taxonID	type	description	language	source
122687EBFFD02161FC11CCAE6CE9BA73.taxon	description	(FIGS 3 – 6, TABLE 1)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD02161FC11CCAE6CE9BA73.taxon	materials_examined	Type material examined. Isops pallida, near Hammerfest, Norway, 71 ° 12 ′ 5 N, 20 ° 30 ′ 5 E, 247 m, Willem Barents Exp. 1878 – 79, RMNH Por 652, wet specimen (only pictures were seen); RMNH, Vosmaer slide collection, box number 37, three spicule preparations with number 64.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD02161FC11CCAE6CE9BA73.taxon	description	Geodia atlantica, holotype, off western Ireland, 54 ° 17 ′ 5 N, 11 ° 33 ′ 5 W, 709 m, stones and rock, 9.15 ° C, number S. R. 151 - 27 / 364 - 1914. External morphology and cortex: Colour alive and in ethanol is whitish (Norway, Flemish Cap) to light brown (Bay of Biscay, Rockall Bank). Choanosome colour alive and in ethanol is brownish (always darker than cortex). Large specimens can be funnel shaped with a deep cavity with irregular swellings and ridges (Fig. 3 A, B). From fragments we have seen in large trawl catches at the Faroe Islands, we had the impression that in very large specimens the bottom of the funnel could disappear with time, and that the sponge maintained a ring-shaped wall, 80 – 100 cm in diameter. The largest specimen found measures 72 ¥ 40 ¥ 39 cm. Other specimens are irregularly plate-shaped and convoluted (Fig. 3 C, D). Young specimens are subspherical (Fig. 3 E, G). The surface is smooth. The lower sides of the specimens are sometimes covered with stones incorporated in the cortex. Uniporal oscules (0.3 – 0.5 mm in diameter) are scattered on the top surface of small specimens, and oscules are sometimes surrounded by a raised lighter-coloured boundary. Oscules are scattered on the inner side of funnel-shaped specimens so that pores and oscules are on opposite sides (Fig. 3 H). Cribriporal pore areas (0.3 – 0.5 mm in diameter) are on the outer side of funnel-shaped specimens and on one side of plate-shaped specimens; they are in small groups, which are evenly scattered over the surface (Fig. 3 I). The cortex is elastic, c. 0.5 mm thick (with ectocortex: 30 – 100 Mm) (Fig. 4). Anatriaenes within the choanosome are fairly common (Fig. 4 F). Description of type material: Three spicule preparations (with number 64) (Fig. 4 A) of the syntype b of Isops pallida. Pictures of the spicules are shown (Fig. 4 B). We have seen only pictures of the wet specimen of syntype b: it is a whole spherical specimen about 4 cm in diameter, with very small roots, a thin cortex (<1 mm thick), slightly raised uniporal oscules and cribriporal pores on opposite sides. It looks similar to specimen ZMAPOR 21406 a from Norway (Fig. 3 F). The holotype of G. atlantica is a small subspherical specimen (2.7 ¥ 2 cm). This specimen is represented in plate II of Stephens (1915) and Figure 3 G. In the Dublin Museum, there are also five spicule preparations made by Stephens (four spicule slides and one section). New thick sections were made for this study (Fig. 4 C, D). Figure 5 shows SEM pictures of this holotype. Spicules (Figs 4 – 5, Table 1): Megascleres: (a) oxeas, straight or bent, length: 1275 – 4440 Mm; width: 11 – 68 Mm. (b) Orthotriaenes, rarely dichotriaenes, straight or slightly bent rhabdome, rhabdome length: 630 – 4400 Mm; width: 18 – 125 Mm; orthotriaene clad length: 95 – 750 Mm; protoclad length: 190 – 430 Mm; deuteroclad length: 90 – 300 Mm. (c) Anatriaenes, straight or slightly bent rhabdome, rhabdome length: 376 – 5200 Mm; width: 2 – 32 Mm; clad length: 9 – 300 Mm. (d) Protriaenes, very rare [one reported in the type (Stephens, 1915) and one observed in PC 626], rhabdome length: 3000 Mm; width: 8 – 15 Mm; clad length: 96 – 130 Mm. Microscleres: (e) sterrasters, slightly elongated, more rarely spherical, length: 80 – 125 Mm; width: 75 – 112 Mm; thickness: 70 – 88 Mm. Rosettes are made of 2 – 6 smooth rays; rosette diameter: 4 – 7 Mm; hilum diameter: 10 – 20 Mm. (f) Spheroxyasters, rough actines, 5 – 16 Mm in diameter. (g) Oxyasters I, 3 – 8 rough actines, diameter: 22 – 110 Mm [maximum measured in type by Stephens, (1915)]. (h) Oxyasters II, 9 – 25 rough actines, usually with a larger centrum than oxyasters I, diameter: 12 – 35 Mm. DNA barcodes: GenBank accession nos. HM 592679, HM 592695, EU 442195 (Folmer COI): we have sequenced specimens from western and northern Norway (10), Rockall Bank (1), and Flemish Cap (1): the Folmer COI is identical in all these specimens. No. KC 481227 (18 S), obtained from ZMBN 77927 (Korsfjord, Norway). Distribution (Fig. 6): This species has an amphi- Atlantic boreal distribution. It has been recorded at depths of 65 – 2338 m, the shallowest record being from divers in Sandsfjord, Rogaland, Norway (Moen & Svensen, 2008), the deepest boreal records being from south of Iceland and off south-east Greenland, while the deepest record overall is from the Bay of Biscay. Geodia atlantica seems only to be present in the south-western Barents Sea and absent in Arctic waters, which might explain why it is not mentioned by Koltun (1966). It has been found at temperatures between 1.4 ° C (Denmark Strait) and 10.5 ° C, but is usually found at temperatures higher than 3 ° C. Biology: We found no indications of asexual reproduction. The predatory chiton Hanleya nagelfar Lovén, 1846 and the parasitic foraminiferan Hyrrokkin sarcophaga Cedhagen, 1994 have been found living on G. atlantica (Warén & Klitgaard, 1991; Cedhagen, 1994; Todt et al., 2009). Sea urchins are also possibly feeding on this sponge; the two species observed in Figure 3 C were tentatively identified from the photo as Cidaris cidaris (L., 1758) and Gracilechinus alexandri (Danielsen & Koren, 1883) (T. Saucède, pers. comm.). Cidaris cidaris (Rouho, 1888; Mortensen, 1928) and other cidarids (Bo et al., 2012) are indeed considered to be sponge predators whereas G. alexandri is more of an omnivore opportunist which may be more interested in the small organisms living around and on the sponge. Other associated fauna has been investigated by Klitgaard (1995). The chemistry (elemental analysis, amino acids, sterols, and quaternary ammonium compounds) has been investigated by Hougaard et al. (1991 a, b). Distinctive characters: External morphology: The deep funnel shape or plate convoluted shape, with smooth surface. The pattern of distribution of pores and oscules: when one finds a fragment of a funnel or plate-shaped specimen, oscules are on one side, pores on the other. Spicules: Lack of microxeas (as in G. phlegraei and G. parva) and very common anatriaenes. Remarks: Burton (1930) synonymized I. pallida with G. phlegraei by stating that he had compared type Means are in bold; other values are ranges; N = 30 unless stated otherwise in parentheses, or unless measurements come from other studies. A dash indicates that this measurement is not given in the literature. n. f., not found; n. o., not observed in the subsample in our possession. slides from the ‘ Norman collection’, and taxonomists followed his conclusions. Even Vosmaer (1933: 141 – 142) accepted the synonymy after having examined a slide of G. phlegraei sent to him by Sollas. However, the only slides of I. pallida that we found in the Norman collection (BMNH 10.1.1.1149 to 1156 and MNHN-DN 45) had labels saying ‘ Isops pallida Vosmaer / Lervig, Norway, 1879 ’. In 1879, Norman did stay in Leirvik (current name of ‘ Lervig’) on the island of Stord in the Hardangerfjord in western Norway (Norman, 1893), so these are clearly not from the type of I. pallida (which was collected near Hammerfest in northern Norway). Interestingly, the thin cortex (0.5 mm), the large spiny oxyasters, and spheroxyasters showed that these slides from the Norman Collection were not from a G. phlegraei but from a G. atlantica. So Burton (1930) had probably not examined type slides and had not noticed the difference between G. atlantica and G. phlegraei spicules. However, in the course of this study, Rob van Soest (Naturalis, Leiden) rediscovered in the Vosmaer slide collection (box 37) five slides with labels stating ‘ Isops pallida Vosm’ and ‘ Bar. I p. 16 ’ (Fig. 4 A) which was understood as a reference to the Barents Sea expedition 1 st publication, and to the page number of the original description of I. pallida in Vosmaer (1882). We therefore concluded that these five slides were the syntype slides of I. pallida. Three slides had the number 64 while two slides had the number 62 (Fig. 4 A); the spicule morphologies and abundance corresponded to the descriptions given of syntype a (= 62) and syntype b (= 64), the latter being the one represented in the original plates (external morphology and spicules). Indeed, Vosmaer (1882) clearly notes that specimen a has considerably fewer oxyasters and anatriaenes than specimen b. This is because