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Myxidium B��tschli, 1882, Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on page 547, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Pseudalataspora Kovaleva & Gaevskaya, 1983, Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on page 543, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Palliatus Schulman et al. 1979, Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on page 551, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: http://dx.doi.org/10.11646/zootaxa.3647.4.4

Pseudalataspora kovalevae n. sp. Material studied Type host: Macruronus magellanicus L��nnberg, 1907 Site of infection: gall bladder Localities, dates and depths: (1) off southern Chile, 55 �� 30��S, 71 �� 30��W, October 2007, 350m; (2) off Chiloe Island, Chile, 43 �� 0 0��S, 73 �� 0 0��W, June 2007, 300m; (3) west of Falkland Islands, 51 �� 00��to 52 �� 30��S, 62 �� 00��to 62 �� 30��W, October 2007, 200��� 250m; (4) west of Falkland Islands, 51 �� 00��to 52 �� 30��S, 62 �� 00��to 62 �� 30��W, October 2009, 200��� 250m. Type locality: (1). Prevalence: (1) 40 % (4 of 9); (2) 64 % (16 of 23); (3) 23 % (7 of 30); (4) 57 % (24 of 42). Host length range: 25���42 cm. Collection numbers: NHMUK 2012.3. 19.1, 2012.3. 19.2, 2012.3. 19.3. Morphological description. Sporoblast globular, disporic (Fig. 1). No pseudopodia observed. Ectoplasm dense and clearly demarcated from fine endoplasm. Dimensions, based on 10 fixed specimens: 28.4���38.4 x 28.0��� 36.0. Spore (Figs. 2���6) triangular with sharply pointed tips in side view, flat anteriorly and curved in apical view. Sutural line prominent and raised. Sporoplasm deeply staining and binucleate. Valves drawn out into two delicate broad alate processes joined together at their proximal extremities to form a parachute-like structure over the valves. Polar capsules spherical, close together. Polar filament thick, number of coils not visible. Dimensions, based on 30 fixed spores, as ranges with means �� SD in parentheses: spore length 8.0��� 10.5 (9.1 �� 0.68); spore thickness, including alate processes 30.0��� 35.5 (31.8 �� 1.52), excluding alate processes 8.8���11.6 (9.0 �� 1.2); spore width 14.0���20.0 (15.7 �� 1.57); polar capsule diameter 2.5 ���3.0 (2.7 �� 0.16); polar capsule length: spore length = 1: 2.8���3.8; spore length: spore width = 1: 3.0��� 4.9. Molecular results: Pseudalataspora kovalevae. PCR amplification of the myxosporean 18 S rRNA gene from four infected gall bladders from sample (4) above generated a product of approximately 1650 nucleotides in both of the samples processed from each fish. Forward and reverse 18 S rRNA gene sequences were obtained for PCR products from both samples from each individual host, and internal sequences from a single sample from each host. All sequences were identical. Two polymorphic positions were found. A final sequence of 1594 nucleotides was submitted to Genebank under Accession No. JX 467675. The sequence was novel with respect to previously generated data from myxosporean species. Pseudalataspora kovalevae 18 S rRNA gene sequence showed closest sequence identity to Ceratomyxa anko Freeman et al., 2008 (DQ 301510) and Ceratomyxa pantherini Gunter et al., 2010 (GU 136393) based on BLAST analysis with 92 % and 97 % identity over 92 % and 85 % of its sequence respectively. Following removal of gaps and ambiguous bases, 1034 nucleotides remained for phylogenetic analyses. Cystodiscus melleni (Jirku et al., 2006) was chosen as the outgroup based on its position in the distance tree generated following initial BLAST analyses of the P. kovalevae sequence. MP and ML trees showed similar topologies, P. kovalevae grouping with C. pantherini and C. anko in both cases, with strong bootstrap support (100) (Fig. 7). Discussion. Twelve species of the genus Pseudalataspora have been described from the gall bladders of marine fishes but none has been sequenced to date. In addition, we were made aware of an unpublished manuscript discovered in the late Professor Kovaleva���s laboratory in Kaliningrad, which includes the morphological description of Pseudalataspora pacifica Kovaleva and Grudnev, a new species found in the gall bladder of M. magellanicus caught off the coast of Chile in 1983. The authors described the new species as being similar to P. umbraculiformis, originally described by Gaevskaya and Kovaleva (1984) from the gadid fish Gaidropsarus mediterraneus (L., 1758) in the Northeast Atlantic. However, they considered the two species to differ sufficiently in certain features to be considered different species. The main differences they identified were in the shape and size of the spores, the diameter of the polar capsules, and the number of coils in the polar filament. Because the description of P. pacifica remains unpublished, it cannot be considered to be a valid species, but in recognition of Professor Kovaleva���s discovery we decided to name our new species after her. Table 1 compares P. kovalevae with both P. umbraculiformis and P. pacifica. Pseudalataspora kovalevae is from the same host and locality as P. pacifica and these two species are clearly more similar in morphology than either is to P. umbraculiformis. Gunter et al. (2009) suggested that the Ceratomyxa clade currently contains a number of as yet undifferentiated genera and that additional sampling and further morphological and genetic data are needed to resolve the divisions within the group. As previously mentioned, there are currently no sequence data deposited in GenBank for any species of Pseudalataspora, and the new P. kovalevae 18 S rRNA gene sequence grouped most closely with some Ceratomyxa species. Morphologically the spores of these two genera are very similar, with the presence or absence of delicate alate processes being