonomy and phylogeny to use homologous morphological characters, and that is not always obvious as clearly shown by Fromont & Bergquist (1990) with the example of sigma microscleres or acanthose megascleres. Here, the most likely explanation is that the “asters” derive from the pseudoastrose acanthostyles with reduced shaft typical of Discorhabdella by complete loss of the shaft. The presence of “asteroid” corpuscules somewhat similar to asters have been described in two other members of the family Crambeidae, Crambe tuberosa Maldonado & Benito, 1991, and Crambe chilensis Esteves et al., 2007; these spicules, which in C. tuberosa have axial filaments in the actines, were interpreted as developmental stages of desmas, otherwise present in Crambe spp. (Maldonado & Benito 1991; Uriz & Maldonado 1995; Maldonado & Uriz 1996). *For D. incrustans, measurements by van Soest (2002) from the holotype. ** From Hinde & Holmes (1892): p. 194, Pl. VIII, Fig 29. *** Tentative identification by Łukowiak (2015) from one Eocene spicule. It has been suggested that pseudoastrose acanthostyles are polyaxial spicules, on the basis of SEM observations (Uriz & Maldonado 1995; Maldonado & Uriz 1996). However, in our opinion, the presence of axial canals in the spines of the head of the pseudoastrose acanthostyles remains doubtful: in a few immature spicules, some actines of the head show a very small hole at their end (Fig. 2 E), but a canal could not be seen in transparency under a light microscope in the actin of mature spicules, while the axial canal of the shaft of the same spicules is clearly visible. Similarly, the subtylostyles with a slightly tuberculated head show by transparency a clear axial canal ending in a vesicle in the head, without division. However, it must be noted that from this vesicle, there are very faint radiating lines, which could indicate remains of axial filaments present during the spicule formation (Fig. 3 A). The same very faint radiating lines are barely visible in the head of pseudoastrose acanthostyles (Fig. 3 B). It is also possible that these extra spines and bulges are produced by secondary axial canals not conected to the main shaft axial canal, but somehow added later, just like the axial canals in the rosette formation of geodiid sterrasters (Cárdenas & Rapp 2013). Then, they could be filled in some cases and be invisible in our observations. To conclude, we raise doubts as to the polyaxial nature of these spicules and warrant more observations at the interface between the tip of the tylostyles and their siliceous additions (spines and bulges). There are currently five known species of Discorhabdella from the Pacific coast of Panama, Western Mediterranean, the Azores and New Zealand (Table 1). Our new species differs from the other species by several characters: (1) presence of two types of pseudoastrose acanthostyles, one of which (pseudoaster) has entirely lost the shaft and is reduced to the inflated, spinose head, similar to an aster; (2) microscleres composed only of a single isochelae type, with sigma and oxydiscorhabd missing. The microsclere composition in Discorhabdella spp. is quite diverse, but only D. tuberosocapitatum (Topsent, 1890) has only isochelae. The isochelae of the new species clearly differ from those of D. tuberosocapitatum, which is from Macaronesia (N-E Atlantic), by a different shape, with teeth nearly joining and the shaft bearing a lamella. This type of isochelae shape is unknown in the other species of the genus. These isochelae, very unusual in Poecilosclerida, remind of the ‘palmate-anchorate-arcuate’ evolutionary sequence suggested by Maldonado & Uriz (1996) for aberrant chelae morphologies in Crambeidae genera Crambe and Discorhabdella. However, the isochelae of D. pseudaster n. sp., although possibly included in such a sequence, appear unique in Crambeidae, and more generally in Poecilosclerida. The most original character is the presence of a lamella on the shaft, a character rarely observed in isochelae. To our knowledge, this character is known only in the small isochelae of Hymenancora sirventi (Topsent, 1927) in Myxillidae. An interesting other example could be one isochela found by Hinde & Holmes (1892, p. 200, Pl. IX, 39) in sediment from the Late Eocene, and tentatively attributed to Melonanchora elliptica Carter, 1874. According to their description and drawing, this chela somewhat resembles those of the new species, although larger (66 µm) and bearing two lamellae instead of a single one. The presence of typical Discorhabdella spicules (tuberculate tylostyles and pseudoastrose acanthostyles) in these same Eocene sediments (Hinde & Holmes 1892; Łukowiak 2015) suggests that this chela may belong to this genus. The discovery of this 4 mm long new species emphasizes the need to carefully examine minute, inconspicuous, incrusting sponges on such rocky bathyal substrates, which certainly represent an underestimated source of biodiversity.

Caminella caboverdensis sp. nov. (Figure 7, Table 1) Holotype. RMNH 3810, Cape Verde, NW of São Vincente (16.9167, -25.0333), 75 m, CANCAP VI expedition (on board HMS Tydeman), station 6.174, 22.06.1982, bottom sand, collecting gear: 1.2 m Agassiz trawl, originally identified as Isops intuta by R. van Soest (unpublished). External morphology (Fig. 7D). Similar to C. intuta. Two dark brown mottled pieces (in ethanol); it is not clear if they belong to the same specimen. Very thin cortex (150 µm). Consistency fleshy, compressible. Spicules. (Fig. 7A–C, Table 1, Supp. Mat. Appendix 1). (a) oxeas, 800– 1525 x 8–23 µm, sometimes bent to double bent (‘wavy’); (b) dichotriaenes (rhabdome: 650–900 x 20–50 µm; protoclad: 80–150 µm, deuteroclad: 50– 230 µm); (c) spherical, immature and mature sterrasters, 35–45 µm; (d) oxyasters, 8–42 µm in diameter, 4–9 actins, actins are finely acanthose; center more or less developed; (e) spherasters, 2.5–8 µm in diameter, spiny, regular with large centrum, occasionally look like spherules. Bathymetric range. 75 m. DNA barcoding. COI. The holotype (MH477614) had a difference of 9 bp with the COI of C. intuta, and 4 bp with C. pustula sp. nov. 28S (C1-C2). The holotype (MH478116) has the same 3 bp difference with the cave specimen from Portugal and with C. pustula sp. nov. Submitted to the Sponge Barcoding Project with accession number 1778. Etymology. Named after its type locality, the Cape Verde Islands (‘Cabo Verde’ in Portuguese). Remarks. Although we only have one specimen, with an external and spicule morphology that closely resembles that of C. intuta, we are convinced it belongs to a new species, based on the important genetic difference found in COI (9 bp in Folmer fragment, 658 bp) and 28S (3 bp in C1-C2, 369 bp). We also noticed two spicule differences: 1) sterrasters are smaller than in C. intuta (35–45 µm versus 40–84 µm); 2) oxeas can be much more bent than in C. intuta. These genetic and morphological differences need to be confirmed with additional material. The fact that this specimen has a substantial number of immature sterrasters and few spherules suggests that it might have been living and collected in a silica-limited environment.

Caminella pustula sp. nov. (Figures 8–9, Table 1) Holotype. MNHN-IP-2008-4, Seamount 2 expedition, St. DW184, Banc d’Hyères, 31°24’N, 28°52’W, 705 m, 16.01.1993, coll. Gofas, Métivier & Warén, barrel 2-1. Paratypes. MNHN-IP-2008-4 (four other specimens); MNHN-IP-2008-8, Seamount 2, St. CP151, Grand Banc Meteor, 30°12’N, 28°25’W, 585 m, 11.01.1993, coll. Gofas, Métivier & Warén, barrel 2-1; MNHN-IP-2008- 148 (2 specimens), Seamount 2, St. DW265, Banc Atlantis, 34°29’N, 30°36’W, 545 m, 0 3. 0 2. 1993, coll. Gofas, Métivier & Warén, barrel 2-9; MNHN-IP-2008-149, Seamount 2, St. DW248, Banc Plato, 33°14’N, 29°32’W, 735 m, 0 1.02.1993, coll. Gofas, Métivier & Warén, barrel 2-9. Other material. MNCN 1.01/1017, El Cachucho (=Le Danois Bank), 44°02.6999' N, 05° 06.3436' W, 660 m, ethanol 96%, SponGES 0 617 expedition, station DR9, field#DR9-430, rock dredge, 16.06.2017, coll. P. Rios. External morphology (Fig. 8). Holotype is an elongated, globular sponge, 3.5 cm long (Fig. 8A, shown with arrow, 8C). MNHN-IP-2008-8 has a more irregular shape. In ethanol, surface color is cream to brown; choanosome is cream-colored. The specimen from El Cachucho is 0.7 x 0.6 cm and cream-colored alive (slightly browner in ethanol); it was growing on a Pachastrella sp. Cortex is 1–0.5 mm thick. Specimens are slightly compressible. Uniporal oscules are 0.5 mm wide, elevated up to 1 mm, resembling pimples, with a dark brown ring. Uniporal pores sometimes with a dark ring as well, can also be elevated, but less than oscules. Most specimens are growing on coral branches. Spicules. (Fig. 9, Table 1, Supp. Mat. Appendix 1). (a) oxeas,> 2600 x 20 –40 µm; (b) dichotriaenes (rhabdome: 581–1100 x 43 –75 µm; protoclad: 70–296 µm; deuteroclad: 108–790 µm); (c) elongated mature sterrasters, 70–107 x 53 –92 µm; (d) oxyasters, 27–78 µm in diameter, 2–8 actins, the actins are finely acanthose; center usually well developed; (e) spiny spherules, 4–13 µm in diameter. Bathymetric range. 545–735 m. DNA barcoding. COI. MNCN 1.01/1017 (MH477615). There is a 6 bp difference with C. intuta and a 4 bp difference with C. caboverdensis. 28S (C1-D2). MNCN 1.01/1017 (MH478117). There is a 17 bp difference with C. intuta from Portugal and 3 bp difference with the shorter 28S (C1-C2) of C. caboverdensis. Submitted to the Sponge Barcoding Project with accession number 1780. Etymology. Named for the external surface, which is covered with ‘pimples’, pustula in Latin. Remarks. The key / diagnostic morphological character that identifies this species, is the elevated, pimple- shaped opening (either oscule or pore) with a brown tip. The spicules are overall much larger than in C. intuta and C. caboverdensis sp. nov. (Table 1, Supp. Mat. Appendix 1): i) the sterrasters are elongate (versus more or less spherical in the other two species) and much larger (70–107 µm in length versus 40–84 µm in C. intuta), ii) the oxyasters are 27–78 µm versus 5–27 µm in C. intuta and 8–42 µm in C. caboverdensis; they are sometimes reduced to only 2–4 actins versus usually>6 actins in C. intuta and C. caboverdensis sp. nov, iii) the dichotriaene rhabdomes are thicker (43–77 µm versus 17–60 µm for C. intuta and 20–50 µm in C. caboverdensis and iv) the dichotriaenes have longer clades (especially the deuteroclades). Finally, C. pustula sp. nov. lives at greater depths than the other two species: 545–735 m versus 2–300 m for C. intuta, 75 m for C. caboverdensis.

Cárdenas, Paco, Vacelet, Jean, Chevaldonné, Pierre, Pérez, Thierry, Xavier, Joana R. (2018): From marine caves to the deep sea, a new look at Caminella (Demospongiae, Geodiidae) in the Atlanto-Mediterranean region. Zootaxa 4466 (1): 174-196, DOI: https://doi.org/10.11646/zootaxa.4466.1.14

Genus Caminella Lendenfeld, 1894 Type species. Caminella loricata Lendenfeld, 1894 (by monotypy) Diagnosis (new). Erylinae with sterrasters with actins linked to each other through bridges (giving a brain-like surface) and cortical spherasters and /or spherules., Published as part of C��rdenas, Paco, Vacelet, Jean, Chevaldonn��, Pierre, P��rez, Thierry & Xavier, Joana R., 2018, From marine caves to the deep sea, a new look at Caminella (Demospongiae, Geodiidae) in the Atlanto-Mediterranean region, pp. 174-196 in Zootaxa 4466 (1) on page 177, DOI: 10.11646/zootaxa.4466.1.14, http://zenodo.org/record/1454321