Vosmaer (1882) simply mixed one specimen of G. phlegraei (a) with one specimen of G. atlantica (b): spicule morphologies (spheroxyasters, large spiny oxyasters, abundant anatriaenes) (Fig. 4 B) and sizes (Table 1) clearly show that specimen b of I. pallida type material is conspecific with G. atlantica. Since Vosmaer, (1882) did not explicitly designate a holotype, we have the possibility to designate a lectotype. We formally designate the syntype a as the lectotype of Isops pallida Vosmaer, 1882. Recommendation 74 B of the ‘ International Code of Zoological Nomenclature’ states that in choosing a lectotype among syntypes, preference should be given to the illustrated specimen, in our case syntype b. But we decide to go against this recommendation for the following reasons: (1) a comes before b in the alphabet, (2) atlantica has been used far more in the literature than pallida and, above all, (3) I. pallida has always been considered a junior synonym of G. phlegraei (so our deci- sion will preserve the stability of the nomenclature). Following our decision, specimen b then becomes a misidentification, and I. pallida does not become a junior synonym of G. atlantica. Just before the publication of this revision, Rob van Soest discovered in 2013 a jar (RMNH Por. 652) labelled: ‘ Isops sphaeroides Vosm (type v. I. pallida Vosm.) W. Barents exp. 1878 / 79, 71 ° 12 ′ 5 ″ N 20 ° 30 ′ 5 ″ O, Coll. G. C. J. Vosmaer 12 Juli 1879 ’. Inside are two specimens and another small label written in pencil ‘ Isops pallida, N. Archive Suppl. 1 ’ which refers to the original description of I. pallida by Vosmaer in ‘ Niederlaendisches Archiv fuer Zoologie Supplementband 1 ’. We therefore believe that these two specimens are the two syntypes of I. pallida. The external morphologies (observed from pictures, courtesy of R. van Soest) of the largest specimen (a = lectotype) and of the smallest specimen (b) confirm the above conclusion based on spicules: a is a G. phlegraei and b a G. atlantica. We noted that G. atlantica had a second smaller category of oxyasters that Vosmaer (1882) has seen in specimen b, but not Stephens (1915). Stephens (1915) states that the cortical spheroxyasters become larger in the choanosome; these are actually the oxyasters II. Admittedly, spheroxyasters and oxyasters II can be difficult to separate in spicule preparations, unless carefully measured and examined in thick sections (spheroxyasters are in the ectocortex, oxyasters II usually in the choanosome just below the cortex). In some specimens anatriaenes may be separated into two size categories (e. g. UPSZMC 78293 from the Flemish Cap), especially based on the rhabdome length (376 – 530 vs.> 2000 Mm), but since a continuum of anatriaene sizes exists in other specimens (e. g. PC 222 from northern Norway), we refrained from doing so. There is usually a clear predominance of orthotriaenes over dichotriaenes (but not always the case, see MNHN-ThalassaZ 407). The main difference between the type and the Norwegian specimens is that, in the Norwegian specimens, the asters are less spiny, and the oxyasters I are smaller and much less abundant (Fig. 5); this may be due to the shallower environment of the Norwegian specimens (200 – 400 m) compared with the type (709 m). No consistent morphological differences were found between specimens from the Flemish Cap and specimens from the NEA. A more NEA southern morph may be present (found in the MNHN Thalassa and Centob collections). These specimens are irregularly plate shaped, with a darker external colour and are usually found growing around coral. We have never seen it with the characteristic funnel shape but we have only seen small specimens (less than 15 cm long).	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD8217FFF7AC8C86E69BF77.taxon	description	(FIGS 7 – 10, TABLE 2)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD8217FFF7AC8C86E69BF77.taxon	materials_examined	Type locality and deposition of holotype Geodia barretti, collected by Robert McAndrew (1802 – 1873). South side of Vikna Island (formerly called Vigten or Vikten Island), North-Trøndelag, Norway, 183 m, BNHM 1877.5.21.1399 (dry specimen), BNHM 1877.5.21.1400 (one slide of surface and one spicule preparation), BNHM 1877.5.21.1401 (slide of section). Geodia simplicissima, Foldenfjord, northern Norway, 10 – 75 m, TSZY 10 (wet specimen). Spicule preparations made during this study are now stored at TSZY. External morphology and cortex: Irregularly massive, up to at least 80 cm in diameter, and up to a weight of c. 38 kg (wet); young specimens are usually spherical to subspherical. Mostly with an obvious attachment area, sometimes formed as several stilt-like projections each attached to a piece of gravel. The surface colour (alive) is usually white (Fig. 7 A-C, E), but with sometimes various shades of light yellow (Fig. 7 D) or light brown (Figs 7 F, 8). The choanosome alive is light brown (Fig. 7 C) and becomes whitish in ethanol. The surface is usually clean and smooth but shallow specimens (30 – 50 m) can be slightly dirty and hispid. Some NWA specimens were very hispid over their entire surface. One to many (more than 30) preoscules (i. e. a depression protecting the true oscules), more or less deep, more or less narrow, with a circular to irregular opening (up to several cm wide) (Fig. 7 A – F). Preoscules are generally on top. The preoscule contains uniporal oscules (Fig. 7 G). Each oscule (1 mm in diameter) has a sphincter. Cribriporal pores are scattered over the entire body surface (Fig. 7 H); single pores are 50 – 80 Mm, and poral sieves are c. 0.5 mm. The cortex is 0.4 – 0.6 mm (ectocortex: c. 250 Mm, endocortex: c. 750 Mm) (Fig. 9 A). In the preoscule, the cortex is without sterrasters and triaenes (Fig. 9 B), and ridges of microxeas and strongylasters surround the uniporal oscules (Figs 7 G, 9 B). Description of type material: Medium-sized oval specimen (length: 12 cm, width: 8 cm) from Bowerbank (1872 a: plate XI) which has been cut into five pieces (the main specimen and four smaller pieces); three Bowerbank slides including spicule preparation, section, and cortex surface. Spicules (Fig. 9, Table 2): Megascleres: (a) oxeas I, straight or bent, length: 1075 – 4450 Mm; width: 15 – 75 Mm. (b) Oxeas II, straight or bent, rarely modified to styles, sometimes slightly centrotylote, length: 190 – 900 Mm; width: 4 – 16 Mm. (c) Dichotriaenes, rare orthotriaenes, rhabdome length: 620 – 4600 Mm; width: 20 – 150 Mm; orthotriaene clad length: 240 – 500 Mm; protoclad length: 100 – 400 Mm; deuteroclad length: 45 – 450 Mm. (d) Anatriaenes, rhabdome length: more than 7.4 mm; width: 9 – 40 Mm; clad length: 50 – 250 Mm. (e) Meso / protriaenes (rare), rhabdome length: up to 2640 Mm; width: 7.5 – 15 Mm; clad length: 25 – 115 Mm; central clad length: 25 – 98 Mm. Microscle- res: (f) sterrasters, spherical to elongated, length: 65 – 130 Mm, width: 51 – 105 Mm, thickness: 60 – 80 Mm; hilum diameter: 12 – 23 Mm. Rosettes are made of 3 – 7 rays, covered with warts; rosette diameter: 4 – 7 Mm. (g) Strongylasters, rough actines, 3 – 11 Mm in diameter. (h) Oxyasters I (only in very deep specimens> 1000 m), rough actines, diameter: 30 – 80 Mm. (i) Oxyasters II, rough actines, diameter: 6 – 32.5 Mm. The spiculogenesis of shallow specimens (30 – 50 m depth) being somewhat disrupted, their spicule measurements have not been included here but they are shown in Table 2 and discussed in Cárdenas & Rapp (2013). DNA barcodes: We found two haplotypes for the COI Folmer marker. GenBank accession nos. HM 592679, HM 592695, and EU 442195: haplotype 1 from Spitsbergen (5), southern, western, and northern Norway (12), Sweden (1), off western Ireland Means are in bold; other values are ranges; N = 30 unless stated otherwise in parentheses, or unless measurements come from other studies. A dash indicates that this measurement is not given in the literature. n. f., not found; n. o., not observed in the specimen in our possession (usually because the sample was too small). (2), Davis Strait (1), Flemish Cap (1), and the Mediterranean Sea (1). No. KC 574389: haplotype 2 (1 - bp difference with haplotype 1 in position 382: ‘ A’ instead of ‘ T’) was found in two specimens from the Flemish Cap (UPSZMC 78262, UPSZMC 78268). Nos. EU 552080, HM 592809 (28 S, C 1 - D 2 domains): we have sequenced 28 S (C 1 - D 2) from specimens from Spitsbergen (1), western Norway (2), and off Ireland (1): we did not observe genetic differences in this marker among NEA specimens. No. KC 481224 (18 S), obtained from ZMBN 77922 (Korsfjord, Norway) and ZMBN 89722 (Spitsbergen): no variation was observed. Distribution (Fig. 10): Geodia atlantica, G. barretti, and G. hentscheli may have been confused in the past, especially until the description of G. atlantica by Stephens (1915) and G. hentscheli by Hentschel (1929), and above all when juveniles were found (e. g. Burton, 1949). This should be kept in mind when examining the G. barretti distribution map that includes a few records not verified by us. However, we did check specimens from Fristedt (1887) (SMNH), Lundbeck (1909) (ZMUC), Boury-Esnault et al. (1994) (MNHN), Voultsiadou & Vafidis (2004), Nichols (2005), and van Soest et al. (2007) (ZMAPOR). Specimens from Voultsiadou & Vafidis (2004) and van Soest et al. (2007) were mis-identifications (cf. Discussion). The record in the Asturias (Spain) given by Ferrer-Hernández (1918) at 150 – 300 m depth is based on slides, and it is dubious as it seems too shallow for this species at this latitude, but he unfortunately gives no description. Other identifications could be confirmed by accurate descriptions and plates (e. g. Vosmaer, 1882). Geodia barretti has been found at depths from 30 to 2000 m. Most NEA records are from between 200 and 500 m, at temperatures of 4 – 8 ° C; Grand Banks, Flemish Cap, Nova Scotia, and Davis Strait specimens were found at 410 – 1852 m, at temperatures of 3 – 5 ° C. Shallow specimens from the western Norwegian coast have been collected at temperatures of 3 – 9 ° C, and possibly experience up to 14 – 15 ° C in September – October (Cárdenas & Rapp, 2013). The only specimen we identified from the Mediterranean Sea was collected at 167 m where the water temperature is around 13 ° C and the salinity usually more than 38 p. p. m. Localities where the species occurs at lower temperatures, down to 0.4 ° C, were only found in the Denmark Strait. Breitfuss (1930) reports G. barretti in the southern part of the Kara Sea at – 1.75 ° C but we have not examined this specimen, and because no other records exist of this species in this area, this record needs to be confirmed and is here considered dubious. Blacker (1957) only gives the coordinates for his 1949 and 1950 trawls; we could not find coordinates for the 1951, 1952, 1954, and 1955 trawls. Likewise, Dyer et al. (1984) do not give coordinates for their 1978 – 81 trawls. We therefore manually copied on Figure 10 the G. barretti records between northern Norway and Spitsbergen from figure 3 c in Dyer et al. (1984), which also integrates the Blacker (1957) localities. Biology: Gametogenesis has been well studied as well as the annual reproductive cycle (Spetland et al., 2007). This study on Scandinavian fjord populations shows that G. barretti is (1) gonochoric and oviparous and that (2) reproduction coincides with phytoplankton blooms. Gametogenesis usually takes place from February to May with a gamete release in early summer; sometimes a second gametogenesis / spawning event takes place later in the summer (Spetland et al., 2007). In our only specimen from the Mediterranean Sea, collected on 22 August 2010, spermatogenesis was observed. We found no indications of asexual reproduction in this species.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD8217FFF7AC8C86E69BF77.taxon	description	Distinctive characters: External morphology: the generally smooth surface (absence of hispidity and epibionts) and white colour. The irregular form, especially in specimens larger than about 15 cm in diameter. The clearly visible sieves in the sometimes numerous preoscular cavities. Spicules: usually dichotriaenes and strongylasters (but these characters are not sufficient as G. hentscheli can also have both). Remarks: As explained before (Cárdenas et al., 2010), we stress that G. barretti ’ s oscules are not covered by a sieve. There is a depression called a preoscule, in which we find single uniporal oscules (without any kind of sieve). Every oscule has its own unique sphincter, and this is clearly visible with the naked eye (Fig. 7 G) or in a thick section (Fig. 9 B). We find the same arrangement in G. hentscheli (cf. below). Burton (1949) identified some very small Geodia specimens as G. barretti; this identification is probably wrong, as the specimens seem to have been buds, and came from ‘ an unspecified point in the Arctic and from an unknown depth’. But he pointed out the similarity of some of his specimens to G. parva. Indeed, what he had in front of him must have been buds from G. hentscheli or G. parva.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD8217FFF7AC8C86E69BF77.taxon	materials_examined	We examined a spicule slide of the specimen identified as G. barretti from the Mediterranean Sea (specimen A 183), collected at a surprising 4 – 6 m depth in the Aegean Sea (Voultsiadou & Vafidis, 2004). In our opinion, this is a misidentification; the dichotriaenes and oxyasters are typical of Geodia conchilega Schmidt, 1862, a common Mediterranean shallow species for which we had comparative material (e. g. MNHN DNBE- 846, G. conchilega from Banyuls, France, collected and identified by N. Boury- Esnault). But we did identify a G. barretti specimen collected in the ‘ Canyon des Moines’ (south Corsica) at 167 m depth (‘ CorSeaCan’ campaign with the ROV Achille). The spicule morphologies match those of our other specimens: there are only oxyasters II (10 – 35 Mm); microxeas can be slightly bent and are occasionally centrotylote; the cortex thickness is standard (0.5 – 0.6 Mm). The main and only difference we could find is that the spherical sterrasters are smaller (56 – 59.9 – 65 Mm) than in the Atlantic G. barretti (Table 2), except those from shallow waters. It had the same COI Folmer haplotype 1 as all the NEA and most of the NWA specimens. This is the first true record of this species in the Mediterranean Sea. At least six additional sightings between 167 and 199 m depth (without collection) of G. barretti - like specimens were made during the ‘ MedSeaCan’ and ‘ CorSeaCan’ campaigns: in the ‘ Banc de Magaud’, ‘ Banc de Nioularge’ (both off the Côte d’Azur), and ‘ Canyon de Cargèse’ (western Corsica) (J. Vacelet & M. Fourt, pers. comm.). Overall, these G. barretti - like specimens seem less smooth than northern ones, with regular small bumps on their surface (where megascleres cross the cortex and retain sediments); unlike the NEA specimens, they always had a single deep, wide preoscule (c. 3 cm in diameter). We cannot be completely sure they are all G. barretti because Geodia megastrella Carter, 1876 can have a very similar external morphology, although it has never been observed in the Mediterranean (although we have found it in the Balgim material collected off Morocco, along with G. barretti). The spiculogenesis of shallow specimens (30 – 50 m) is disrupted so that spicule morphologies are somewhat different (Cárdenas & Rapp, 2013). Thus, it has been shown that G. simplicissima from northern Norway is actually a G. barretti growing in shallow waters. G. simplicissima has therefore been put in synonymy with G. barretti (Cárdenas & Rapp, 2013). We examined the holotype of Geodia barretti divaricans [MOM 04 - 1333 (wet specimen) and MNHN DT- 1299 (type slide)]. In the choanosome, it has slightly spined oxyasters which can reach a very large size (25 – 70 Mm in diameter) and just under the cortex strongly spined oxyspherasters (17 – 22 Mm in diameter), which are different from the oxyasters II of G. barretti. Geodia divaricans is clearly different from G. barretti.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFD8217FFF7AC8C86E69BF77.taxon	description	No morphological differences were found between specimens from the NWA (Flemish Cap, Nova Scotia, Davis Strait) and specimens from the NEA. We just note that some NWA specimens were very hispid (e. g. UPSZMC 78268 – 78269), a feature never observed in NEA specimens. NWA Specimens deeper than 1000 m depth do have oxyasters I but they are much smaller (c. 32 – 42 Mm) than in NEA specimens deeper than 1000 m (up to 58 – 80 Mm), and more difficult to consider as a separate size category. Also, NWA specimens deeper than 1000 m have subspherical sterrasters, not elongated like in NEA specimens collected at the same depths. No morphological differences were observed between the two haplotypes (Table 2: UPSZMC 78269 vs. other specimens). Interestingly, haplotype 2 is closer by 1 bp to the sequence of G. hentscheli than haplotype 1: it is therefore closer to the common ancestor of these sister-species, which would suggest that the common ancestor lived in the NWA. It so happens that UPSZMC 78268 (haplotype 2) was initially identified as G. hentscheli due to its important hispidity, spherical shape, and narrow unique preoscule (see a complete description of UPSZMC 78268 on the Sponge Barcoding Project, http: // www. spongebarcoding. org). Further work is needed to see if haplotype 2 has a consistently G. hentscheli - like morphology (as in Fig. 7 A). Blacker (1957) and Dyer et al. (1984) sampled extensively between northern Norway and Spitsbergen between 1949 and 1981 and found extensive sponge grounds. But when O. S. T. participated