Caminella intuta (Topsent, 1892) (Figures 2–6, Table 1) Synonyms. Cydonium intutum Topsent, 1892: Topsent 1892, p. XVIII. Isops intuta (Topsent, 1892): Topsent 1893, p. XLIII; Topsent 1894, p. 336, pl. XI, pl. XVI; Lendenfeld 1903, p. 95–96; Sarà 1961, p. 31; Boury-Esnault 1971, p. 296; Pouliquen 1972, Table 1; Templado et al. 1986, Table 1. Isops intutus (Topsent, 1892): Vosmaer 1933, p. 145. Isops intuta (Vosmaer, 1894): Maldonado 1992, Table 1 (mistake in authority). Caminella intuta (Topsent, 1892): Cárdenas et al. 2011; Sitjà & Maldonado 2014, Table 2. Isops maculosus Vosmaer, 1894: Vosmaer 1894, p. 273–274. Isops maculosa Vosmaer, 1894: Lendenfeld 1903, p. 96. Caminella loricata Lendenfeld, 1894: Lendenfeld 1894, p. 150–151, pl. II, pl. III, pl. VIII; Lendenfeld 1903, p. 89–90. Not Isops intuta (Topsent, 1892): Boury-Esnault et al. 1994, p. 41 = Geodia sp. (this study) Holotype. MNHN DT-2290 (slide with sections, in bad state), Cap l’Abeille, Banyuls, France, 25-30 m. Material examined. Caminella intuta. France: SME, collection ‘Topsent’, box 1, slide#12 labeled “ Isops intuta ”, Banyuls-sur-Mer (type locality), dredge; SME #90S, dry piece and slide, 17.03.1958, Le Petit Congloué, Archipel de Riou, Marseille, cliff facing NE, 40 m, coll. J. Vacelet; SME #193S, wet specimen and slide, 26.08.1998, Gameau cave, La Ciotat, 6-8 m, preserved in formalin, coll. J. Vacelet; SME PL617PC-7 (Fig. 2C, 2E) and PL617PC-10b, 18.05.2016, Cosquer cave, Calanque de la Triperie, Parc National des Calanques, coll. P. Chevaldonné, ethanol 96 %; Lebanon: SME 21/09/2002 -56a, 5/07/2003 -1 (sac 9), 13/07/2003 -2, preserved in formalin, Chak El Hatab (north of Selaata, Lebanon), " lithistid cave ", in dark area, 2-3 m, colls. T. Pérez and J. Vacelet; SME field# 080524 - Lb 2-03, 24.05.2008, Chak El Hatab, “lithistid cave”, 2 m, coll. T. Pérez, ethanol; Alboran Sea: field# Sp175, station BV 41, 35.99362, -2.86795, 102-112 m, beam trawl, INDEMARES- ALBORAN, 21.07.2012, coll. S. Gofas, identified by Sitja & Maldonado (2014), preserved in formalin (Fig. 2F); Portugal: ZMAPOR 21653, field# B.05.09.268, Gruta do Carreiro Maldito, Berlengas, Portugal, 6-8 m, 20.09.2005, ethanol 96%, coll. A. Cunha (Fig. 2D, 3F–G). Isops maculosus, lectotype (here designated), RMNH Por 644 (not seen), Gulf of Naples, between Capri and Naples, 150-200 m, wet specimen labeled as ' Isops intutus Tops. Corallieri, 17 Mei 1884 coll. Vosmaer N.249', and 18 slides, some of which are labeled Isops maculosus, but all have the number 249. Slides are mostly histological sections, but two are spicule slides. According to GBIF records from the RMNH collection (https://www.gbif.org/ occurrence/search?taxon_key=5892880), there are at least seven other paralectotypes (here designated): RMNH Por 78, 645, 646, 647, 648, 649, 650; BMNH 1955.3.24.2 (seen), two fragments of paralectotype RMNH Por 647 (~2.5 x 1.5 cm), 12.12.1890, Gulf of Naples. Caminella loricata, holotype (not seen), ZMB Por 2423, Lesina (=Hvar), Adriatic Sea, Croatia, one small piece 1 cm x 0.5 cm, purchased in 1897, no slides; NHMW-3Zoo-EV-MP345 (Fig. 3A, 3C–E), slide from holotype with a thick section (seen high resolution pictures); NHMW-3Zoo-EV-MP346 (Fig. 3B), slide from holotype with histological sections (seen high resolution pictures). Geodia sp. SME, Balgim slide box ‘Tetractinellida’, three slides labeled ‘CP–63 (185)’ (two spicule preparations and one piece of cortex), off Morocco, 1510 m, originally identified as Isops intuta by M. J. Uriz (Boury-Esnault et al. 1994). External morphology and skeleton organization. (Figs. 2–3) Massive subglobular, up to 10 cm wide (Fig. 2C) with smooth, clean surface. Cave specimens from France and Portugal area are brown, to dark brown (Fig. 2A, 2C, 2D). Cave specimens from Lebanon (Fig. 2B) are pure white (small specimens), white to cream or light brown (large specimens). Deep-sea specimens are cream-colored to brown. Internal color is white to light brown. Colors are retained in ethanol and formalin. One to several uniporal oscules (0.2–2 mm in diameter) can be present, each leading into a cloaca (Fig. 3F). Cave specimens from Lebanon generally have a single oscule at the top, more rarely two or three, while other cave specimens often have several oscules. Oscules can have a slightly elevated margin in life, especially visible in underwater photographs (Fig. 2A, 2B), often flush with surface after preservation (Fig. 2E). Surface punctured by numerous uniporal pores (20–200 µm in diameter in cave specimens; 20–55 µm in deep- sea specimen), which can have elevated surrounding walls (Fig. 2E), or not (Fig. 2F). Oscules and pores can be surrounded by a conspicuous dark ring (especially in cave specimens) (Fig. 2A, 2C), or not (deep-sea specimens) (Fig. 2F). Consistency fleshy. The cortex is thin (0.2–0.5 mm thick), more or less flexible and easily detachable from the underlying choanosome. The ectocortex is composed of a dense aggregation of spherasters/spherules while the endocortex is made of sterrasters (Fig. 3D–E). Dichotriaenes support the cortex, but do not cross it (Fig. 3G). In the choanosome, a few oxeas are more or less radially organized (Fig. 3F–G). In the choanosome, oxyasters and sterrasters are common while spherasters/spherules are rare and predominantly found around the canals. Spicules. (Figs. 4–6) (Table 1) (a) oxeas, a few styloids, 900– 2500 x 6 -50 µm; (b) dichotriaenes (rhabdome: 370– 2000 x 12 –60 µm; protoclad: 60–237 µm; deuteroclad: 25–410 µm), rarely orthotriaenes; (c) sterrasters, spherical to oval, 40–84 µm, surface without clear rosettes, instead the actins build bridges between them making a brain-like surface, which is then covered with small warts; immature sterrasters have blunt actins covered with small branches that sometimes link two actins together making a honeycomb surface (similar to Placospongia selenasters). From here on we will distinguish ‘young’ (not fully grown) from ‘immature’ (fully grown but underdeveloped) sterrasters; (d) oxyasters, 3–8 actins, 10–36 µm in diameter, actins are finely acanthose and blunt at the very tip; center is more or less developed; (e) spiny spherasters to spherules, different ratios depending on the specimens, sometimes with very irregular spherasters, 3–15 µm in diameter. We observed four types of spicule phenotypes depending on the origin of the specimens. 1) Cave specimens from France and Portugal, as well as C. loricata from Croatia had immature sterrasters associated with irregular to regular spherasters, which rarely become spherules (Figs. 3E, 4). Dichotriaenes can also be irregular; orthotriaenes are occasionally found. 2) Specimens from shallow-water but deeper (25–40 m) and not in caves (specimens from the type locality identified by Topsent and from 40 m in Marseille) have immature sterrasters associated with irregular to regular spherasters, many of which are spherules. 3) Cave specimens from Lebanon (all coming from the same cave) had mature sterrasters and only spherules. Megascleres are larger and robust (Fig. 5). 4) Deep-sea specimens (Isops maculosus paralectotype and specimen from the Alboran Sea) had mature sterrasters with regular spherasters, which have on average shorter actins, and often become spherules (Fig. 6). Bathymetric range. In caves, C. intuta lives in semi-dark to dark areas, where it can be found as shallow as 2 m depth. Outside caves, it has been recorded from 25–30 m in Banyuls-sur-Mer (Topsent, 1892) to 300 m in the Alboran Sea (Templado et al. 1986). DNA barcoding. COI. ZMAPOR 21653 (HM592740) and cave specimen from Lebanon (080524 -Lb2-03) had a 100% identical COI (MH477613). 28S (C1-D2). ZMAPOR 21653 (HM592804) and cave specimen from Lebanon (080524 -Lb2-03, MH478114) were 100 % identical. Both had a 5 bp difference with PL617PC-10b from Cosquer cave (MH478115). 18S. ZMAPOR 21653 (MH478118). ZMAPOR 21653 was submitted to the Sponge Barcoding Project (http://www.palaeontologie.geo.uni-muenchen.de/SBP/) with accession number 1777. Remarks. Suspicions of synonymy with Isops maculosus and Caminella loricata were raised early on by Topsent (1895, p. 580–581) who suggested that i) Isops maculosus might be a synonym of Isops intuta, although its colour seemed different and its sterrasters larger, ii) Caminella loricata might be a synonym of Isops intuta since the ‘microdesmen’ described by Lendenfeld (1894) were probably spherasters. However, Topsent (1895) did not conclude since he had not seen type material and Vosmaer (1894) had not given any measurements or illustrations of I. maculosus. This issue was later taken up by Vosmaer (1933), finally giving some spicule measurements of I. maculosus. He concluded that his species, I. maculosus and C. loricata, are junior synonyms of I. intuta. Once again, these conclusions were not based on the comparison of type material. The present study is the first one to examine the type material from Cydonium intutum, Isops maculosus, and Caminella loricata. We confirm that I. maculosus and C. loricata are junior synonyms of Caminella intuta. And yet, we do note some external morphology and spicule differences within C. intuta specimens, which will be discussed below. ……continued on the next page The singularity of Caminella sterrasters was foreseen by Topsent (1894) who noted that sterrasters had complex and ornate surfaces. Later Vosmaer (1933, p. 148) very accurately observed that it “appears as if spines of neighboring actins were fused together” but concludes this is probably not the case and that “the truth being that they are only crowded close together”. The unique pattern and microstructures of the sterrasters’ surfaces is revealed here for the first time with SEM (Figs. 4–7, 9). The surface of Caminella sterrasters is indeed quite different from Geodia sterrasters which have characteristic star shaped structures called ‘rosettes’ (4–8 µm in diameter) which do not fuse with one another, and can be smooth or covered with small warts (Cárdenas et al. 2009; Cárdenas et al. 2013). In Caminella, actins produce at their tips, small perpendicular bridges (Fig. 4D) that reach towards the other actins, creating a complex fused network, which gives first a honeycomb-like structure (especially when observed with an optical microscope) then a brain-like structure when these bridges thicken. Finally, these surfaces are covered with small warts. Interestingly, all cave specimens (Fig. 4) except the ones from Lebanon have immature sterrasters, which are very similar to immature sterrasters observed in shallow-water Geodiidae in Norway: Pachymatisma normani Sollas, 1888 and Geodia barretti Bowerbank, 1858 (Cárdenas & Rapp 2013). Cárdenas & Rapp (2013) hypothesize that the lower silica concentration in shallow waters is primarily responsible for the immature sterrasters and we may contemplate a similar hypothesis for most cave specimens, which are also amongst the shallowest specimens (6–8 m). Disrupted spiculogenesis is particularly important in the cave specimen from Berlengas, Portugal, where the sterrasters are the smallest (Table 1) and dichotriaenes quite irregular. On the contrary, the Lebanese specimens have larger and fully mature sterrasters (Fig. 5) and significantly thicker spicules (Table 1), which may be linked to higher availability of dissolved silica. Interestingly, the cave where the specimens were collected was densely populated by two species of lithistids (Pérez et al. 2004), which may support this hypothesis, since lithistids have higher needs of silica than other demosponges. Spicule variations between Mediterranean caves have already been observed for the lithistid tetractinellid Discodermia polymorpha (Pisera & Vacelet 2011) but in none of those cases did it seem clear that this species was lacking silica. Shallow-water C. intuta living outside caves and slightly deeper (25–40 m) still have immature sterrasters. Only deeper specimens (>100 m) (Fig. 6), living in waters where silica availability is usually higher, share the same mature sterrasters as the cave specimens from Lebanon. This is in accordance with other Geodiidae, where mature sterrasters are usually found below 40 m (Cárdenas & Rapp 2013). The cortical spheraster morphology also seems to be affected by environmental parameters, possibly silica availability: in cave specimens with immature sterrasters, spherasters are irregular, with small centers, and therefore few spherules; deep-sea specimens have more regular spherasters with larger centers (thus hiding the actins and making spherules); Lebanese specimens reach the extreme of having only spherules, probably indicative of high silica availability. To conclude, ectocortical microsclere morphology seems to be influenced by silica availability, such as in P. normani, but unlike in G. barretti (Cárdenas & Rapp 2013). We wonder whether this means that microscleres in Erylinae (C. intuta, P. normani) and Geodinae have different origins and/or spiculogenesis mechanisms. Like G. barretti and P. normani, C. intuta may be a deep-sea species that manages to survive in shallow-waters, despite lacking silica most of the time. Actually, C. intuta, like several other Mediterranean tetractinellids (e.g. Penares helleri (Schmidt, 1864), Penares euastrum (Schmidt, 1868), Geodia cydonium (L., 1767), Caminus vulcani (Schmidt, 1862), Calthropella pathologica (Schmidt, 1868), Thrombus abyssi (Carter, 1873), Alectona millari Carter, 1879, Discodermia polymorpha Pisera & Vacelet, 2011, Neoschrammeniella bowerbanki (Johnson, 1863) and Neophrissospongia nolitangere (Schmidt, 1870)) (Pisera & Vacelet 2011; Pouliquen 1972) are typically found from marine caves to the deep sea. Currently, we do not have enough data from either population of C. intuta to assess the connectivity between shallow and deep populations. However, Vosmaer (1933) did note differences between his deeper specimens and the shallower specimens of Topsent and Lendenfeld, especially the absence of raised openings and surrounding dark rings around them. Our morphological observations confirm there are differences in external morphology: the deep specimens (from the Alboran Sea and Gulf of Naples) share the same dull color, the absence of a brown ring around the openings, and openings without walls and very small pores (20– 50 µm). The significantly different 28S from one of the Cosquer Cave specimen (5 bp difference) is rather surprising since the Lebanese and Portuguese specimens have identical 28S. These underwater caves are only 20,000 years old (before, they were above water) so we doubt it could be due to a speciation event. Either i) the population that colonized this cave originated from a very different population than the ones present in Portugal and Lebanon (maybe from the deep sea) or ii) we sequenced a variant of 28S in this species, since intragenomic 28S polymorphism has been observed in some demosponges (Plotkin et al. 2017). Unfortunately, we did not manage to sequence COI from the Cosquer Cave specimens, but the morphology clearly suggests it is C. intuta. More cave and deep-sea specimens need to be examined and sequenced, especially for a population genetics study that should unveil the genetic ties between the marine cave, mesophotic zone and deep populations. We noticed that the surface of the sterrasters of the ‘ Isops intuta ’ specimen from the Balgim expedition (Boury-Esnault et al., 1994, fig. 112e–f) was not typical of Caminella. The two tiny Balgim specimens (3 mm in diameter), dredged off Morocco at 1510 m, clearly have sterrasters with rosettes typical of Geodia species. Furthermore, we examined three slides (2 spicule preparations and 1 piece of cortex) from Balgim specimen CP63– 185. This specimen has i) slightly asymmetrical oxeas (versus symmetrical in C. intuta), ii) shorter proto+deuteroclades, iii) presence of several anatriaenes and one protriaene (overlooked by the authors), but iv) we could not find the spherasters (although they are claimed to be present), instead we found v) smaller oxyasters (8– 12 versus 10–40 in C. intuta) with a different morphology than in C. intuta (Boury-Esnault et al., 1994, fig. 112d), the oxyasters are more spiny, especially at the tip of the actins. To conclude, we are sure that the Balgim specimens have been misidentified and are a Geodia sp., probably a juvenile, which might explain the rarity of sterrasters in the cortex., Published as part of Cárdenas, Paco, Vacelet, Jean, Chevaldonné, Pierre, Pérez, Thierry & Xavier, Joana R., 2018, From marine caves to the deep sea, a new look at Caminella (Demospongiae, Geodiidae) in the Atlanto-Mediterranean region, pp. 174-196 in Zootaxa 4466 (1) on pages 177-187, DOI: 10.11646/zootaxa.4466.1.14, http://zenodo.org/record/1454321, {"references":["TOpsENT, E. (1892) DIAgNOsEs d'EpONgEs NOuvELLEs dE LA MEdITERRANEE ET pLus pARTIcuLIeREmENT dE BANyuLs. Archives de Zoologie Experimentale et Generale, 2 (NOTEs ET REvuE 6), xvII - xxvIII.","TOpsENT, E. (1893) NOuvELLE sERIE dE dIAgNOsEs d'EpONgEs dE ROscOFF ET dE BANyuLs. Archives de Zoologie Experimentale et Generale, SERIEs 3, 1 (NOTEs ET REvuE 10), xxxIII - xLIII.","POuLIquEN, L. (1972) LEs spONgIAIREs dEs gROTTEs sOus-mARINEs dE LA REgION dE MARsEILLE: EcOLOgIE ET sysTEmATIquE. Tethys, 3 (4), 717 - 758.","VOsmAER, G. C. J. (1894) PRELImINARy NOTEs ON sOmE TETRAcTINELLIds OF ThE BAy OF NApLEs. Tijdschrift der Nederlandsche Dierkundige Fereeniging, 2, 269 - 286.","REdmONd, N. E., MORROW, C. C., ThAckER, R. W., DIAz, M. C., BOuRy-EsNAuLT, N., CaRdENAs, P., HAjdu, E., LobO-HAjdu, G., PIcTON, B. E., POmpONI, S. A., KAyAL, E. & COLLINs, A. G. (2013) PhyLOgENy ANd SysTEmATIcs OF DEmOspONgIAE IN LIghT OF NEW SmALL SubuNIT RIbOsOmAL DNA (18 S) SEquENcEs. Integrative and Comparative Biology, 53, 388 - 415.","JOhNsON, J. Y. (1863) DEscRIpTION OF A NEW sILIcEOus spONgE FROm ThE cOAsT OF MAdEIRA. Proceedings Zoological Society London, 3, 257 - 259."]}