in the Meteor 1990 cruise in the same area, very few Geodia were collected (Barthel, Tendal & Witte, 1991). Eight triangular dredges were made in the southern area off Bear Island, and only one small Geodia was collected. In the northern area off Spitzbergen numerous triangular dredges were made and no Geodia were collected; a single large specimen of G. macandrewii was taken by a hyperbenthic sledge. There is a possibility that the reason for this sampling discrepancy is the sampling method – Blacker (1957) and Dyer et al. (1984) used a trawl while a large triangle-dredge was essentially used on the Meteor – but this does not seem very likely. Alternatively, the large masses of sponges earlier reported may have disappeared since 1981 due to an inflow of very cold water from the north, intensive trawling activity in the area, or disease, although it is difficult to believe that any of these would hit such a large area. It is also possible that the Meteor cruise was rather unlucky in finding sponge grounds. The ‘ Ecosystem Barents Sea’ cruise in 2007 collected tonnes of Geodia in station 2562, but this was much closer to the Norwegian coast (c. 80 km). Mass mortality of G. barretti was actually observed in the Kosterfjord area (southern Norwegian and western Swedish waters) and started in the winter of 2006 / 07; this may be due to unusually high temperature and a deepening of the thermocline in 2006 and 2008 in this region (Guihen et al., 2012). Maximum temperatures in the autumn 2006 / 08 at Tisler reef were 12.5 ° C instead of 9 ° C in other years. However, shallow specimens at 30 m depth on the western Norwegian coast seem to experience up to 14 – 15 ° C in September – October (Cárdenas & Rapp, 2013). So the dramatic rate of change in temperature in the Kosterfjord (4 ° C in less than 24 h) is more likely to be one of the causes of this mass mortality. The population still suffers from the incident and the mortality is still high (P. C. & M. T., ROV observations at c. 80 m depth in Swedish waters of the Kosterfjord in May 2012).	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFC12171FCF8C8C86E3DB8BE.taxon	description	(FIGS 11 – 14, TABLE 3)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFC12171FCF8C8C86E3DB8BE.taxon	materials_examined	Type locality and deposition of types: Hentschel (1929) explicitly designated a holotype (ZMB Por 7549 has an old label saying ‘ Sidonops mesotriaena n. sp. Typ’) from north of Spitsbergen, station 41, 81 ° 20 ′ N, 20 ° 30 ′ E, 1000 m, 11 th of August 1898. Paratypes are ZMB Por 7545 – 7546, 7548, 7550 – 7551, 8421, all from st. 41 and st. 40 (81 ° 22 ′ N, 21 ° 21 ′ E, 650 – 1000 m). External morphology and cortex: The body is spherical (Fig. 11 A – D), up to about 15 cm in diameter. The colour alive is white to light yellow or brownish. With the use of ROVs, we found that this species can be entirely covered with long spicules, making them look very fury (Fig. 11 D), which may be lost when dredged, and specimens then seem completely smooth (Fig. 11 A – C). Some specimens are budding (Fig. 11 C), and buds are usually columnar with sometimes a small peduncle. A preoscule opening, usually narrow (up to 1.8 cm in diameter), occasionally up to three, is found on the top side. This preoscule is even observed in young specimens (8 mm in diameter). These preoscule openings are often surrounded by a narrow elevated ring with a thickened cortex. Sometimes, in large specimens, the preoscule opening ‘ sinks’ in the sponge body (one such specimen can be seen in the upper right corner of Fig. 11 B). As in G. barretti, uniporal oscules are concentrated in the preoscule (Fig. 11 E); the cortex there is without sterrasters, and ridges of microxeas and strongylasters surround oscules. These ridges can be much more developed than in G. barretti. The cribriporal pores are scattered over the sides of the body (total diameter of the sieve: 0.1 – 0.2 mm) (Fig. 11 F). The sterraster layer is elastic, 0.25 – 1.4 mm thick (Fig. 12), with the ectocortex poorly developed (0.1 mm thick, in holotype) (Fig. 12 B) to well developed (0.3 mm thick) (Fig. 12 C). Description of type material: The holotype ZMB Por 7549 is cut into four pieces. For this study, we have examined only a small slice of the paratype ZMB Por 7551. Thick sections of ZMB Por 7551 made during this study (Fig. 12 A) are now stored at the ZMB. Figure 13 shows SEM pictures of the spicules of the paratype ZMB Por 7551. Spicules (Fig. 13, Table 3): Megascleres: (a) oxeas I, straight or bent, length: 1200 – 5175 Mm; width: 29 – 82 Mm. (b) Oxeas II, usually straight (sometimes slightly bent), sometimes slightly centrotylote, length: 142 – 610 Mm; width: 5 – 23 Mm. (c) Ortho- to dichotriaenes, rhabdome length: 252 – 4060 Mm (maximum length was measured by Hentschel, 1929); width: 22 – 145 Mm; orthotriaene clad length: 196 – 835 Mm; protoclad length: 60 – 520 Mm; deuteroclad length: 96 – 492 Mm. (d) Anatriaenes, rhabdome length: more than 6000 Mm; width: 17 – 43 Mm; clad length: 90 – 308 Mm. (e) Meso / protriaenes, rhabdome length: up to 4185 Mm; width: 17 – 36 Mm; clad length: 87 – 224 Mm; central clad length: 98 – 196 Mm. Microscleres: (f) sterrasters, usually spherical, some are very irregular, 56 – 102 Mm in diameter; thickness: 55 – 80 Mm; hilum: 12 – 20 Mm. Rosettes are made of 3 – 7 rays, covered with warts. Rosette diameter: 5 – 6 Mm. (g) Strongylasters to sphero-strongylasters, spiny, 4 – 22 Mm in diameter. (h) Oxyasters, spiny, with a more or less inflated centrum, with 4 – 20 rays, diameter: 10 – 62 Mm. DNA barcodes: GenBank accession nos. HM 592671, EU 442197 (Folmer COI): we have sequenced COI from specimens from northern Iceland (1), the Schultz Massive Seamount in the Greenland Sea (4), and the Davis strait (1): the Folmer COI is identical in all these specimens. No. EU 552083 (28 S, C 1 - D 2 domains): we have sequenced 28 S from two specimens from the Schultz Massive Seamount in the Greenland Sea, and no variation was observed. No. KC 481226 (18 S), obtained from UPSZMC 78042 (Schultz Massive Seamount). Distribution (Fig. 14): Geodia hentscheli is an Arctic species. The species has been recorded at depths of 130 – 2000 m, at temperatures of – 1.76 (eastern Greenland) to 4.5 ° C (west of Iceland and Reykjanes Ridge). The shallowest records (less than 200 m deep) come from the Canadian Ice Island at 81 ° N (Wagoner et al., 1989) and eastern Greenland (Burton, 1934; Koltun, 1964), the deepest records being off eastern Greenland, at temperatures of – 1.76 to 0.4 ° C. This species has not been found off Newfoundland. Biology: Budding seems to be fairly common in this species (Fig. 11 C). The isopod Caecognathia robusta (G. O. Sars, 1879) is a common epibiont living in the preoscule of this species (Barthel & Brandt, 1995). We have observed very few sponges living on the fur of G. hentscheli (e. g. Calcarea spp.). Distinctive characters: External morphology: the almost spherical form with one narrow preoscular cavity on top. Also, usually there is a thickening of the cortex just around the oscule, and there might be more or less high ridges between the small oscules (inside the preoscule). Spicules: on average, small size and ‘ bumpiness’ of the sterrasters, some sterrasters are very irregular, short and thick microxeas, large and sometimes irregular strongylasters. But one or all of these characters may be absent. Remarks: We examined specimen B 331 (ZMO) from East Greenland (137 m depth), identified as G. nodastrella by Burton (1934). A misidentification was suspected as G. nodastrella is a typical Lusitanian deep-sea species, never formerly described from arctic waters. B 331 (ZMO) is a small spherical specimen (8 mm in diameter) with two bundles of long spicules (mainly mesoprotriaenes) sticking out from it and a 0.5 - cm-thick cortex. The presence of one very small preoscule opening already suggests that this is not G. nodastrella (which has cribriporal pores and oscules, no preoscules). Furthermore, the spicules clearly match those of G. hentscheli. Burton (1934) was probably misled by the strongylasters which, in Greenland and Icelandic specimens, can become large sphero-strongylasters (up to 22 Mm in diameter in this specimen, larger than those measured in Table 3). We also examined the larger specimen B 330 (ZMO) from the same catch and identified by Burton (1934) as G. mesotriaena (now hentscheli); this identification is correct. In Table 3, the maximum size of oxyasters measured is 38 Mm. But we also examined more specimens and we found that oxyasters could reach sizes of 48 Mm (ZMBN 85205, Iceland, 604 m depth), 55 Mm (UPSZMC 78266, Davis Strait, 847 m depth) or even 62 Mm (PC 18, Iceland, 800 m depth). These large sizes of oxyasters are not mentioned by Koltun (1966). We also noted that the NEA specimens have oxyasters