GEODIA SIMPLEX SCHMIDT, 1870 Geodia simplex, Schmidt, 1870: p. 70; Arndt, 1913: p. 112; Burton, 1930: p. 490; 1946: p. 856. Type locality and deposition of holotype: Egedesminde, West Greenland, 50–90 m, ZMUC-DEM-319 (wet specimen). Burton (1946) also speaks of a Schmidt spicule preparation from the type, still in the BMNH collection today (BMNH 70.5.3.79). Discussion: Arndt (1913) identified with hesitation a small specimen from Norway as G. simplex; the specimen has not been located. Burton (1946), after an examination of spicule preparations from the type material from Greenland, concluded that G. simplex is probably identical to G. cydonium, which explains why Burton (1959) later mentioned G. cydonium as occurring in Greenland and Norway. Also, Koltun (1966: 57) doubted the existence of G. simplex as an independent species. We have inspected the holotype, a whole specimen cut in two. It is a rounded lump, measuring c. 7 cm in diameter and 3 cm in height; the surface is damaged in some areas, and algae are growing on it. The cortex is 1 mm thick. The spicule repertoire is clearly that of G. cydonium from the Mediterranean Sea. However, there must be a mistake, most likely from Schmidt’s side, as the label is in his handwriting. The algae growing on the specimen do not occur in Greenland; on the contrary, one of the species is Mediterranean, another one Mediterranean–southern boreal (Dr Poul Møller Pedersen, pers. comm.). We therefore confirm that G. simplex is a junior synonym of G. cydonium. As molecular results suggest that G. cydonium is a species complex (Cárdenas et al., 2011), only a thorough morphological revision of this complex will tell us to which species group G. simplex belongs., Published as part of Cárdenas, Paco, Rapp, Hans Tore, Klitgaard, Anne Birgitte, Best, Megan, Thollesson, Mikael & Tendal, Ole Secher, 2013, Taxonomy, biogeography and DNA barcodes of Geodia species (Porifera, Demospongiae, Tetractinellida) in the Atlantic boreo-arctic region, pp. 251-311 in Zoological Journal of the Linnean Society 169 (2) on page 298, DOI: 10.1111/zoj.12056, http://zenodo.org/record/5287119, {"references":["Schmidt O. 1870. Grundzuge einer Spongien-Fauna des atlantischen Gebietes. (Wilhelm Engelmann: Leipzig): iii - iv, 1 - 88, pls I - VI.","Arndt W. 1913. Zoologische Ergebnisse der ersten Lehr- Expedition der P. Scottlanderschen Jubilaums-Stifting. Jahresberichte der Schlesischen Gesellschaft fur vaterlandische Cultur 90: 110 - 119.","Burton M. 1930. Norwegian sponges from the Norman collection. Proceedings of the Zoological Society of London 1930: 487 - 546, pls I - II.","Burton M. 1946. Notes on certain species of Geodia described by Oscar Schmidt. Annals and Magazine of Natural History Ser. 11, xiii: 856 - 860.","Burton M. 1959. Spongia. In: Fridriksson A, Tuxen SL, eds. The zoology of Iceland. Copenhagen: Ejnar Munksgaard, 1 - 71.","Koltun VM. 1966. Four-rayed sponges of Northern and Far Eastern seas of the USSR (order Tetraxonida). Opredeliti Faunei SSSR 90. (Zoological Institute of the Academy of Sciences of the USSR: Moscow, Leningrad): 1 - 112, pls I - XXXVIII (in Russian).","Cardenas P, Xavier JR, Reveillaud J, Schander C, Rapp HT. 2011. Molecular phylogeny of the Astrophorida (Porifera, Demospongiae) reveals an unexpected high level of spicule homoplasy. PLoS ONE 6: e 18318."]}

GEODIA CYDONIUM (JAMESON, 1811) Records in the boreo-arctic region North-eastern Kara Sea: Off Sewernaja Semlja (Gorbunov, 1946: p. 37). South-western Barents Sea: Kola Fjord (Breitfuss, 1912: p. 62, as Cydonium mülleri; also referred to by Hentschel, 1929: p. 920, as Geodia mülleri). Norway: Off Vadsø, Varanger Fjord (Burton, 1930: p. 490, G. mülleri); Røberg, Trondheim Fjord (Arndt, 1913: p. 112, G. mülleri); Korsfjord near Bergen (Norman, 1879: p. 13, Geodia sp.; Brunchorst, 1891: p. 31, Geodia sp., according to Arndt, 1935: p. 30, both G. cydonium); off Haugesund (Schmidt, 1875: p. 120, Geodia gigas, according to Arndt, 1935: p. 30, G. cydonium); off Stavanger (Burton, 1930: p. 490, G. mülleri). Iceland: 64°56′N, 11°48′W, 216 m, 25.08.1902 (Burton, 1959: p. 9); Faxa Bay (Einarsson, 1941: p. 23, as G. mülleri). Discussion: Koltun (1966) reinvestigated the specimens of Gorbunov (1946) and Breitfuss (1912) and found that they are G. phlegraei. The specimen of Arndt (1913) could not be traced. A.B.K. and H.T.R. have sampled intensively on the same locality, Røberg in the Trondheimsfjord, and found many specimens of G. barretti and G. phlegraei, but not a single specimen referable to G. cydonium; note that the specimen of Arndt was probably about 15 cm in diameter. We conclude that Arndt’s specimen must have been misidentified. Arndt (1935) referred to some Geodia ‘sp.’s in the literature as G. cydonium. Nothing indicates that Arndt ever saw any of these specimens, rather he just felt certain that G. cydonium was an inhabitant of Norwegian waters. We have worked along most of the Norwegian coast and we have not found specimens that could be referred to G. cydonium. Probably all those referred to above represent G. barretti, which is very common along the entire Norwegian coastline. The Icelandic records are doubtful, too. We reinvestigated the specimen of Burton (1959) stored at ZMUC; in our opinion it is a fragment of G. barretti. Einarsson (1941) wrote ‘... enormous masses of sponges (G. mülleri ?) are encountered...’ Unfortunately we have no other samples from the same area, but everything considered, if it is a Geodia at all, it is presumably G. barretti. Our conclusion is that there are no certain records of G. cydonium north of the line Shetland Islands– Lousy Bank, west of the Faroe Islands. The last mentioned locality is listed by Burton (1959), and no description or further reference is given; accordingly, it may also be considered doubtful until a control is possible., Published as part of Cárdenas, Paco, Rapp, Hans Tore, Klitgaard, Anne Birgitte, Best, Megan, Thollesson, Mikael & Tendal, Ole Secher, 2013, Taxonomy, biogeography and DNA barcodes of Geodia species (Porifera, Demospongiae, Tetractinellida) in the Atlantic boreo-arctic region, pp. 251-311 in Zoological Journal of the Linnean Society 169 (2) on page 298, DOI: 10.1111/zoj.12056, http://zenodo.org/record/5287119, {"references":["Jameson R. 1811. Catalogue of animals of the class Vermes found in the Firth of Forth, and other parts of Scotland. Memoirs of the Wernerian Natural-History Society 1: 556 - 565.","Gorbunov GP. 1946. Benthonic populations of the novosibirsk shallow and central part of the Arctic Ocean. Tr. dreif. eksped. Glavsevmorputi na 1 / p ' G. Sedov' 1937 - 1940 III: 30 - 138 (in Russian).","Breitfuss JS. 1912. Zur Kenntnis der Spongio-Fauna des Kola Fjords. Travaux de la Societe Imperiale des Naturalistes de St. Petersbourg, Section Zoologie 41: 61 - 80, pls I - II.","Hentschel E. 1929. Die Kiesel- und Hornschwamme des Nordlichen Eismeers. In: Romer F, Schaudinn F, Brauer A, Arndt W, eds. Fauna Arctica. Eine Zusammenstellung der arktischen Tierformen mit besonderer Berucksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nordliche Eismeer im Jahre 1898. Jena: G. Fischer, 857 - 1042, pls XII - XIV.","Burton M. 1930. Norwegian sponges from the Norman collection. Proceedings of the Zoological Society of London 1930: 487 - 546, pls I - II.","Arndt W. 1913. Zoologische Ergebnisse der ersten Lehr- Expedition der P. Scottlanderschen Jubilaums-Stifting. Jahresberichte der Schlesischen Gesellschaft fur vaterlandische Cultur 90: 110 - 119.","Norman AM. 1879. The Mollusca of the fjords near Bergen, Norway. Journal of Conchology 2: 8 - 77.","Brunchorst J. 1891. Die biologische Meeresstation in Bergen. Bergens Museums Aarsberetning for 1890 5: 1 - 31.","Arndt W. 1935. Porifera. III. a Die Tierwelt der Nord- und Ostsee. Leipzig. 1 - 140.","Schmidt O. 1875. Spongien Die Expedition zur physikalischchemischen und biologischen Untersuchung der Nordsee im Sommer 1872. Jahresbericht der Commission zur Wissenschaftlichen Untersuchung der Deutschen Meere in Kiel. 115 - 120, pl. I.","Burton M. 1959. Spongia. In: Fridriksson A, Tuxen SL, eds. The zoology of Iceland. Copenhagen: Ejnar Munksgaard, 1 - 71.","Einarsson H. 1941. Survey of the benthonic animal communities of Faxa Bay (Iceland). Meddelelser fra Kommissionen for Danmarks fiskeri- og HavundersOgelser. Serie: Fiskeri 11: 1 - 46.","Koltun VM. 1966. Four-rayed sponges of Northern and Far Eastern seas of the USSR (order Tetraxonida). Opredeliti Faunei SSSR 90. (Zoological Institute of the Academy of Sciences of the USSR: Moscow, Leningrad): 1 - 112, pls I - XXXVIII (in Russian)."]}

Thenea muricata (Bowerbank, 1858) (Figures 18 E���F, 21���23, Table 5) Synonymy (modified from Maldonado (2002)). Tethea muricata Bowerbank, 1858: Bowerbank 1858, p. 308, pl. 25. fig. 18; Bowerbank 1872, p. 115, pl. V, figs. 1-6. Thenea muricata: Gray 1867, p. 541; Sollas 1888, p. 95; Topsent 1892, p. 37; Topsent 1894, p. 375; Lambe 1896, p. 202; Lambe 1900, p. 26; Topsent 1913 a; Stephens 1915, p. 11; Ferrer-Hern��ndez 1922, p. 2; Sar�� 1958, p. 216; Gamulin-Brida 1969, p. 89; Pulitzer-Finali 1972, p. 348; Uriz 1981, p. 48; Steenstrup & Tendal 1982, p. 259; Pulitzer-Finali 1983, p. 474; Pansini 1987 b, p. 43; Uriz & Rosell 1990, p. 48; Pansini & Musso 1991, Table 2; Boury-Esnault et al. 1994, p. 51; Maldonado 2002, p. 156; Voultsiadou 2005, Table 1; van Soest et al. 2007, Table 2; C��rdenas et al. 2011, Table S 1. Ancorina (Thenea) muricata: von Lendenfeld 1903, p. 53. ? Ancorina (Thenea) muricata: Babi�� 1914, p. 152. Wyvillethomsonia wallichii Wright, 1870: Wright 1870, p. 8. Thenea wallichii: Sollas 1882, p. 427. Dorvillia agariciformis Kent, 1870: Kent 1870, p. 293. Tisiphonia agariciformis: Thomson 1873, p. 167. Clavellomorpha minima Hansen, 1885: Hansen 1885, p. 19. ? Thenea (muricata) schmidti Sollas, 1886: Babi�� 1915, p. 389; Babi�� 1922, p. 282. Thenea intermedia Sollas, 1888: Sollas 1888, p. 97. Not Thenea schmidti Sollas, 1886: Sollas 1886, p. 183; Sollas 1888, p. 67 (this study) Material. ZMBN 85232, Brattholmen, Hjeltefjord, western Norway, 100 m; ZMBN 85252 (a/b), B��mlafjorden, Midtvik��y, western Norway, 59 �� 40 'N, 5 �� 24 'E, 230 - 90 m, triangular dredge; ZMBN 85231, west of Marstein, western Norway, 60 �� 8 ' 18 ''N, 4 �� 50 ' 47 ''E, 300 m, Agassiz trawl; ZMBN 87920, Sognefjorden deep basin, off L��nefjorden, Western Norway, 61 ��08.3'N, 6 �� 10 'E, 1232 - 1225 m, Agassiz trawl; ZMBN 85253, mid Norway, 63 ��08.54'N, 8 �� 13.37 'E, 165 m, M��re 2006 cruise; NTNU-VM 54908, Trollsteinen, mid Norway, 64 �� 35 ' 59.9 "N, 11 �� 1 ' 30 "E, 420 m; NTNU-VM 54909, Kinebbneset, Trondheimsfjorden western Norway, 63 �� 32 ' 28 "N, 9 �� 49 ' 47.2 "E; NTNU-VM 55655, Tautra, Trondheimsfjorden, mid Norway, 63 �� 32 ' 17.99 "N, 10 �� 30 ' 33.99 "E, 160 m; NTNU-VM 55622, north of Sm��la Island, mid Norway, 63 �� 37 ' 50 "N, 7 �� 59 ' 14.9 "E, 300 m; NTNU-VM 55843, Haltenbanken, mid Norway, 64 �� 14 ' 56 ''N, 9 ��0' 3.7 ''E. Comparative material. Thenea muricata, ZMAPOR 19481, Rockall Bank, 55 �� 31 ' 7.9 "N, 15 �� 48 ' 27.54 "W, 844-857 m; MNHN- DCL 3436, Banuyls, France, 800 m, Ecomarge 1985; MNHN-DCL 4083, off Cape Santa Maria di Leuca, Italy, 39 �� 34.01 'N, 18 �� 26.16 'E, 585 m, MEDECO 2007 cruise; ZMBN 85254, Gulf of Cadiz, Balgim campaign, St. CP25, 36�� 41 ' 5 ''N, 7 �� 19 ' 4 ''W, 544 m. Thenea schmidti, ZMAPOR 18036, Gulf of Cadiz, 35 �� 47 ' 59.9 "N, 7 �� 47 ' 59.9 "W, 443 m. Outer morphology (Fig. 21 A-B). Massive sub-spherical sponge. ZMBN 85231 is 2.8 cm long and 2.6 cm high. ZMBN 85252 (a/b) are made of ���fused specimens��� (4-8 cm of diameter). These larger specimens have many stringlike buds on its top surface, these can reach 2 cm long. ZMBN 85232 is a half damaged specimen. All specimens are slightly compressible. Surface is minutely hispid, like sand-paper. Color alive and in ethanol is brown. Choanosome is whitish to brownish (alive and in ethanol). Single, circular to ellipsoid oscule situated in the top (3 mm in ZMBN 85231, 1- 4 mm in ZMBN 85252); a conspicuous whitish sphincter surrounds it and there is no sieve. The sieved pore area runs along the equatorial side of the specimen (ZMBN 85231) or can extend itself in the entire lower part of the specimen (ZMBM 85252). All specimens have roots. Skeleton (Fig. 21 C-D). Like T. abyssorum. Depending on where the section is made, plesiasters are rare (especially in the choanosome) to very abundant (especially under the ectosome). Protriaenes are found under the ectosome as well (Fig. 21 D), they can be rare to common depending on the specimen. Anatriaenes were observed in the choanosome associated in bundles with triaenes (Fig. 21 D); they were also observed coming out of the top surface. Spicules (ZMBN 85231) (Figs. 18 E-F, 21 D-F). (a) oxeas, slightly bent, length: 4000- 5196.4 - 5925 ��m (N= 7); width: 30- 41.4 - 55 ��m (N= 7). (b) dichotriaenes (Fig. 21 D-E), rhabdome is straight to bent, or even flexuous, length: 1925- 2972 - 3875 ��m (N= 9); width: 23- 49.6 - 80 ��m; protoclad length: 190- 284.5 - 370 ��m; deuteroclad length: 350- 564 - 900 ��m. (c) anatriaenes (Fig. 21 F), common, straight, bent or flexuous rhabdome, length: 1550- 3976.1 - 4675 ��m (N= 23); rhabdome width: 5- 11.8 - 22 ��m; two shapes, the smaller ones, probably younger forms (with clad length: 60-85 ��m and rhabdome width: 5-9 ��m) have a regular umbrella-shape, the larger more common ones have bent clads, clad length: 60- 157.1 - 242 ��m. (d) protriaenes, rare, straight, rhabdome length> 1000 ��m; rhabdome width: 50 ��m (N= 1); clad length:> 600 ��m (N= 1). (e) spirasters (and few metasters) (Fig. 18 F), spined, length: 13- 23.2 - 32.5 ��m. (f) plesiasters (Fig. 18 E), 3-7 actines, faintly spined (hardly visible with an optical microscope), actine length: 16.6 - 37.0 - 70.3 actine width: 1.7- 3.5 - 10 Reproduction. We observed oocytes in vitellogenetic phase (A. Ereskovsky, pers. comm.) in the thick sections of T. schmidti (ZMAPOR 18036, collected at the end of August 2004) and T. muricata (ZMBN 85231, collected mid-June 2008). Spermatogenesis has been observed in T. muricata from Norway, possibly collected in May (Sollas 1882). Oocytes and spermatocytes have already been observed simultaneously in T. muricata from the Adriatic Sea indicating that this species can be hermaphrodite (Babi�� 1915). Elongated thread-like buds are common on the upper surface of large Norwegian specimens (e.g. ZMBN 85252) whereas they seem to be more subglobular in Mediterranean samples (Uriz 1981, Fig. 28). Distribution (Fig. 22). Greenland, Denmark Strait and southern Iceland (Steenstrup & Tendal 1982); Norway (Bowerbank 1872; Sollas 1882; Topsent 1913 a; Steenstrup & Tendal 1982); Rockall Bank (van Soest et al. 2007); northern Spain (Ferrer-Hern��ndez 1922); Azores (Topsent 1892; 1904); Gulf of Cadiz (Boury-Esnault et al. 1994); Mediterranean Sea (Sollas 1888; Sar�� 1958; Gamulin-Brida 1969; Pulitzer-Finali 1972; Uriz 1981; Pulitzer-Finali 1983; Uriz & Rosell 1990; Pansini & Musso 1991; Boury-Esnault et al. 1994; Voultsiadou, 2005);? Canada (Topsent 1892; Lambe 1896). As T. schmidti: Puerto Rico (Sollas 1888); Azores (Topsent 1892; 1904); Gulf of Cadiz (Sollas 1888; this study); Cape Verde (van Soest 1993); Mediterranean Sea (Babi�� 1915; 1922; Pansini 1987 b). Depth. For T. muricata: 60-2940 m (Steenstrup & Tendal 1982; Voultsiadou 2005); for T. schmidti in the Azores: 349���4020 m (Topsent 1904). Discussion. The Norwegian specimens fully agree with the redescription of this species (Steenstrup & Tendal 1982). It is the most widespread Thenea species (Fig. 22): its northernmost localities are at the moment in Eastern Greenland and the Denmark Strait while its southernmost localities are off Morocco. We have never identified T. muricata in northern Norway or the Barents Sea. We re-examined the T. muricata specimens (ZMAPOR 2385) collected by Vosmaer (1885) in the Barents Sea, and re-identified them as T. valdiviae based on the presence of sieves covering their oscules (vs. ���naked��� oscules with a conspicuous whitish sphincter in T. muricata). We therefore agree with Steenstrup and Tendal (1982) that other arctic records of T. muricata are likely to be T. valdiviae (e.g. Hansen 1885; Koltun 1966). T. muricata from the Zanzibar area is suspicious and probably a misidentification (Burton 1959 a). T. schmidti is distributed from the Azores to Gibraltar (Fig. 22) and originally characterized by i) a highdensity of larger plesiasters, ii) a poorly developed choanosome tissue, iii) larger choanocyte chambers and iv) a large oscule sphincter (Sollas, 1888). Our specimen of T. schmidti (Fig. 23) corresponds in every point to this description. Stephens (1915) compared her Irish T. muricata with T. schmidti and often could not tell which one had the most plesiasters. Indeed, Norwegian T. muricata could show abundant plesiasters but these were localized in certain areas (e.g. under the ectosome), whereas T. schmidti appeared uniformly filled with plesiasters (Fig. 23 C���D). The choanocyte chambers seemed a bit bigger (50-60 ��m) for T. schmidti, but not as much as the ones measured by Sollas (1888): up to 90 ��m. Mediterranean specimens of Thenea have raised doubts concerning the validity of T. schmidti. Sollas (1888) described T. intermedia with few plesiasters associated with poorly developed choanosome tissue and large choanocyte chambers (with measurements intermediate between T. schmidti and T. muricata from Norway). On the contrary, Topsent (1904) described a Mediterranean specimen with numerous plesiasters (as large as in T. schmidti) associated with dense choanosomal tissue. Therefore, these characters have been considered to be variable and T. schmidti was synonymized with T. muricata (Uriz 1981; Steenstrup & Tendal 1982). The Mediterranean specimen we examined is small (6 mm high, without the roots). It has numerous plesiasters, but not as large as in T. schmidti (Table 5). Genetic data (Fig. 25) show that our Mediterranean specimen has the same COI and 28 S sequence as T. muricata (ZMBN 85231) so Mediterranean specimens are likely to be T. muricata with numerous plesiasters. Meanwhile, even though T. schmidti (ZMAPOR 18036) has the exact same COI Folmer fragment as T. muricata, its faster evolving 28 S (C 1 ���D 2) sequence is significantly different (Fig. 25). This strongly suggests that T. schmidti is a valid NEA species and that we are not focusing on the right characters to differentiate it. The amount of plesiasters and the size of the choanocyte chambers do not seem to be reliable characters; the size of plesiasters does not seem to be a good character as they can be fairly large in T. muricata as well (Table 5). The very large plesiasters (actins of 420���540 ��m length) found by Babi�� (1915) in Thenea from the Adriatic Sea (Table 5) are puzzling at the moment. The white broad sphincter around the oscule of T. schmidti might be a more reliable character, but we need more specimens to confirm this. An even more southern species is Thenea bojeadori von Lendenfeld, 1907 (and its possible synonyms T. microclada von Lendenfeld, 1907 and T. megastrella von Lendenfeld, 1907) which can be found all along the northern coast of Africa (L��vi 1959; Cruz 2002) (Fig. 22). The status of T. bojeadori would also need to be reevaluated in order to clearly separate it from T. schmidti and T. muricata (Table 5). Material Depth Plesiaster Spiraster/metaster (m) (actin length) (length) ..... continued on the next page Material Depth Plesiaster Spiraster/metaster (m) (actin length) (length) MNHN-DCL 3436 * 800 31��� 126.9 ��� 247 18��� 25.2 ��� 39 Banyuls France (abundant) (10) Adriatic Sea - 420���540 21 (Babi�� 1915) (abundant), Published as part of C��rdenas, Paco & Rapp, Hans Tore, 2012, A review of Norwegian streptaster-bearing Astrophorida (Porifera: Demospongiae: Tetractinellida), new records and a new species, pp. 1-52 in Zootaxa 3253 on pages 39-43, DOI: 10.5281/zenodo.280590, {"references":["Bowerbank, J. S. (1858) On the Anatomy and Physiology of the Spongiadae. Part I. On the Spicula. Philosophical Transactions of the Royal Society of London, 148, 279 - 332, pls XXII - XXVI.","Maldonado, M. (2002) Family Pachastrellidae Carter, 1875. In: Hooper, J. N. A. & van Soest, R. W. M. (Eds.) Systema Porifera. A Guide to the classification of Sponges. Kluwer Academic / Plenum Publishers, New York, pp. 141 - 162.","Bowerbank, J. S. (1872) Contributions to a General History of the Spongiadae. Part I. Proceedings of the Zoological Society of London, 1872, 115 - 129, pls V - VI.","Gray, J. E. (1867) Notes on the arrangement of sponges, with the descriptions of some new genera. Proceedings of the Zoological Society of London, 2, 492 - 558, pls XXVII - XXVIII.","Sollas, W. J. (1888) Report on the Tetractinellida collected by H. M. S. Challenger, during the years 1873 - 1876. Report on the Scientific Results of the Voyage of H. M. S. Challenger, 1873 - 1876. Zoology, 25, 1 - 458, pls I - XLIV, 1 map.","Topsent, E. (1892) Contribution a l'etude des Spongiaires de l'Atlantique Nord (Golfe de Gascogne, Terre-Neuve, Acores). Resultats des campagnes scientifiques accomplies par le Prince Albert I. Monaco, 2, 1 - 165, pls I - XI.","Topsent, E. (1894) Etude monographique des spongiaires de France I. Tetractinellida. Archives de Zoologie Experimentale et Generale, 3, 259 - 400, pls. XI - XVI.","Lambe, L. M. (1896) Sponges from the Atlantic coast of Canada. Transactions of the Royal Society of Canada, 2, 181 - 211, pls I - III.","Lambe, L. M. (1900) Sponges from the coasts of Northeastern Canada and Greenland. Transactions of the Royal Society of Canada, VI, 19 - 48, pls I - VI.","Topsent, E. (1913 a) Spongiaires de l'Expedition Antarctique Nationale Ecossaise. Transactions of the Royal Society of Edinburgh, 49, 579 - 643, pls I - VI.","Stephens, J. (1915) Sponges of the Coasts of Ireland. I. - The Triaxonia and part of the Tetraxonida. Fisheries, Ireland Scientific Investigations, 1914, 1 - 43, pls I - V.","Ferrer-Hernandez, F. (1922) Mas datos para el conocimiento de las esponjas de las costas espanolas. Boletin de pescas, Sept. Oct. y Nov., 1 - 26, pl. 1 - 2.","Sara, M. (1958) Contributo all conoscenza dei Poriferi del Mar Ligure (1). Annali di Museo Civico di Storia Naturale Genova, 70, 207 - 244.","Gamulin-Brida, H. (1969) A contribution to the study of the tetractinellid sponge Thenea muricata with special consideration of its importance in the bionomics of the Adriatic Sea. Thalassia Jugoslavica, 5, 89 - 93.","Pulitzer-Finali, G. (1972) Report on a collection of sponges from the Bay of Naples. 1. Sclerospongiae, Lithistida, Tetractinellida, Epipolasida. Pubblicazioni della Stazione zoologica di Napoli, 38, 328 - 354.","Uriz, M. J. (1981) Estudio sistematico de las esponjas Astrophorida (Demospongia) de los fondos de pesca de Arrastre, entre Tossa y Calella (Cataluna). Boletin (Instituto Espanol de Oceanografia), 6, 8 - 58.","Steenstrup, E. & Tendal, O. S. (1982) The genus Thenea (Porifera, Demospongiae, Choristida) in the Norwegian Sea and adjacent waters; an annotated key. Sarsia, 67, 259 - 268.","Pulitzer-Finali, G. (1983) A collection of Mediterranean Demospongiae (Porifera) with, in appendix, a list of Demospongiae hitherto recorded from the Mediterranean Sea. Annali del Museo Civico di Storia Naturale Giacomo Doria, 84, 445 - 621.","Pansini, M. (1987 b) Report on a collection of Demospongiae from soft bottoms of the Eastern Adriatic Sea. Publications of the Sherkin Island Marine Station, 1, 41 - 43.","Uriz, M. - J. & Rosell, D. (1990) Sponges from bathyal depths (1000 - 1750 m) in the Western Mediterranean Sea. Journal of Natural History, 24, 373 - 391.","Pansini, M. & Musso, B. (1991) Sponges from Trawl-Exploitable Bottoms of Ligurian and Tyrrhenian Seas: Distribution and Ecology. Marine Ecology, 12, 317 - 329.","Boury-Esnault, N., Pansini, M. & Uriz, M. J. (1994) Spongiaires bathyaux de la mer d'Alboran et du golfe ibero-marocain. Memoires du Museum national d'Histoire naturelle, 160, 1 - 174.","Voultsiadou, E. (2005) Sponge diversity in the Aegean Sea: Check list and new information. Italian Journal of Zoology, 72, 53 - 64.","van Soest, R. W. M., Cleary, D. F. R., de Kluijver, M. J., Lavaleye, M. S. S., Maier, C. & van Duyl, F. C. (2007) Sponge diversity and community composition in Irish bathyal coral reefs. Contributions to Zoology, 76, 121 - 142.","Cardenas, P., Xavier, J. R., Reveillaud, J., Schander, C. & Rapp, H. T. (2011) Molecular phylogeny of the Astrophorida (Porifera, Demospongiae p) reveals an unexpected high level of spicule homoplasy. PLoS ONE, 6, e 18318. doi: 10.1371 / journal. pone. 0018318.","von Lendenfeld, R. (1903) Porifera. Tetraxonia. In: Schulze, F. E. (Ed.) Das Tierreich., Friedlander: Berlin, pp. vi - xv, 1 - 168.","Babic, K. (1914) Uber Ancorina (Thenea) muricata (Bowerbank). Zoologischen Anzeiger, XLV, 152 - 158.","Wright, P. (1870) Notes on Sponges. 1, On Hyalonema mirabilis, Gray. 2, Aphrocallistes Bocagei sp. nov. 3, On a new Genus and Species of Deep Sea Sponge. Quarterly Journal of Microscopical Science, 10, 1 - 9, pls. I - III.","Sollas, W. J. (1882) The sponge-fauna of Norway; a Report on the Rev. A. M. Norman's Collection of Sponges from the Norwegian Coast. Annals and Magazine of Natural History, 5, 426 - 453, pl XVII.","Kent, W. S. (1870) On a New Anchoring Sponge, Dorvillia agariciformis. Monthly Microscopical Journal, 4, 293 - 295, pl. LXVI.","Thomson, C. W. (1873) The Depths of the Sea. Macmillan and Co., London, 527 pp.","Hansen, G. A. (1885) Spongiadae. The Norwegian North-Atlantic Expedition 1876 - 1878. Zoologi, 13, 1 - 26, pls I - VII, 1 map.","Sollas, W. J. (1886) Preliminary account of the Tetractinellid sponges Dredged by H. M. S. ' Challenger' 1872 - 76. Part I. The Choristida. Scientific Proceedings of the Royal Dublin Society (new series), 5, 177 - 199.","Babic, K. (1915) Zur Kenntnis der Theneen. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere, 40, 389 - 408, pl. 16 - 18.","Babic, K. (1922) Monactinellida und Tetractinellida des Adriatischen Meeres. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere, 46, 217 - 302, pls 8 - 9.","Topsent, E. (1904) Spongiaires des Acores. Resultats des campagnes scientifiques accomplies par le Prince Albert I. Monaco, 25, 1 - 280, pls 1 - 18.","van Soest, R. W. M. (1993) Affinities of the marine Demosponge fauna of the Cape Verde Islands and tropical West Africa. Courier Forschungsinstitut Senckenberg, 159, 205 - 219.","Vosmaer, G. C. J. (1885) The Sponges of the ' Willem Barents' Expedition 1880 and 1881. Bijdragen tot de Dierkunde, 12, 1 - 47, pls I - V.","Koltun, V. M. (1966) Four-rayed sponges of Northern and Far Eastern seas of the USSR (order Tetraxonida). Opredeliti Faunei SSSR 90. (Zoological Institute of the Academy of Sciences of the USSR: Moscow, Leningrad), 1 - 112, pls I - XXXVIII.","Burton, M. (1959 a) Sponges. In: Scientific Reports. John Murray Expedition, 1933 - 34. British Museum (Natural History): London, 151 - 281.","von Lendenfeld, R. (1907) Die Tetraxonia. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf der Dampfer Valdivia 1898 - 1899, 11, i - iv, 59 - 374, pls IX - XLVI.","Levi, C. (1959) Resultats scientifiques des Campagnes de la ' Calypso'. Campagne de la ' Calypso' dans le Golfe de Guinee et aux iles Principe, Sao Tome et Annobon. 5. Spongiaires. Annales de l'Institut Oceanographique, 37, 115 - 141, pls 5 - 6.","Cruz, T. (2002) Esponjas marinas de Canarias. Consejeria de Politica Territorial y Medio Ambiente del Gobierno de Canarias, S / C Tenerife, 260 pp."]}