with fairly thin actines (2 Mm thick) whereas the three specimens from Davis Strait (PA 2010 - set 155) we examined have oxyasters with less numerous, thicker actines (up to 5 Mm thick). There is also quite Means are in bold; other values are ranges; N = 30 unless stated otherwise in parentheses, or unless measurements come from other studies. A dash indicates that this measurement is not given in the literature. n. f., not found; n. o., present but not observed in the sample in our possession, or broken. a lot of variation of the strongylasters (more so than in G. barretti). As we noted earlier, they can be fairly large and fairly irregular, and their actines can also be so small that they look like irregular spheres. We also noted that some specimens from Davis Strait (e. g. UPSZMC 78266) have particularly large spicules overall and a much thicker cortex (1 – 1.4 mm) than other specimens we examined (Table 3), probably because they are larger specimens (> 10 cm). The smaller specimen UPSZMC 78267 (Fig. 11 A), c. 3 cm in diameter, from the same station has a cortex of 0.55 mm. So cortex thickness may increase with size of specimens. Based on their morphology, G. barretti and G. hentscheli have previously been considered sister species (Koltun, 1966): their spicule and external morphologies are very similar so that they can be easily confused. But spicule measurements suggest that, on average, G. hentscheli has smaller sterrasters, thicker and shorter oxeas II, and larger somewhat more irregular strongylasters than G. barretti, but their ranges overlap so that these characters are not sufficient. Orthotriaenes are more common in G. hentscheli than in G. barretti. At all depths, G. hentscheli sterrasters are usually spherical whereas G. barretti sterrasters tend to become elongated in the NEA below 1000 m depth (but not in the NWA). Also, G. hentscheli sterrasters may have a bumpier surface than in G. barretti, due to slightly larger rosettes with more spines, but again, this character is not always present or easy to distinguish for a non-specialist. On the other hand, G. hentscheli sterrasters can often be irregularly developed (Fig. 13 F), and this is never observed in G. barretti. As for external morphology, confusion is still possible because G. barretti can sometimes have a subspherical shape with a narrow preoscule as well (Fig. 7 A). Genetically G. barretti and G. hentscheli are clearly different: 6 – 7 bp difference in the COI Folmer fragment, 8 bp difference in the 28 S (C 1 - D 2) fragment, and even 1 bp difference with 18 S.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFCF214BFF76CDE16A69B9B3.taxon	description	(FIGS 15 – 17, TABLE 4)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFCF214BFF76CDE16A69B9B3.taxon	materials_examined	Type locality and deposition of holotype: Collected by Robert McAndrew. Vikna Island (formerly called Vigten or Vikten Island), North Trøndelag, Norway, 183 m. The holotype specimen was not found (E. Sherlock, BNHM, pers. comm.); only type slides were examined: BNHM 1877.5. 21.1396 (surface and two spicule preparations) and BNHM 1877.5. 21.1398 (one section, one spicule preparation). External morphology and cortex: Young specimens are spherical (Fig. 15 E). The regular spherical body form persists during growth until the body reaches a size of about 10 cm in diameter; from that on the diameter is larger than the height, and our largest specimens are somewhat flattened, measuring 35 – 42 cm in maximum diameter and 20 – 24 cm in height (Fig. 15 A – C). Some mid-sized specimens from eastern Greenland and the Flemish Cap (Fig. 15 D) are unusually flat, the diameter measuring more than twice the height. The colour alive is whitish yellow to light grey; whitish in ethanol. The choanosome is yellowish in live specimens (Fig. 15 B); whitish in ethanol. Small specimens have smooth surfaces; larger ones develop a fur of long spicules, essentially on the sides (Fig. 15 A), which is frequently damaged during the catching procedure, so that large surface areas appear smooth. Cribriporal oscules (Fig. 15 F) are regularly scattered over the upper surface; each sieve is 0.5 – 1 mm in diameter. Cribriporal pores (Fig. 15 G) are scattered over the sides of the body; sieves are also 0.5 – 1 mm in diameter. The sterraster layer is very strong and tough, and usually 1 – 2 mm thick (occasionally up to 4 mm thick) (Fig. 15 H, I). The ectocortex with spheroxyasters and microxeas is very thin (45 – 100 Mm thick) compared with the endocortex (Fig. 15 H, I). Spicules (Fig. 16, Table 4): Megascleres: (a) oxeas, straight or bent, length: up to 14 mm; width: 5 – 106 Mm. (b) Microxeas, straight or slightly bent, rarely centrotylote, length: 220 – 445 Mm; width: 3 – 13 Mm. (c) Ortho- to dichotriaenes, orthotriaenes are more common, straight rhabdome, rhabdome length: 1650 – 9625 Mm (maximum from Brøndsted, 1932); width: 40 – 165 Mm. Clads can often end with a small downward bend, orthotriaene clad length: 210 – 1125 Mm; protoclad length: 220 – 500 Mm; deuteroclad length: 70 – 450 Mm. (d) Anatriaenes, common swelling on top of the cladome, rhabdome length up to 22 mm; width: 3.5 – 63 Mm; clad length: 24 – 285 Mm. (e) Meso / protriaenes, rhabdome length: up to 12 mm; width: 5 – 79 Mm; clad length: 32 – 330 Mm; central clad length: 49 – 685 Mm. Microscleres: (f) sterrasters, spherical to subspherical, 124 – 360 Mm in diameter; thickness: 128 – 230 Mm; hilum diameter: 20 – 30 Mm. Rosettes are made of 5 – 7 warty rays; rosette diameter: 6 – 10 Mm. (g) Spheroxyasters, rough actines (difficult to see with the optical microscope), with centrum more or less developed, diameter: 4 – 18 Mm. (h) Oxyasters, thin rough actines, diameter: 10 – 88 Mm. DNA barcodes: GenBank accession nos. EU 442198, HM 592689, HM 592696 (Folmer COI): we have sequenced COI from specimens from western and northern Norway (4), Spitsbergen (2), Davis Strait (1), and Flemish Cap (1): the Folmer COI is identical in all these specimens. No. EU 552082 (28 S, C 1 - D 2 domains): we have sequenced 28 S from the Bergen area (2), and Spitsbergen (1), 1 - bp difference was observed between the two Bergen specimens. No. KC 481225 (18 S), obtained from ZMBN 89717 (Spitsbergen). Two specimens from Spitsbergen sequenced: no variation was observed. Distribution (Fig. 17): We characterize the species as northern boreal, with the ability to invade some neighbouring cold-water areas, but not penetrating into real Arctic conditions. It has been recorded at depths from 157 m (Trondheimsfjord) to 1900 m (eastern Greenland). The temperature range is – 0.82 ° C (north of the Faroe Islands, BIOFAR st. 901) to 8.3 ° C (south-west of the Faroe Islands, BIOFAR st. 69). Most eastern records are from between 230 and 400 m, at temperatures of 5 – 8 ° C. Occurrences at temperatures below 2 ° C were essentially found in the Denmark Strait (282 – 467 m depth). The records on the southern flanks of the Bill Bailey and Faroe Banks came from rather deep water, 1140 and 650 m, respectively. They hardly represent the southern distribution limit of the species but could indicate that off the Shetland Isles and Scotland it can be expected to occur on the upper slope. Brøndsted (1932) and Koltun (1966) mentioned the Shetlands as part of the distribution area; although this is to be expected, there is so far no proof, the record being a mistake by Brøndsted for the Norwegian record of the type specimen. Koltun (1966) mentioned occurrences in the south-western Barents Sea and the Denmark Strait, but gave no detailed information. Biology: We found no indications of asexual reproduction. The predator chiton H. nagelfar and the parasitic foraminiferan H. sarcophaga have been found living on G. macandrewii (Warén & Klitgaard, 1991; Cedhagen, 1994; Todt et al., 2009). More associated fauna has been investigated by Klitgaard (1995). The chemistry (elemental analysis, amino acids, sterols, and quaternary ammonium compounds) has been investigated (Kingston et al., 1979; Hougaard et al., 1991 a, b). Note that G. macandrewii off the Labrador coast (Canada) and from the Faroe Islands have very similar sterol composition (Kingston et al., 1979; Hougaard et al., 1991 b). Distinctive characters: External morphology: the regularly round, almost spherical form with no conspicuous openings, the hard consistency, the uniform distribution on the top side of cribriporal oscules, and the very thick cortex. Spicules: very large sterrasters (124 – 360 Mm in diameter). Remarks: It was named after Robert MacAndrew who collected this species. Spicule measurements of this species are scarce apart from Sollas (1888), Brøndsted (1932), and Koltun (1966). Although this is a fairly common species in some areas, it is rarely mentioned in the literature compared with G. barretti or G. phlegraei, perhaps because it is