Family Theneidae Carter, 1883 Diagnosis: Astrophorida with long-shafted triaenes (sometimes lost) in combination with diverse categories of streptasters: spirasters, metasters and plesiasters (sometimes with annulate actines) (C��rdenas et al. 2011). Genera: Annullastrella, Cladothenea, Thenea., Published as part of C��rdenas, Paco & Rapp, Hans Tore, 2012, A review of Norwegian streptaster-bearing Astrophorida (Porifera: Demospongiae: Tetractinellida), new records and a new species, pp. 1-52 in Zootaxa 3253 on page 33, DOI: 10.5281/zenodo.280590, {"references":["Carter, H. J. (1883) Contributions to our Knowledge of the Spongida. - Pachytragida. Annals and Magazine of Natural History, 5, 344 - 369, pls XIV - XV.","Cardenas, P., Xavier, J. R., Reveillaud, J., Schander, C. & Rapp, H. T. (2011) Molecular phylogeny of the Astrophorida (Porifera, Demospongiae p) reveals an unexpected high level of spicule homoplasy. PLoS ONE, 6, e 18318. doi: 10.1371 / journal. pone. 0018318."]}

Genus Pachastrella Schmidt, 1868 Diagnosis: Pachastrellidae with large four rayed clathrops (never becoming mesotrider desmas). Microscleres always include microstrongyles., Published as part of C��rdenas, Paco & Rapp, Hans Tore, 2012, A review of Norwegian streptaster-bearing Astrophorida (Porifera: Demospongiae: Tetractinellida), new records and a new species, pp. 1-52 in Zootaxa 3253 on page 11, DOI: 10.5281/zenodo.280590, {"references":["Schmidt, O. (1868) Die Spongien der Kuste von Algier. Mit Nachtragen zu den Spongien des Adriatischen Meeres (Drittes Supplement). Wilhelm Engelmann, Leipzig, i - iv, 1 - 44, pls I - V. pp."]}