often confused Means are in bold; other values are ranges; N = 30 unless stated otherwise in parentheses, or unless measurements come from other studies. A dash indicates that this measurement is not given in the literature. n. f., not found. with these species. In spicule preparations, it can be difficult to separate the largest ectocortical spheroxyasters from the smallest choanosomal oxyasters as there is no clear transition from one form to the other, so we recommend measuring them on a section, to ensure not to mix both categories (this was done for the holotype and ZMBN 77924). There is some variation within oxyasters: small (up to 25 – 28 Mm) with thin actines (1 – 2 Mm thick) usually in specimens from shallower depths (183 – 600 m), to very large oxyasters (40 – 88 Mm) with thick actines (4 – 7 Mm thick) in specimens living deeper than 1000 m. The large sizes are not mentioned by Koltun (1966). The large dichotriaenes with atypical forward orientated cladomes represented by Bowerbank (1872 a: plate X, fig. 4) were indeed observed on the type slide number ‘ Bk. 1398 ’. However, in our opinion, they result from a contamination from another Astrophorina (probably Stelletta normani Sollas, 1880 a, a common species on the Norwegian coast at these same depths). Molecular phylogenetic studies suggest that G. macandrewii belongs to Cydoniump along with G. cydonium, G. conchilega, and Geodia papyracea Hechtel, 1965 (Fig. 2). G. macandrewii shares the possession of identical cribriporal oscules and pores with G. cydonium and G. conchilega; G. papyracea, by contrast, has uniporal oscules (Cárdenas et al., 2009). We could nonetheless suggest that the association of cribriporal oscules and oscules could be an autapomorphy of the clade, and that the cribriporal oscules were later modified and lost in G. papyracea. These four species also share spiny euasters but spiny euasters are also present in the neighbouring Geodia clades (Depressiogeodiap and Geodiap) (Fig. 2). Geodia macandrewii, G. cydonium, and G. papyracea share orthotriaenes but G. conchilega has dichotriaenes. So additional sampling is needed to better understand the Cydoniump clade and find morphological apomorphies that would support it. No spicule differences were observed between NWA and NEA specimens. Sterrasters are usually subspherical but we noted that one specimen from Davis Strait (UPSZMC 78255) had some atypically shaped sterrasters, most of them ‘ lemon-shaped’, but other specimens from the same locality had ‘ normal’ sterrasters.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFCF214BFF76CDE16A69B9B3.taxon	materials_examined	We examined slides (74 - 8 - 27.1) made by H. M. Reiswig from material published under the name Geodia megastrella and collected off the Labrador coast (53 ° 24 ′ 50 ″ N, 52 ° 15 ′ 00 ″ W) at 732 m depth (Kingston et al., 1979). We observed a thick cortex (2 mm thick), orthotriaenes, common ana / protriaenes, very large sterrasters (up to 360 Mm), and spiny oxyasters with thin actines (10 – 30 Mm). This leaves no doubt that this is in fact G. macandrewii.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFF52140FC0ACCD36B21BB64.taxon	description	(FIGS 18 – 20, 23, TABLE 5)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFF52140FC0ACCD36B21BB64.taxon	materials_examined	Type locality and type material examined Isops phlegraei, Korsfjord near Bergen, Norway. 60 ° 10 ′ N, 05 ° 10 ′ E, 330 m. Collected by Rev. A. M. Norman in 1878, BNHM 1910.1. 1.840.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFF52140FC0ACCD36B21BB64.taxon	description	Synops pyriformis, near Hammerfest, Norway, 71 ° 12 ′ 5 ″ N, 20 ° 30 ′ 5 ″ E, 247 m, Willem Barents Exp. 1878 – 79, RMNH Por. 660 (wet specimen); MNHN- DN 23, Norman Collection, spicule preparation and section; BMNH 10.1.1.1147 – 1148, Norman Collection, spicule preparation and section (not seen). Isops sphaeroides, near Hammerfest, Norway, 71 ° 12 ′ 5 ″ N, 20 ° 30 ′ 5 ″ E, 247 m, Willem Barents Exp. 1878 – 79, MNHN-DN 24, syntype 1, Norman Collection, spicule preparation and section; RMNH, Vosmaer slide collection, box number 37, syntype 2, two spicule preparations and one stained thin section; BMNH 10.1.1.1158 – 1159, Norman Collection, two slides (not seen). Isops pallida, near Hammerfest, Norway, 71 ° 12 ′ 5 ″ N, 20 ° 30 ′ 5 ″ E, 247 m, Willem Barents Exp. 1878 – 79, RMNH Por. 652, wet specimen (seen only from pictures); RMNH, Vosmaer slide collection, box number 37, two spicule preparations with number 62. External morphology and cortex: More or less spherical, the largest specimens somewhat flattened on the top, cup-shaped, generally up to 20 cm in diameter, and 15 cm high (Fig. 18 A, B, E); our largest specimen measures 43 cm in maximum dimension. Juveniles are spherical (Fig. 18 C). Specimens growing on vertical cliffs (i. e. fjords) can be flattened or more irregular (Fig. 18 D). Lower part sometimes formed as a base, with root-like outgrowths (Fig. 19 B). The colour alive usually varies from whitish grey to light brown, to slightly rose. One shallow fjord specimen (PC 111, shallower than 175 m) was faintly purple in ethanol, like shallow G. barretti and Pachymatisma normani Sollas, 1888 (a NEA boreal Geodiidae) (Cárdenas & Rapp, 2013). Other specimens fixed in ethanol seemed to be reddish, although this staining comes from the epibiont yellow sponge Hexadella dedritifera Topsent, 1913 (Fig. 18 A, F) which turns dark red during the fixation. Many specimens are very hispid on the sides, but never on the top surface (Fig. 18 A – C). The fur can be up to 10 mm long, but is not a regularly occurring feature in dredged specimens as it may be easily lost during the collection. The cortex (Fig. 18 I) is thin to fairly thick (0.7 – 2 mm thick); it is usually difficult to cut and tends to break in pieces. Many epibionts (especially sponges) are present in this hispid part. Uniporal oscular openings are up to 1 mm in diameter, and are found mainly on the upper surface (Fig. 18 A – F). Oscular openings may be at the tip of thick conical elevations, more or less pointy, which can be lighter-coloured (Fig. 18 G) (but not always) and which gave the name to this species: Campi phlegraei is a local name of the Naples volcanic area (Sollas, 1880 b). Uniporal pores (c. 300 – 400 Mm in diameter) (Fig. 18 H – - J) are scattered on the sides and partly on the underside of the body. Pores are usually not elevated but can also be surrounded by a white margin. Description of type material: The holotype of G. phlegraei is a small subspherical specimen (diameter: 2.5 cm) cut up into four parts (Fig. 19 A); it has conical-shaped oscules. There is also one BMNH slide Means are in bold; other values are ranges; N = 30 unless stated otherwise in parentheses, or unless measurements come from other studies. A dash indicates that this measurement is not given in the literature. n. f., not found; n. o., not observed in the specimen in our possession. N, Norway; S, Sweden. of the type but it is damaged and the embedding medium has blackened. Figure 20 shows SEM pictures of the spicules from the holotype. The type of S. pyriformis is a medium-sized specimen (length: 10 cm, width: 8 cm) cut into five pieces. The main piece (Fig. 19 B) is the elongated cup-shaped half represented by Vosmaer (1882: plate IV). We have only seen pictures of the wet specimen of the lectotype (specimen a) of I. pallida (Fig. 19 C, D): the pearshaped specimen is cut into two fragments (one-half and one-third of a single specimen). The half fragment is about 6 ¥ 4 cm with a thick cortex (1 – 2 mm thick), uniporal oscules, and uniporal pores. There are also two spicule preparations of the lectotype (slide 62: a) of I. pallida (Fig. 4 A). One slide is broken so the label is gone but the spicules are identical to those of slide 62. Spicules (Fig. 20, Table 5): Megascleres: (a) oxeas, length: 1173 – 7600 Mm; width: 10 – 100 Mm. (b) Orthotriaenes, rare dichotriaenes, rhabdome length: 586 – 6655 Mm; width: 12 – 150 Mm; orthotriaene clad length: 80 – 1125 Mm; protoclad length: 220 – 250 Mm; deuteroclad length: 100 – 250 Mm. (c) Anatriaenes, rare (some clads were dichotomized in the type of I. sphaeroides) rhabdome length: up to 11 mm; width: 8 – 25 Mm; clad length: 30 – 130 Mm (minimum according to Koltun, 1966). (d) Protriaenes, very rare, rhabdome length: 11 600 – 12 750 Mm; width: 25 – 30 Mm; clad length: 190 – 200 Mm. Microscleres: (e) sterrasters, subspherical (NEA specimens) or spherical (in some NWA specimens), length: 82 – 144 Mm; width: 70 – 124 Mm; thickness: 65 – 80 Mm; hilum: 12 – 15 Mm. Rosettes are made of 4 – 12 smooth rays; rosette diameter: 6 – 8 Mm. (f) Spherasters with spiny conical actines (more rarely with blunt ends), 8 – 26 Mm in diameter. (g) Oxyasters, smooth (rough actines were rarely observed in very large oxyasters), 10 – 70 Mm in diameter (maximum is from measurements of the type by Sollas, 1888). DNA barcodes: GenBank accession nos. EU 442196, HM 592701 (Folmer COI). We have sequenced COI from specimens from Spitsbergen (1), western and northern Norway (10), Mingulay Reef (1), and Orphan Knoll (1): the Folmer COI is identical in all these specimens. No. KC 481222 (18 S), obtained from ZMBN 77929 (Korsfjord, Norway). 