Thenea abyssorum Koltun, 1964 (Figures 17, 18 A���B) Synonymy. Thenea muricata abyssorum Koltun, 1959 (nomen nudum): Koltun 1959, p. 662. Thenea abyssorum: Koltun 1964, p. 146; Koltun 1966, p. 37; Steenstrup & Tendal 1982, p. 259; Barthel & Tendal 1993, p. 83; Witte 1996, p. 571; Weslawski et al. 2003, p. 75; C��rdenas et al. 2011, Table S 1. Thenea sp.: Babi�� 1915, p. 408. Material. NTNU-VM 54948, off Vester��len, northern Norway, 69 �� 30.3 'N, 13 �� 55 'E, 2130 m; NTNU-VM 66585, R��stbanken, northern Norway, 68 �� 36.5 'N, 11 �� 54 'E, 675- 850 m. Comparative material. Thenea abyssorum, ZMBN 85228, Arctic mid-ocean ridge, Greenland Sea, 73 �� 34 'N, 07�� 45 'E, H 2 DEEP cruise 2008, 2425��� 2463 m, Sneli sled (Sneli 1998). Outer morphology (Fig. 17 A). Small sub-spherical sponge (2.5 cm high, 2 cm wide) with root-like structures. Very hispid. Slightly compressible. Color alive and in ethanol is dirty brown. Choanosome is whitish to brownish (alive and in ethanol). Pores and oscule are on opposite sides (in upper part of sponge). The pore area is covered by a sieve and protected by an overhang. There is no sieve over the single oscule (ca 1 mm in diameter), it is circular and surrounded by a fringe of long oxeas. Skeleton (Fig. 17 B���C). Radial bundles of oxeas start at the center of the specimen. Many go beyond the surface, especially on the top surface, very hispid. Triaenes are found only at the surface (Fig. 17 C), either crossing the ectosome or supporting it. Spirasters are abundant in the ectosome, but spirasters and plesiasters can be found throughout the choanosome (spirasters being much more common than plesisasters). The roots are made of bundles of oxeas and anatriaenes and they originate in the central part of the sponge. Anatriaenes are especially common in the basal part, always crossing the ectosome. Spicules (ZMBN 85228) (Figs. 17 C���F, 18 A���B). (a) oxeas, straight, length> 3750 ��m; width: 20���40 ��m. (b) dichotriaenes (Fig. 17 C-D), rarely modified to orthotriaene or plagiotriaene, straight rhabdome or slightly bent, rhabdome length> 4165 ��m; rhabdome width: 20- 47.2 - 70 ��m; protoclad length: 60- 132.4 - 210 ��m; deuteroclad length: 240- 480.5 - 850 ��m. (c) anatriaenes I (Fig. 17 E), common, straight rhabdome with a rather blunt tip, rhabdome length: 920-1257.5 - 1750 ��m (N= 22); rhabdome width: 5 - 9.0 - 15 ��m; clad length: 45- 87.8 - 130 ��m. A more or less developed swelling can be found above the cladome. (d) anatriaenes II (Fig. 17 F), common, very long and flexuous rhabdome, length between 2000 ��m to more than 3600 ��m; rhabdome width: 10 ��m; very long clades running somewhat parallel to the rhabdome, giving to the anatriaene a closed-umbrella shape, in some cases the cladome is aborted and only an inflated tip is present, clad length: 40- 279.7 - 548 ��m (N= 16). (e) protriaenes, absent in this specimen. (e) spirasters (Fig. 18 B), strongly spined, length: 25.0- 36.6 -57.3 ��m. (f) plesiasters (Fig. 18 A), 4��� 6 actines, strongly spined or more rarely smooth, actine length: 60- 84.3 - 100 ��m; actine width: 13���16 ��m. Reproduction. This species has been shown to be oviparous and gonochoristic, but successive hermaphroditism cannot be excluded (Witte 1996). Oogenesis takes place between June and August with spermatic cysts found only in August (Witte, 1996). Subglobular buds are relatively common in this species, they are often placed at the lower half of the specimens (Barthel & Tendal 1993; Witte 1996). Distribution. Arctic Ocean, Norwegian Sea, Greenland Sea (Steenstrup & Tendal 1982, Fig. 1 D). Depth. 850���3670 m (this study; Steenstrup & Tendal 1982). Discussion. T. abyssorum is the northernmost and deepest (> 1000 m) Thenea species of the North Atlantic. On the field, this species can be easily confused with T. muricata or small T. valdiviae: a spicule preparation is therefore necessary. T. abyssorum is characterized by its large strongly spined plesiasters and streptasters. We report here for the first time a second category of anatriaene in this species, with unusually long clads (Fig. 17 F)., Published as part of C��rdenas, Paco & Rapp, Hans Tore, 2012, A review of Norwegian streptaster-bearing Astrophorida (Porifera: Demospongiae: Tetractinellida), new records and a new species, pp. 1-52 in Zootaxa 3253 on pages 33-36, DOI: 10.5281/zenodo.280590, {"references":["Koltun, V. M. (1964) Sponges (Porifera) collected in the Greenland seas and from the region to the north of the Spitzbergen and Franz Josef Land, from expeditions of the ' F. Litke' 1955, ' Obb' 1956 and ' Lena' 1957 and 1958. Scientific results of the high- latitudes Oceanographic Expeditions to the northern part of the Greenland Sea and adjacent areas of the Arctic basin between 1955 - 1958. Trudy Arkticheskogo i antarkticheskogo Nauchno-Issledovatel'skogo Instituta, 259, 143 - 166.","Koltun, V. M. (1959) Donnaia fauna abissalnuich glubin Zhentralnovo Poljarnovo basseina [Fauna from the abyssal of central polar basin]. Doklady Akademii Nauk SSSR. Seriya: Biologiya, 129, 662 - 665.","Koltun, V. M. (1966) Four-rayed sponges of Northern and Far Eastern seas of the USSR (order Tetraxonida). Opredeliti Faunei SSSR 90. (Zoological Institute of the Academy of Sciences of the USSR: Moscow, Leningrad), 1 - 112, pls I - XXXVIII.","Steenstrup, E. & Tendal, O. S. (1982) The genus Thenea (Porifera, Demospongiae, Choristida) in the Norwegian Sea and adjacent waters; an annotated key. Sarsia, 67, 259 - 268.","Barthel, D. & Tendal, O. S. (1993) The sponge association of the abyssal Norwegian-Greenland sea: species composition, substrate relationships and distribution. Sarsia, 78, 83 - 96.","Witte, U. (1996) Seasonal reproduction in deep-sea sponges-triggered by vertical particle flux? Marine Biology, 124, 571 - 581.","Weslawski, J. M., Wlodarska-Kowalczuk, M. & Legezynska, J. (2003) Occurrence of soft bottom macrofauna along the depth gradient in High Arctic, 79 ° N. Polish Polar Research, 24, 73 - 88.","Cardenas, P., Xavier, J. R., Reveillaud, J., Schander, C. & Rapp, H. T. (2011) Molecular phylogeny of the Astrophorida (Porifera, Demospongiae p) reveals an unexpected high level of spicule homoplasy. PLoS ONE, 6, e 18318. doi: 10.1371 / journal. pone. 0018318.","Babic, K. (1915) Zur Kenntnis der Theneen. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere, 40, 389 - 408, pl. 16 - 18.","Sneli, J-A. (1998) A simple benthic sledge for shallow and deep-sea sampling. Sarsia, 83, 69 - 72."]}

Ecionemia megastylifera Wintermann-Kilian & Kilian, 1984 (Figures 5–6 – 7) Synonyms. Ecionemia megastylifera Wintermann-Kilian & Kilian, 1984: 122, figs. 1–2. Ancorina megastylifera (Wintermann– Kilian & Kilian, 1984): Alcolado 2002: p. 56. Stellettinopsis dominicana Pulitzer-Finali, 1986: 67, figs. 1–2 (new synonym). Ecionemia dominicana (Pulitzer-Finali, 1986): Rützler et al. 2000: Table 1 (new synonym). Holotype. INVPOR 1148, Bahia Nenguange, Colombia, 3 m depth. Material. 5 specimens, from STRI Point, Solarte Island and Bastimento Island, 0.4–20 m depth. Two specimens deposited: ZMBN 81782–81783. Additional material examined. Ecionemia megastylifera, INVPOR 1148, holotype, Bahia Nenguange, Colombia, 3 m; ZMAPOR 7772, Bahamas; ZMAPOR 14265, off Slangebaai, Curacao, 3.8 m. Ecionemia dominicana (Pulitzer-Finali, 1986), MSNG 47679, holotype, Boca Chica, Dominican Republic, 15– 25 m. Ecionemia demera (de Laubenfels, 1934), MNHN DNBE- 17, slide of holotype, Puerto Rico, 60–73 m; ZMAPOR 0 3512, Puerto Rico. Outer morphology (Fig. 5 A–D). Thickly encrusting to massive lobate sponge. One very large specimen found on coral rubble in Solarte south was sub-circular, 8 cm high and with a diameter of 15 cm (Fig. 5 B); it had cemented and integrated coral rubble pieces. Other specimens had an elongated shape and were 20 cm long (Fig. 5 C). Surface is rugose. Slightly compressible. Choanosome has a hairy aspect due to the high density of large oxeas. External color on living specimen and in ethanol is light brownish to black. Internal color on living specimen is whitish to cream, whitish in ethanol. Oscules are 3–10 mm of diameter and distributed on the top surface of the sponge (Fig. 5 A–C). There is a thin contractile membrane around the oscule. Oscules lead into one or more cloacae chambers where uniporal exhalant openings are present. Pores are cribriporal (Fig. 5 D) and evenly distributed over large surfaces of the sponge, there is no separation between pore plates. A single pore plate has a diameter of around 0.5 mm. Macro-epibionts (e.g. sponges, algae) can be found growing on large specimens. Gammarid amphipods were found living in an oscule. Skeleton (Fig. 5 E). Small acanthostrongylasters lay on a paratangential layer of acanthomicrorhabds, thus forming a very thin cortex (ca 140–170 µm thick) supported by bundles of long–shafted robust dichotriaenes. Thin microxeas, sometimes in bundles, cross the cortex. In certain parts where the sponge is hispid, large oxeas, dichotriaenes, anatriaenes and protriaenes also cross the cortex. Sub-cortical canals cross transversely this layer of triaenes. Under the triaenes, large oxeas paratangentially positioned form a dense layer (ca 1.2 mm thick). Deeper, this arrangement becomes looser and more irregular; oxeas II, some dichotriaenes and rare anatriaenes can be found there also. Acanthoxyasters are found dispersed in the choanosome, and are abundant around canals. Large granular cells (diameter: 16–27 µm) are especially abundant in the cortex but are also present, albeit in lower density, in the choanosome. The cortex of cloacae chambers is somewhat different. It has fewer acanthomicrorhabds and is not supported by triaenes; it is mainly tangential large oxeas that stretch and support the acanthostrongylaster layer. It is only crossed by a few microxeas. Spicules (measurements of ZMBN 81782, except for protriaenes, only found in ZMBN 81783) (Figs. 6– 7). Megascleres: (a) oxeas I, very large, slightly bent, sometimes modified to styles, length: 1332– 1524 – 1764 µm; width: 56– 64.9 – 70 µm. (b) oxeas II, straight or slightly bent, length: 520– 815.2 – 1260 µm; width: 14– 22.9 – 42 µm. (c) microxeas, usually straight but can be slightly bent, length: 182– 215.4 – 317 µm; width: 2.5– 4 – 5 µm. (d) dichotriaenes to orthotriaenes, stout, rare in some specimens, the rhabdome can be slightly bent, and in some rare cases extra spines are present, thus becoming acanthotriaenes, rhabdomes can have an oxeote or strongylote end, rhabdome length: 790– 997.6 – 1222 µm; width: 50– 64.7 – 80 µm; clad length: 120– 151.7 – 180 µm (some young triaenes observed, rhabdome length: 400– 530 – 651 µm; width: 18– 27 – 41 µm; clad length: 53– 76.2 – 104 µm). (e) anatriaenes, rare, rhabdome slightly inflated, rhabdome length: 228– 278.3 – 334 µm (N= 3); rhabdome width: 1– 1.3 – 2 µm (N= 6); clad length: 3– 4 – 5 µm (N= 6). (f) protriaenes, rare, sometimes modified to mesoprotriaenes, rhabdome length: 417 µm (N= 1); rhabdome width: 2.5– 3.75 – 5 µm (N= 2); clad length: 27– 32.5 – 38 µm (N= 2). Microscleres: (g) acanthomicrorhabds (Fig. 6 A), with oxeote or strongylote ends, sometimes centrotylote, length: 59– 98.6 – 138.3 µm; width: 4.5– 8.2 – 10.6 µm. (h) acanthoxyasters (Fig. 6 B), 6–9 actines, a characteristic spike at the end of each actine, diameter: 9– 13.1 – 19 µm. (i) acanthostrongylasters (Fig. 6 C), 4–11 actines, diameter: 4– 6.7 – 10 µm. Habitat in the Bocas del Toro region. Common in coral rubble or in sheltered areas of reefs (such as small cavities). Usually sciophilous 1–25 m depth. Distribution. Bahamas (this study); Cuba (Alcolado 2002); Dominican Republic (Pulitzer-Finali 1986); Belize (Rützler et al. 2000); Costa Rica (Cortés 1996); Panama (this study); Colombia (Wintermann-Kilian & Kilian 1984; Valderrama 2001; Díaz 2007); Curaçao (this study). Remarks and discussion. This species possesses true spiny microrhabds (Fig. 6 A) (not sanidasters) so it does not belong to the Ancorina genus. The original description by Wintermann– Kilian and Kilian (1984) mentions only one size class of euasters, but after reexamination of the holotype it is quite obvious that there are two kinds of euasters: the small strongylasters placed in the ectocortex and the larger oxyasters in the choanosome. Wintermann– Kilian and Kilian (1984) note the presence of microstrongyles, but these are simply variations of microrhabds. They also overlooked the presence of microxeas present in the holotype (Fig. 6 E). Furthermore, we observed in the Bocas material and the holotype two size classes of oxeas that had not been identified before. There were some differences between the Bocas del Toro specimens and the holotype. The holotype had smaller oxeas, smaller triaenes (Fig. 6 D), thinner and smaller microrhabds (Fig. 6 E) (length: 48– 69.2 – 92, width: 2– 3.1 – 5), smaller microxeas, asters with thinner actines (Fig. 6 F), and more oxeas II (most of them found under the oxea I layer) (Fig. 5 F). Anatriaenes and protriaenes observed in the Bocas del Toro specimens were not found in the holotype. The holotype of E. dominicana had also two size classes of euasters and the microxeas, both of which had been overlooked by Pulitzer-Finali (1986). The microrhabds’ measurements in the E. dominicana holotype were very similar to those found in the holotype of E. megastylifera. As a result of these observations, E. dominicana becomes a junior synonym of E. megastylifera. Lehnert (1993) also described a Stelletinopsis sp. in Cozumel (Mexico) which resembles E. megastylifera. However, the vase shape, the blue and yellow colors and the absence of triaenes do not match with our species. The other Caribbean species of Ecionemia, E. demera from Puerto Rico differs from E. megastylifera in having plagiotriaenes, much larger anatriaenes, smaller acanthomicrorhabds (ca 7 µm) and only one kind of euasters. The dark color of E. megastylifera is due to large pigmented cells, more or less present with respect to light conditions. These pigments may serve for light protection since their abundance is greater in light environments; this has been suggested in other sponges (Renouf 1934; Bandaranayake et al. 1996; Cavalcanti et al. 2007; Valderrama et al. 2009). This is the species collected in Bocas del Toro by Nichols (2005) and identified as Ecionemia sp. We sequenced three of our specimens and obtained strictly identical COI sequences as Nichols (2005).

Cinachyrella apion (Uliczka, 1929) (Figure 3) Synonyms (R��tzler & Smith 1992). Cinachyra apion Uliczka, 1929: 43, figs. 16���21, pl. I, fig. 4. Cinachyra rizophyta Uliczka, 1929: 38, figs. 1���10, pl. I, fig. 1. Cinachyra cavernosa Lamarck, 1815 sensu de Laubenfels, 1950: 128, fig. 56, pl. II, fig. 7. Cinachyra subterranea van Soest & Sass, 1981: 337, fig. 4, pl. II, fig. 2. Cinachyrella apion (Uliczka, 1929): R��tzler 1987: 200, figs. 2g; 3 b���g; 4 a, c; 5 d, e. Holotype. ZMB 4911, St. Thomas, Virgin Islands (not seen). Material. ZMBN 80958, Adriana���s reef, 1 m depth; ZMBN 81785, STRI Point, 1 m depth; 4 other specimens collected. Additional material examined. Cinachyrella apion, ZMBN 81789, Key Largo, Florida; UFBA-POR 2232, Barra/ Ondina, Salvador, Bahia, Brazil, 13 ��00' 42 '' S / 38 �� 31 ' 12 '' W, C. Menegola col., intertidal. Outer morphology (Fig. 3 A). Massive, small spherical sponge that can reach 7 cm in diameter. External color alive is orange, as well as the choanosome. In ethanol, color becomes yellowish. The surface is very hispid, which hides the true color of the sponge, and gives it a rather sandy color. Generally one oscule (ca 3 mm of diameter), which can be found in any part of the sponge. Porocalices (2 mm of diameter) are numerous and evenly distributed, especially on the sides. No macro-epibionts were found growing on this species, except filamentous red algae. No budding was observed. Skeleton (Fig. 3 B). The skeleton organization is similar to that of C. alloclada except for the presence of trichodragmata. Radial bundles of oxeas cross the choanosome until and beyond the surface. A whitish ectosome layer is visible with the naked eye (ca 0.9 mm), it has less sigmas than in the choanosome. Anatriaenes are tangled with the oxea bundles. The cladomes of anatriaenes are placed in or just under the ectosome layer, or protruding beyond the surface. Protriaenes I and protriaenes II are present only around the oscules and porocalices (Fig. 3 C). Protriaene cladomes were not found in the sponge, but only at its surface. Trichodragmata were common in the choanosome (never in the ectosome), with no particular orientation. Smaller stout oxeas, Haplosclerida-like, dispersed in the choanosome with no particular orientation. A few crystalline round structures (diameter of 91���127 ��m) are present in the choanosome, only visible in thick sections. In the choanosome there are also some foreign spicules, foraminifera and diatoms. Spicules (measurements of ZMBN 80958) (Fig. 3 D���F). Megascleres: (a) oxeas I, large, length: 2088��� 3526.2 ��� 4800 ��m (N= 14); width: 11��� 31.3 ��� 37 ��m (N= 14). (b) oxeas II (Fig. 3 D), foreign?, short, stout, length: 141��� 169.9 ��� 219 ��m; width: 6��� 9.4 ��� 13.6 ��m. (c) protriaenes I, rhabdome length:> 2040 ��m (rhabdomes broken); rhabdome width: 4��� 5.2 ��� 7 ��m; clad length: 33��� 74 ��� 113 ��m. (d) protriaenes II, rhabdome length:> 1440 (rhabdomes broken); rhabdome width: 2��� 2.9 ��� 4 ��m; clad length: 8��� 20.7 ��� 29 ��m. (e) anatriaenes (Fig. 3 E), very common, rhabdome with whip���like end, rhabdome length: 2232��� 3708 ��� 4800 ��m (N= 4); rhabdome width: 2��� 4.1 ��� 7 ��m; clad length: 11��� 44.7 ��� 68 ��m. Microscleres: (f) trichodragmata, length: 135��� 203.3 ��� 248 ��m; width: 7��� 14.1 ��� 30 ��m. (g) sigmaspires (Fig. 3 F), spiny, length: 8.7��� 10.4 ��� 12.5 ��m (N= 13); width: 0.9��� 1.2 ��� 1.4 ��m (N= 13). Habitat in the Bocas del Toro region. Common on very shallow reefs and sand (0.4���1.5 m depth), often below mangrove trees. Other reports at Bocas del Toro found it growing on mangrove roots (Collin et al. 2005; D��az 2005), as it is often found in Belize (R��tzler & Smith 1992). Distribution. North and South Carolina (R��tzler & Smith 1992); Bermuda (de Laubenfels 1950; R��tzler & Smith 1992); Florida (R��tzler & Smith 1992); Bahamas (van Soest & Sass 1981; R��tzler & Smith 1992); Virgin Islands (Uliczka 1929); Belize (R��tzler & Smith 1992; R��tzler et al. 2000); Panama (Wulff 2000; D��az 2005); Colombia (Wintermann-Kilian & Kilian 1984); Brazil (Lazoski et al. 1999; Cedro et al. 2007). Remarks and discussion. C. apion was considered to be rare in Bocas del Toro (D��az 2005), but we found it to be fairly common, especially in Adriana���s reef. Despite the presence of oxeas II (Fig. 3 D), the morphology of our specimens completely agreed with the description of C. apion. It is interesting to note that the short oxeas II were never previously reported in C. apion, and that they were not observed in our specimens from Florida. Therefore, we cannot rule out the possibility that these oxeas II were foreign. The crystalline structures in the choanosome were previously reported in this species and C. alloclada (R��tzler & Smith 1992; Lazoski et al. 1999). These structures have also been observed in other species of Tetillidae: Cinachyrella levantinensis Vacelet et al. (Vacelet et al. 2007), Craniella cranium (M��ller) and Craniella zetlandica (Carter) (P. C��rdenas, personal observation). Their function is still unknown, as well as their diagnostic relevance. We nonetheless emphasize the fact that these were never found in C. kuekenthali (in previous records or our specimens). Our COI sequence (FJ 711645) was strictly identical to those of a C. apion from Twin Cays Mangroves, Belize (EF 519601) and from Flatts Inlet, Bermuda (AJ 843895)., Published as part of C��rdenas, Paco, Menegola, Carla, Rapp, Hans Tore & D��az, Maria Cristina, 2009, Morphological description and DNA barcodes of shallow-water Tetractinellida (Porifera: Demospongiae) from Bocas del Toro, Panama, with description of a new species, pp. 1-39 in Zootaxa 2276 on pages 7-9, DOI: 10.5281/zenodo.191088, {"references":["Uliczka, E. (1929) Die tetraxonen Schwamme Westindiens (auf Grund der Ergebnisse der Reise Kukenthal-Hartmeyer). Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Thiere, supplement 16, 35 - 62, pl I.","Rutzler, K. & Smith, K. P. (1992) Guide to the Western Atlantic species of Cinachyrella (Porifera: Tetillidae). Proceedings of the Biological Society of Washington, 105, 148 - 164.","Lamarck, J. B. P. (1815) Suite des polypiers empates. Memoires du Museum d'Histoire Naturelle, Paris, 1, 69 - 80, 162 - 168, 331 - 340.","de Laubenfels, M. W. (1950) The Porifera of the Bermuda Archipelago. Transactions of the Zoological Society of London, 27, 1 - 154, pls I - II.","van Soest, R. W. M. & Sass, D. (1981) Marine sponges from an island cave on San Salvador Island, Bahamas. Bijdragen tot de Dierkunde, 51, 332 - 344.","Rutzler, K. (1987) Tetillidae (Spirophorida, Porifera): a taxonomic reevaluation. In: Vacelet, J. & Boury-Esnault, N. (Eds.) Taxonomy of Porifera. NATO ASI Series, vol. G 13. Springer-Verlag, Berlin, Heidelberg, pp. 187 - 203.","Collin, R., Diaz, M. C., Norenburg, J., Rocha, R. M., Sanchez, J. A., Schulze, A., Schwartz, M. & Valdes, A. (2005) Photographic identification guide to some common marine invertebrates of Bocas Del Toro, Panama. Caribbean Journal of Science, 41, 638 - 707.","Rutzler, K., Diaz, M. C., van Soest, R. W. M., Zea, S., Smith, K. P., Alvarez, B. & Wulff, J. (2000) Diversity of sponge fauna in mangrove ponds, Pelican Cays, Belize. Atoll Research Bulletin, 476,","Wintermann-Kilian, G. & Kilian, E. F. (1984) Marine sponges of the region of Santa Marta (Colombia). Part II. Homoslerophorida, Choristida, Spirophorida, Hadromerida, Axinellida, Halichondrida, Poecilosclerida. Studies on Neotropical Fauna and Environment, 19, 121 - 135.","Lazoski, C., Peixinho, S., Russo, C. A. M. & Sole Cava, A. M. (1999) Genetic confirmation of the specific status of two sponges of the genus Cinachyrella (Porifera: Demospongiae: Spirophorida) in the Southwest Atlantic. Memoirs of the Queensland Museum, 44, 299 - 305.","Cedro, V. R., Hajdu, E., Sovierzosky, H. H. & Correia, M. D. (2007) Demospongiae (Porifera) of the shallow coral reefs of Maceio, Alagoas State, Brazil. In: Custodio, M. R., Lobo-Hajdu, G., Hajdu, E. & Muricy, G. (Eds.). Porifera research: biodiversity, innovation and sustainability. Serie Livros 28, Museu Nacional, 233 - 237.","Vacelet, J., Bitar, G., Carteron, S., Zibrowius, H. & Perez, T. (2007) Five new sponge species (Porifera: Demospongiae) of subtropical or tropical affinities from the coast of Lebanon (eastern Mediterranean). Journal of the Marine Biological Association of the UK, 87, 1539 - 1552."]}