18 S of ZMBN 89719 (Spitsbergen) was also sequenced: no variation was observed. Distribution (Fig. 23): Geodia phlegraei has a boreal distribution and seems to avoid arctic waters; it can be found from 40 m (Trondheimsfjord) to 3000 m (Orphan Knoll). It is commonly found at depths from 100 – 300 m (Norwegian continental shelf) to 725 m (Faeroes), at temperatures of 0.3 ° C (BIOICE, st. 2926) to 7.9 ° C (BIOFAR, st. 297). It has also been recorded by divers in Norwegian fjords at shallower depths: for example, it has been photographed by A. Salesjö in the Trondheimsfjord at only 40 – 50 m depth (http: // www. anderssalesjo. com /? id = 3306 & lang = 42, accessed 7 May 2013). We also identified it in material collected in Mingulay Reef (western Scotland) at 128 – 139 m depth. In Mingulay reef (R. van Soest, pers. comm.) and Norwegian fjords at shallow depths, temperatures can reach 10.5 ° C. The specimen identified as G. phlegraei from Rockall Bank (van Soest et al., 2007) was actually a Geodia cf. nodastrella. Interestingly, all but one of the NWA specimens from the Flemish Cap that we examined (a total of eight specimens) collected during NEREIDA 2009 – 10 and originally identified as G. phlegraei were in fact G. parva. The only specimen from the Flemish Cap that we identified as G. phlegraei (DR 24 - 69 d = UPSZMC 78280) has an external morphology similar to G. phlegraei and large elongated sterrasters (c. 132 – 136 Mm in diameter). Unfortunately, we could not get a COI sequence from it to confirm this identification. However, we did get a COI sequence for R 1340 - 04 (= UPSZMC 78308) confirming it was G. phlegraei: it has large spherical sterrasters (88 – 107 Mm in length) and it was collected at 3000 m depth at Orphan Knoll, where the temperature was 2.4 ° C (Fig. 18 F shows R 1341 - 18 collected at 2347 m in the same area). The most western specimen of G. phlegraei found is from western Greenland (‘ Shinkai Maru’, st. 32, 64 ° 13.5 ′ N, 54 ° 42.1 ′ W. 970 m) (Fig. 18 G, H), but this material was not suited for molecular studies. Biology: When he described this species, Sollas (1880) immediately noticed that it was ‘ covered by various foreign bodies’. Indeed, G. phlegraei is the boreoarctic Geodia which is most often found covered with epifauna, especially sponges and including individuals of its own species. Here are a few sponges found on G. phlegraei: Craniella sp., Cyamon spinispinosum (Topsent, 1904), Hexadella dedritifera (mis-identified as Aplysilla sulphurea in Klitgaard (1995 )), Leucandra spp., Lissodendoryx (L.) fragilis Fristedt, 1885, Polymastia grimaldii (Topsent, 1913), Stelletta normani, Ute gladiata Borojevic, 1966, etc. G. phlegraei can settle on other sponges as well such as other large Astrophorina [S. normani, Stryphnus fortis (Vosmaer, 1885)]. The predator chiton H. nagelfar and the parasitic foraminiferan H. sarcophaga have been found living on G. phlegraei (Warén & Klitgaard, 1991; Cedhagen, 1994; Todt et al., 2009). More associated fauna has been investigated by Klitgaard (1995). The chemistry (elemental analysis, amino acids, sterols, and quaternary ammonium compounds) has been investigated by Hougaard et al. (1991 a, b). The associated microsymbionts of G. phlegraei collected in the Sula Ridge reef (Norway) have been studied (Graeber et al., 2004; Dieckmann et al., 2005) and led to the isolation and description of a new gammaproteobacterium (Oceanospirillales group): Spongiispira norvegica (Kaesler et al., 2008). We observed on the type section of I. sphaeroides made by Vosmaer many subglobular oocytes without pseudopodes. This specimen has been collected in the Barents Sea on 2 July 1879. In the NEA, G. phlegraei can easily be confused with another Geodiidae, Pachymatisma normani, which also has raised white-rimmed uniporal oscules, but P. normani has microrhabds in the cortex, instead of spheroxyasters. Geodia phlegraei can also be easily confused with its sister species G. parva, and in that case only spicule and genetic characters can differentiate them (cf. below). Distinctive characters: External morphology: round to sometimes cup-shaped, with only uniporal openings (i. e. no sieve). The numerous small oscules on the top of specimen, each with a whitish rim making it look like a little wart. Often overgrown with other sponges, hydrozoa, bryozoa, etc. Spicules: large spherasters, smooth oxyasters along with fairly large sterrasters (70 – 144 Mm). Remarks: Choanosomal oxyasters can sometimes be separated into two size categories (10 – 25 and 45 – 60 Mm) but in some specimens we have more of a continuum of sizes so we decided to treat oxyasters as one category (Table 5). Koltun (1966) notes the presence of rare small slightly curved oxeas (230 – 420 ¥ 8 – 10 Mm); we never observed those and wonder whether these could have been contamination. Dichotriaenes have been reported by Koltun (1966) and Vosmaer (1882), but it should be stressed here that they are fairly rare as we only found a few (notably in the holotype). Anatriaenes are rare and we report for the first time the presence of protriaenes in this species (found in the type of I. sphaeroides), but they seem to be very rare.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFF52140FC0ACCD36B21BB64.taxon	description	We also compared type specimen and type slides of S. pyriformis with our material. The cortex thickness of S. pyriformis (1.1 – 1.3 mm) agrees well with G. phlegraei (Table 5). Spicule measurements also match those of G. phlegraei (Table 5). Finally, observation of the external morphology (Fig. 19 B) and new spicule preparations from the wet type of S. pyriformis further confirmed this. All in all, we therefore follow Burton (1930) and confirm the synonymy for S. pyriformis, I. pallida, and I. sphaeroides. Koltun (1966) had already underlined the morphological variability of G. phlegraei. He notably states that in deeper and colder waters the morphology of G. phlegraei is somewhat different. Klitgaard & Tendal (2004) also noticed this arctic water morphotype and considered it as a subspecies of G. phlegraei: G. phlegraei pyriformis. After having examined many specimens from the whole boreo-arctic area, we confirm the existence of different morphotypes, and, after incorporating additional morphological data as well as molecular data, it was decided that the arctic morphotype represented a valid species which had in fact been previously described under the name Geodia parva Hansen, 1885, before being synonymized with G. phlegraei (Burton, 1930). Below, we resurrect and redescribe G. parva.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFFE2145FC3CCEA66E68B94E.taxon	description	(FIGS 21 – 23, TABLE 5)	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFFE2145FC3CCEA66E68B94E.taxon	materials_examined	Type material examined Geodia parva, unknown station, Norwegian North Sea Exp. 1876 – 78, ZMBN 100 (wet specimen). External morphology and cortex: It seems there are two morphotypes for this species. The first morphotype can be found in true Arctic waters (Norwegian Sea, Greenland Sea, Davis Strait) – it is very characteristic and different from G. phlegraei; this morphotype is the one described below. The second morphotype, henceforth called the mixed-water mor- photype, is found in areas where Arctic and Atlantic waters mix (Denmark Strait, Flemish Cap) and has a mix of characters of the Arctic morphotype and G. phlegraei (cf. Discussion for its description). Arctic morphotype: Juveniles are spherical. Larger specimens are more or less spherical, flattened and cupshaped (Fig. 21 A – C); the largest specimens are 26 cm in diameter (Klitgaard & Tendal, 2004), so G. parva reaches smaller sizes than G. phlegraei. Specimens from the Schultz Massive Seamount (Biodeep and H 2 deep Expeditions) can be fairly irregular (Fig. 21 D). Root-like structures at the base are fairly common. Budding is commonly observed, and the buds are string like (Fig. 21 D) to more massive (e. g. club-shaped). Most specimens have a dense fur covering the whole body, even the top surface where the oscules are found (Fig. 21 A, B). In dredged specimens, this fur is often lost except in sheltered folds (Fig. 21 E). Colour alive is whitish to light brownish. With the openings usually lighter-coloured this gives the sponge a characteristic mottled appearance. The cortex is very thin to thin (0.15 – 0.7 mm thick) (Fig. 21 G, H), flexible, and easily cut (no