Geodia gibberosa Lamarck, 1815 (Figures 14���15) Synonyms (modified from da Silva, 2002). Geodia gibberosa Lamarck, 1815: 334. Pyxitis gibberosa Lamarck, 1815: Schmidt 1870: 70. Geodia (Geodia) gibberosa Lamarck, 1815: Hechtel 1965: 68, pl. VIII, fig. 2. Geodia cariboea Duchassaing de Fonbressin and Michelotti, 1864 (in part): 105, pl. XXV, fig. 8. Geodia tumulosa Bowerbank, 1872: 628, pl. XLVII. Geodia media Bowerbank, 1873 (non G. m e d ia Lendenfeld, 1910): 13, pl. II. Geodia dysoni Bowerbank, 1873: 14, pl. III. Geodia reticulata Bowerbank, 1874: 300, pl. XLVI, figs. 14���20. Sidonops stromatodes Uliczka, 1929: 54, figs. 51���56, pl. I, fig. 10. Geodia media var. leptoraphes Uliczka, 1929: 56, figs. 57���67, pl. I, fig. 11. Geodia flexisclera Pulitzer-Finali, 1986: 76, figs. 10���11. Holotype. MNHN DT ��� 608, dry, French Guiana. Material. 5 specimens, all collected in Solarte lagoon, on mangrove roots, 0.5���1 m depth. Two specimens (fragments) are deposited: ZMBN 77928 and 81780. Additional material examined. Geodia gibberosa, MNHN DT ��� 608, holotype, French Guiana; YPM 5302, 5304, 5311, mangrove boat channel, Port Royal, Jamaica (from Hechtel, 1965); USNM 4997, off Florida, USA, 14 m; ZMAPOR 03772b, Plaja Kalkie, Westpunt, Cura��ao; UFBA-POR 207, Barra do Pote, Veracruz, Bahia, Brazil, 12 �� 59 '00" S / 38 �� 36 '00" W, V. Almeida coll., intertidal. Outer morphology (Fig. 14 A���D). Massive, irregularly lobate large sponge (���gibberosa��� means ���hunchbacked���) (Fig. 14 A���B). Size can be up to 30 cm in diameter. Color alive is dark���green to light���brown when exposed to light, otherwise whitish. Color in ethanol is whitish, except the oscule area that stays dark brown. Choanosome color, alive and in ethanol, is whitish. However, in live green specimens, the choanosome just below the cortex can also be greenish. It is a slightly compressible sponge with a dense choanosome, and a tough cortex, difficult to break. Surface is usually smooth, but can be hispid in some areas. Often overgrown by other sponges (e.g. Tethya actinia de Laubenfels, Chalinula molitba (de Laubenfels), Haliclona spp.), ascidians, polychaetes, clams and algae. Oscules are uniporal (0.5���1.5 mm in diameter), each with a sphincter (Fig. 14 F), grouped in circular slightly depressed areas (2���4 cm diameter) (Fig. 14 C). These oscular plates can be situated at the end of lobate projections or not; they are never covered by ectosymbionts. These plates usually are of darker color, brownish. Pores (Fig. 14 D) are cribriporal (diameter of a pore plate: 0.5���1 mm), numerous and evenly distributed over the whole surface. Skeleton (Fig. 14 E���F). The cortex is 0.4���0.9 mm thick and is subdivided between a very thin ectocortex of acanthoxyasters III (ca 20 ��m) and a thicker endocortex of sterrasters (ca 500 ��m). Oxeas I and plagiotriaenes are radially positioned under the cortex with cladomes supporting the endocortex. Under this layer of plagiotriaenes the radial arrangement is less obvious. Most acanthoxyasters II are present right under the cortex while acanthoxyasters I and III are quite abundant throughout the choanosome, as well as developing sterrasters. Oxeas II can be found in the choanosome but are especially placed on the cortex around the oscules (Fig. 14 F). Spicules (measurements from ZMBN 77928, except for the rosette diameter measured in ZMBN 81780) (Fig. 15). Megascleres: (a) oxeas I, stout, straight or very slightly bent, length: 1044��� 1342.1 ��� 1824 ��m; width: 16��� 31.2 ��� 42 ��m. (b) oxeas II, usually straight, length: 157��� 201.5 ��� 238 ��m; width: 3.6��� 4.3 ��� 7.2 ��m. (c) plagiotriaenes (Fig. 15 A), rhabdome length: 792��� 1266 ��� 1620 ��m; rhabdome width: 35��� 48.1 ��� 70 ��m; clad length: 98��� 211.4 ��� 308 ��m. (d) anatriaenes (Fig. 15 B), rare, rhabdome length: 1433 ��m (N= 1); rhabdome width: 6��� 7.9 ��� 11 ��m (N= 3); clad length: 23��� 26.5 ��� 33 ��m (N= 3). Microscleres: (e) sterrasters (Fig. 15 C���D), oval, with smooth 3���5 branched rosettes at their surface (diameter: ca. 4 ��m), length: 75��� 84.1 ��� 93.1 ��m; width: 54��� 79.5 ��� 93.1 ��m. (f) acanthoxyasters I (Fig. 15 E), 6���12 thin actines, diameter: 16��� 21.3 ��� 34 ��m. (g) acanthoxyasters II (Fig. 15 F), large centrum, with thicker and shorter actines than acanthoxyasters I, 14���20 actines, diameter: 9.7��� 13.4 ��� 16.2 ��m. (h) acanthoxyasters III (Fig. 15 G), 9���17 actines, diameter: 4.3��� 6.5 ��� 7.6 ��m. Habitat in the Bocas del Toro region. Common on mangrove roots, 1���2 m depth. Distribution. Georgia, Florida, Texas, U.S.A. (de Laubenfels 1936 b; 1953; Little 1963; Freeman et al. 2007); Bermudas (de Laubenfels 1950); Bahamas (de Laubenfels 1949; Wiedenmayer 1977); Cuba (Alcolado 2002); Jamaica (Bowerbank 1872; Hechtel 1965; Lehnert & van Soest 1998); Dominican Republic (Bowerbank 1873); Puerto Rico (Pulitzer-Finali, 1986); St. Thomas (Duchassaing de Fonbressin & Michelotti 1864); St. John (Uliczka 1929); Barbados (Uliczka 1929; van Soest & Stentoft 1988); Mexico (Lehnert 1993); Honduras (Bowerbank 1872); Belize (R��tzler et al. 2000); Costa���Rica (Loaiza Coronado 1991; Cort��s 1996); Panama (de Laubenfels 1936 a; Wulff 2000); Colombia (D��az 2007); Cura��ao (van Soest 1981); Venezuela (Carter 1882; Sutherland 1980); French Guiana (Lamarck 1815); Brazil (da Silva 2002). Remarks and discussion. After the Galeta and Panama Canal locality records, this is the third record of G. gibberosa in Panama. In fact, G. gibberosa is one of the sponge species that have been able to cross to the Pacific side using the Panama Canal (de Laubenfels 1936 a). The spicule sizes and morphologies of our specimens fitted previous descriptions (da Silva 2002) and the comparative material, except for the Florida specimen (USNM 4997) which had a more regular gross morphology and smaller sterrasters with a different rosette pattern (data not shown). We suggest that the status of Florida populations should be tested in the future. This is the first observation of anatriaenes in G. gibberosa but due to their rarity, they could have easily been overlooked in previous observations. It is also the first time that oxeas II are observed in high density around the oscules (Fig. 14 F). The lobate morphology of our specimens (Fig. 14 B) was very similar to that of the holotype of Geodia tumulosa (Bowerbank 1872, pl. XLVII), later synonymized with G. gibberosa (Carter, 1882; da Silva, 2002). G. gibberosa is a common wide-spread Tropical western Atlantic species. In the literature it appears that G. gibberosa is very polymorphic when it comes to its gross morphology (lobate to flat, massive to encrusting) and its color (white, brown, green and black). A pattern emerges when one considers its two habitats (reef and mangrove). Reef specimens tend to be smaller in size with common encrusting forms, and are usually white to brown. On the other hand, mangrove specimens like ours tend to be large, massive, lobate, and darker colored. Ecology studies have shown that G. gibberosa is very palatable for reef fishes and has no chemical defenses (Pawlik et al. 1995). Therefore, it uses secondary metabolites to promote overgrowth of other species better equipped to defend themselves from fish predation (Engel & Pawlik 2005). All the specimens we observed in Bocas del Toro were indeed covered with numerous sponges, ascidians, algae, etc. Fish predation being lower in mangroves (Dunlap & Pawlik 1996), G. gibberosa can reach bigger sizes and grow in more open habitats thereby receiving more sunlight and having darker colors. Conversely, predation pressure in reefs being higher, G. gibberosa is usually smaller and prefers cryptic habitats (under rocks or other sponges, crevices). When not exposed to light it is of lighter color, often white. Seemingly, two other mangrove sponges (Tedania (Tedania) ignis (Duchassaing de Fonbressin & Michelotti) and Chondrosia sp.) were found in cryptic habitats when collected on reefs (Dunlap & Pawlik 1996). An alternative to the polymorphism hypothesis is that we simply have two (or more) cryptic species. Both hypotheses should be tested combining morphology and molecular data., Published as part of C��rdenas, Paco, Menegola, Carla, Rapp, Hans Tore & D��az, Maria Cristina, 2009, Morphological description and DNA barcodes of shallow-water Tetractinellida (Porifera: Demospongiae) from Bocas del Toro, Panama, with description of a new species, pp. 1-39 in Zootaxa 2276 on pages 28-31, DOI: 10.5281/zenodo.191088, {"references":["Lamarck, J. B. P. (1815) Suite des polypiers empates. Memoires du Museum d'Histoire Naturelle, Paris, 1, 69 - 80, 162 - 168, 331 - 340.","Schmidt, O. (1870) Grundzuge einer Spongien-Fauna des atlantischen Gebietes. (Wilhelm Engelmann: Leipzig), iii - iv, 1 - 88, pls I - VI.","Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103.","Duchassaing de Fonbressin, P. & Michelotti, G. (1864) Spongiaires de la mer Caraibe. 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(1977) Shallow-water sponges of the western Bahamas. Experientia Supplementum 28, 1 - 287, pls 1 - 43.","Alcolado, P. M. (2002) Catalogo de las esponjas de Cuba. Avicennia, 15, 53 - 72.","Lehnert, H. & van Soest, R. W. M. (1998) Shallow water sponges of Jamaica. Beaufortia, 48, 71 - 103.","van Soest, R. W. M. & Stentoft, N. (1988) Barbados Deep-Water Sponges. In: Hummelinck, P. W. & Van der Steen, L. J. (Eds.) Studies on the Fauna of Curacao and other Caribbean Islands. Uitgaven van de Natuurwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen, No. 122., Amsterdam, pp. 1 - 175.","Lehnert, H. (1993) The sponges from Cozumel (Mexico). Inventory, critical comparison of taxonomic characters and description of a new species. Acta Biologica Benrodis, 5, 35 - 127.","Rutzler, K., Diaz, M. C., van Soest, R. W. M., Zea, S., Smith, K. P., Alvarez, B. & Wulff, J. (2000) Diversity of sponge fauna in mangrove ponds, Pelican Cays, Belize. 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Stelletta sp. (Figure 10) Material. ZMBN 81643, Solarte lagoon (9 ° 18.35 N, 82 ° 10.39 W), on mangrove root, 1 m depth. Additional material examined. Stelletta soteropolitana Cosme & Peixinho, 2007, UFBA-POR 500, holotype, Salvador, Bahia State, Brazil, Outer morphology (Fig. 10 A–B). Massive, sub–globular sponge, 10 cm high, with a diameter of 9 cm. Surface color alive is dark–purple to whitish. Choanosome color is cream In ethanol, cortex and choanosome are grayish. One large oscule (diameter: 2 cm) with a contractile membrane is on the top surface. Under the oscule is a large cloaca (6 cm deep) with cribriporal exhalant apertures. Unconspicuous cribriporal pores are evenly distributed, each pore is 100–1000 µm large. Slightly–compressible, rugose surface with irregular ridges, grooves and knobs (2–7 mm high). Mussels and oysters are embedded in it. A second specimen was observed (but not collected): it was about twice the size of ZMBN 81643, with two oscules. Skeleton (Fig. 10 C–D). Dense radial bundles of plagiotriaenes and anatriaenes, especially near the surface. Their cladomes are often found at or beyond the surface of the sponge. There is a slightly differentiated cortex, not visible with the naked eye, except for the pigmentation. Bundles of megascleres are looser under the cortex and triaenes less abundant. Acanthotylasters, oxeas I and II are abundant throughout the sponge. Spicules (Fig. 10 E–G). Megascleres: (a) oxeas I, large, slightly bent or straight, length: 1080– 1236.3 – 1440 µm; width: 18– 24.4 – 32 µm. (b) oxeas II (Fig. 10 E), smaller, with very thinly pointed ends, almost whip–like, generally with a bend or a double bend, length: 303– 724.7 – 1000 µm; width: 3– 9.5 – 14 µm. (b) plagiotriaenes (Fig. 10 F), with rare dichotriaenes, rhabdome can have a straight pointed or whip–like end, rhabdome length: 511– 850.9 – 1056; rhabdome width: 13– 31.5 – 39 µm; clad length: 50– 121.3 – 152 µm. (c) anatriaenes (Fig. 10 G), common but less frequent than plagiotriaenes, and rare dichoanatriaenes, with depressed apex, rhabdome length: 716– 1363.2 – 2280 µm; rhabdome width: 8– 21.7 – 29 µm; clad length: 38– 65.1 – 91 µm. Microscleres: (d) acanthotylasters (Fig. 10 H), 5–8 actines with scarce spines, actine is only slightly tylote, one single spine at the tip of each actine, diameter: 9– 13.5 – 20 µm. Habitat in the Bocas del Toro region. Mangrove roots, rare, 1 m depth. Distribution. Panama (this study). Remarks and discussion. This species appears to be rare in Bocas del Toro. We only sighted two specimens during our survey in 2007, both a few meters apart in Solarte lagoon. At this time, we only took a small piece from ZMBN 81643. In July 2009, both specimens were still there, but were slightly bigger and bleached (Fig. 10 B). The bleaching could be due to shading from the algae growing around it, because when we finally collected ZMBN 81643 and kept it alive alone in an open tank, it started to gain back some purple colouration. At present there are four known species of Stelletta with tylasters in the Caribbean (S. fibrosa, S. variabilis, S. kallitetilla and S. pudica) and three in Brazil (S. anancora, S. beae Hajdu & Carvalho, 2003 and S. soteropolitana). Of these, only the three Brazilian species share long clads (average of clad length> 100 µm) with our Bocas del Toro specimen. However, when compared with Stelletta sp., S. anancora has (i) only one category of oxeas, (ii) 2–3 triaene sizes, (iii) no anatriaenes and (iv) no dense accumulation of megascleres in its cortex; S. beae has (i) shorter plagiotriaenes (rhabdome length ca 490 µm) with longer clads (ca 200 µm), (ii) shorter anatriaenes with reduced clads (ca 23 µm), (iii) one size of oxea, and (iv) smaller acanthotylasters (ca 11 µm). When it comes to S. soteropolitana it has i) plagiotriaenes twice the size of those in Stelletta sp., ii) protriaenes and iii) no anatriaenes. As we have already shown for S. fibrosa, anatriaenes can be present or absent in the same species, and the missing anatriaenes in S. soteropolitana, at the present time, cannot be used to distinguish the species from our S. sp. The protriaenes found in S. soteropolitana could simply be plagiotriaenes that are more forward oriented. Also, it is important to stress that S. soteropolitana is described from a single specimen and therefore, we have no data on its intra-specific morphological variation, especially concerning spicule sizes. To conclude, we cannot exclude S. soteropolitana as being conspecific with our specimen but in our opinion more data is required to settle this matter.

Lamellomorpha Bergquist, 1968 Lamellomorpha Bergquist, 1968: 30. Diagnosis (modified from Hooper & Maldonado 2002) Massive, lamellate stalked-palmate, or paddle-shaped sponges, with a relatively smooth, granular, or fleshy, slightly conulose surface. Ectosomal skeleton a skin-like membrane packed with microstrongyles. Choanosomal skeletal architecture a core of megascleres, which are straight, curved, sinuous, or contort oxeas, frequently modified with one or both ends rounded as in strongyloxeas. These radiate through the stalk and fan. Straight oxeas arise as short subectosomal tracts that emerge oblique to the surface. Roughened microstrongyles or microxeas and streptasters (amphiasters, metasters, and spirasters) scattered throughout the body. Type species Lamellomorpha strongylata Bergquist, 1968 (by monotypy).