breakage in large pieces as in G. phlegraei). Many epibionts (e. g. sponges) are present in this hispid part (Fig. 21 B). Uniporal oscular openings are up to 1 mm in diameter, and are found mainly on the upper surface (Fig. 21 E). Oscular openings are often wide and surrounded by a white rim (more rarely conical elevations), as in many specimens of G. phlegraei. Uniporal pores (up to 1 mm in diameter) (Fig. 21 F) are scattered on the sides and partly on the underside of the body. Pores can be very slightly elevated and are usually surrounded by a white margin. Description of holotype: The type material of G. parva is composed of one very small spherical sponge (4 mm in diameter) and a small piece of cortex of another specimen (3 mm). This explains the name given to this species: ‘ parva’ means ‘ small’ in Latin. The small piece of cortex has been completely used for spicule and SEM preparation (Fig. 22 E, F), and the resulting slides and SEM stub are now stored at the ZMBN under the same number. Spicules (Fig. 22, Table 5): Megascleres: (a) oxeas, sometimes modified to styles, length: 773 – 7935 Mm; width: 14 – 102 Mm. (b) Orthotriaenes, dichotriaenes are fairly rare, rhabdome length: 360 – 5395 Mm; width: 20 – 108 Mm; orthotriaene clad length: 102 – 1008 Mm; protoclad length: 56 – 405 Mm; deuteroclad length: 44 – 521 Mm. (c) Anatriaenes are fairly rare, rhabdome length: 1 – 16 mm (minimum length measured by Hentschel, 1929); width: 15 – 34 Mm; clad length: 29 – 78 Mm. (d) Meso / protriaenes, very rare, clades are slightly forward or even slightly backward, with or without a central clad, rhabdome length: 840 – 8371 Mm; width: 9 – 68 Mm; clad length: 36 – 161 Mm; central clad length: 60 – 334 Mm. Microscleres: (e) sterrasters, spherical with a ‘ bumpy’ appearance and commonly irregular (Arctic morphotype), usually spherical, but also sometimes elongated, similar to the sterrasters of G. phlegraei (mixed-water morphotype), 56 – 104 Mm in diameter (Arctic morphotype); length: 68 – 124 Mm (mixed-water morphotype); thickness: 52 – 56 Mm; shallow and large hilum: 16 – 25 Mm. Rosettes are made of 10 – 15 piled smooth rays. Rosette diameter: 6 – 8 Mm. (f) Spherasters with fairly spiny actines which look almost blunt under an optical microscope and commonly irregular (Arctic morphotype), spherasters with less spiny actines, which look more conical and pointy under an optical microscope as in G. phlegraei (mixed-water morphotype), 10 – 30 Mm in diameter. (g) Oxyasters, smooth or more rarely slightly rough actines, 10 – 72 Mm in diameter. DNA barcodes: GenBank accession no. HM 592690 (Folmer COI). We have sequenced COI from specimens from Spitsbergen (1), the Schultz Massive Seamount in the Greenland Sea (3), the Davis Strait (1), the Flemish Cap (2), and Orphan Knoll (1): the Folmer COI is identical in all these specimens. No. KC 481223 (18 S), obtained from ZMBN 85210 (Schultz Massive Seamount). Distribution (Fig. 23): Arctic distribution. It has been found at depths from 100 m [Canadian Ice Island (Wagoner et al., 1989)] to 2747 m (Orphan Knoll), at temperatures of – 1.5 ° C (Wagoner et al., 1989) to 4.4 ° C (Ingolf Exp. st. 90). Biology: Associated fauna has never been closely investigated but, as in G. phlegraei, our observations suggest that many macrosymbionts grow on its fur, especially sponges (Fig. 20 B): Hexactinellida, Hexadella dedritifera, Asbestopluma (A.) lycopodium (Levinsen, 1887), Craniella infrequens (Carter, 1876), etc. H. nagelfar (chiton) and the parasitic foraminiferan H. sarcophaga have not been observed on G. parva, but these two species do not actually thrive in cold waters. Distinctive characters: External morphology (Arctic morphotype): bumpy / wrinkled surface and thin flexible cortex (c. 0.5 mm). Budding. Hispidity all over the sponge and overgrown with other sponges, etc. Spicules: spherical small sterrasters (56 – 92 Mm), some irregular sterrasters. Remarks: Koltun (1966) noted a different G. phlegraei morphotype of the Norwegian Sea, the Greenland Sea, and the central part of the Arctic Ocean. They are smaller, brighter in colour, with a thinner cortex (0.5 – 0.9 mm) and their spherasters have blunt rays (instead of being pointy). Klitgaard & Tendal (2004) also recognize this morph and consider the boreal and arctic form to be subspecies by calling them Isops phlegraei phlegraei and I. phlegraei pyriformis (arctic subspecies). They in fact sometimes occur in the same catch, in the hydrographically mixed regions of the Denmark Strait (Stations 78, 90, and 92 of Ingolf Exp.), the south-western Barents Sea, and at Orphan Knoll (Fig. 23). As we have shown above, I. pyriformis is a synonym of I. phlegraei and is therefore not an available name. On the other hand, a re-examination of the type material of G. parva (Table 5, Fig. 22) showed that it belonged to the arctic population. The most obvious differences between G. phlegraei and G. parva are that G. parva show budding and have a thin flexible cortex which gives a characteristic bumpy wrinkled surface appearance. We have never seen buds in G. phlegraei, and its cortex is thick and stiff. Koltun (1966) also notes that oxeas are often modified to styles in G. phlegraei; we have observed this, but only in G. parva. The presence of irregular sterrasters is fairly common in G. parva (Fig. 22 F) but never observed in G. phlegraei. Furthermore, the 1 - bp difference (position 370, A in G. phlegraei, C in G. parva) between the COI of the two species is consistent. It reflects the close phylogenetic relationship of these species but also suggests that they may have completely diverged. 18 S being far more conserved than the Folmer COI marker, we observed no differences between the 18 S of G. phlegraei and G. parva. We therefore gather here enough morphological and molecular evidence to upgrade these two subspecies to two sister species: G. phlegraei and G. parva (here officially resurrected). We remain troubled by the specimens collected in areas where Atlantic and Arctic waters mix (Denmark Strait, Flemish Cap), which essentially includes specimens from St. 90 (Ingolf Exp.) and from the NEREIDA campaign off Newfoundland. Using the COI marker, specimens from the Flemish Cap were identified as G. parva. However, their external morphology may in some occasions be closer to G. phlegraei [thicker cortex, up to 1.8 mm (UPSZMC 78279), oscules with conical elevations, smooth surface]. Their spicules make us also think of G. phlegraei (larger sterrasters and regular less spiny spherasters). Without a molecular marker they are almost impossible to identify for some of them. The status of these populations is therefore questioned: these could be G. parva populations in different environmental conditions (mixing of waters) or G. phlegraei / parva hybrids. Faster evolving genetic markers are clearly needed to settle this matter. The specimens identified by Hentschel (1929) as I. pyriformis were collected in arctic deep waters (1000 m depth) along with typical arctic species (e. g. G. hentscheli, Stelletta rhaphidiophora Hentschel, 1929). Furthermore, sterraster measurements (81 – 91 Mm) fit well with those of G. parva (Table 5). Pictures of the specimens described by Hentschel (ZMB Por 7542, 7543, 7544, and 8420, courtesy of C. Lueter) confirm that these are G. parva. Also, Koltun (1964) records G. phlegraei from the Greenland Sea (Obb, 1956, st. 7, 1441 m, - 0.4 ° C), south-west of Spitsbergen (Lena, 1958, st. 2, 759 m, ca 0.65 ° C) and north-west of Franz Josef Land (F. Litke, 1955, st. 26, 415 m, 0.4 ° C). We suppose that G. phlegraei specimens from station 7 (Obb, 1956) are G. parva because only typical Arctic species were collected at this deep station with negative temperature: Craniella infrequens, Stelletta rhaphidiophora, and Thenea abyssorum Koltun, 1964. Concerning stations 26 and 2, we cannot be sure. The phylogenetic position of the G. phlegraei + G. parva clade in the Geodiinae is still very uncertain, except that it does not belong to the three wellsupported clades Cydoniump, Depressiogeodiap, or Geodiap (Fig. 2). Contrary to the Cydoniump and Depressiogeodiap clades which, for the time being, only include Atlantic species, the G. phlegraei + G. parva clade forms a well-supported clade with Geodia intermedia (Wiedenmayer, 1989) from Southern Australia.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFFB2145FE94CCB66CB2B9F1.taxon	description	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.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
122687EBFFFB215AFC21CD2A6D8BBFF9.taxon	materials_examined	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.	en	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. Zoological Journal of the Linnean Society 169 (2): 251-311, DOI: 10.1111/zoj.12056, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12056