the main distinguishing feature between them. The morphological and molecular similarities between these genera therefore suggest that they are closely related, but they are currently classified in different families. The alate processes are fragile and not always easily observed, which raises the possibility that some species currently assigned to the genus Ceratomyxa may turn out on closer inspection to belong in Pseudalataspora. Feature P. umbraculiformis P. ��� pacifica ���(*) P. kovalevae Plasmodium shape Club-shaped or spherical, Spherical, disporous Spherical, disporous disporous Plasmodium dimensions 18.0���21.0 29.28���32.28 28.4���38.4 x 28.0���36.0 Spore shape Wedge-shaped, expanded Rounded tops, sharply Triangular with pointed anteriorly, narrowed curved anteriorly extremities in side view; curved posteriorly and flat anteriorly in apical view Spore length 8.0��� 9.3 7.98���12.63 8.0��� 10.5 Spore thickness with alate 14.6���17.3 27.93���35.91 30.0��� 35.5 processes Spore thickness without alate 6.7 ���8.0 Not given 8.8���11.6 processes Spore width 14.6���17.3 Not given 14.0���20.0 Sutural line Clear and straight Clear and straight Prominent, raised Polar capsule diameter 2.7 3.0- 3.32 2.5 ���3.0 No. of polar filament coils 5 6 Number not observed, Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on pages 543-547, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Myxidium baueri Kovaleva and Gaevskaya, 1982 Material studied Host: Macruronus magellanicus L��nnberg, 1907 Type hosts: Macrourus holotrachys G��nther, 1878 and Merluccius hubbsi Marini, 1933 Site of infection: gall bladder Localities, dates and depths: (1) off Chiloe Island, Chile, 43 ��S, 73 ��W, June 2007, 300m; (2) west of Falkland Islands, 51 �� 00��to 52 �� 30��S, 62 �� 00��to 62 �� 30��W, October 2007, 200��� 250m; (3) west of Falkland Islands, 51 �� 00��to 52 �� 30��S, 62 �� 00��to 62 �� 30��W, October 2009, 200��� 250m. Type locality: (1). Prevalence: (1) 4 % (1 of 23); (2) 13 % (4 of 30); (3) 7 % (3 of 42). Host length range: 25���42 cm Collection numbers: NHMUK 2012.3. 19.3, 2012.3. 19.4. Morphological description. Vegetative stages not found. Spore elongate with rounded ends in valvular view (Figs. 8, 10), flat elongate with attenuated ends in sutural view (Fig. 9). Sutural line straight and fine. Valve shells each with 4���5 surface striations. Sporoplasm located between the polar capsules, binucleate and deeply staining (Fig. 11). Polar capsules long, pyriform, rounded posteriorly and of equal size, at opposite ends of the spore and opening at the tips of the spores. Polar filament thick and with 4���6 coils. Dimensions, based on 25 fixed spores, as ranges with means �� SD in parentheses: spore length 14.4���20.8 (17.0 �� 2.0); spore width 5.4���9.6 (6.7 �� 1.2); spore thickness 3.2���5.2 (4.6 �� 0.6); polar capsule length 3.2 ���4.0 (3.6 �� 0.2); polar capsule width 1.6���3.2 (2.3 �� 0.5); polar capsule length: spore length = 1: 4.5���5.2; spore length: spore width = 1: 2.2���2.7. Molecular result. PCR amplification of myxosporean 18 S rRNA gene from one infected gall bladder from sample (3) above generated a product of approximately 1700 nucleotides in both of the samples processed from the individual fish in question. Forward and reverse sequences were obtained for each purified product. All sequences were identical. Seven polymorphic positions were observed. A final sequence of 1632 nucleotides was submitted to Genebank under Accession No. JX 467674. BLAST analysis revealed that the sequence was novel when compared to data currently deposited in Genbank. Based on the Max scores obtained following BLAST analysis, the sequence showed closest identity to Myxidium coryphaenoideum Noble, 1983 (DQ 377697), Cystodiscus melleni (DQ003031) and Sphaeromyxa zaharoni Diamant et al., 2004 (AY 538662) with 89 %, 87 % and 86 % identity over 100 %, 100 % and 98 % of nucleotide coverage respectively. A sequence of 874 nucleotides remained for phylogenetic analysis following removal of gaps and ambiguous bases. Tetracapsuloides bryosalmonae Canning et al., 1999 was used as the outgroup. MP and ML trees showed similar topologies, with M. baueri grouping with M. coryphaenoideum in both cases, though bootstrap support was relatively poor at 58 and 62 for MP and ML respectively (Fig. 12). Discussion. The genus Myxidium includes more than 230 named species (Eiras et al., 2011), with more than 70 of these reported from marine fishes. The spores found in M. magellanicus most closely resemble those of Myxidium baueri Kovaleva & Gaevskaya 1982, described from two deepwater fishes in the Southwest Atlantic, one of which, Merluccius hubbsi, is closely related to M. magellanicus, both species being merluccids. Kalavati et al. (1995) also reported M. baueri from Merluccius australis (Hutton, 1872) and M. hubbsi caught in the Southwest Atlantic. These authors noted certain differences between their own observations and the original description, most notably the difference between the numbers of spores in the trophozoite and the presence of distinctive knob-like extremities on the spores in their material. We did not find any vegetative stages and we did not observe the knoblike extremities on the spores of our material from M. magellanicus. Despite these differences and because of the otherwise similar morphology, the common locality and the relatedness of the hosts (Table 2), we tentatively identify our material as M. baueri, for which M. magellanicus is a new host record. Unfortunately no sequence data are currently available for M. baueri in the public databases. Of the species of Myxidium for which molecular sequences are available, our material most closely resembles M. coryphaenoidium, collected from Coryphaenoides rupestris Gunnerus, 1765 caught in the North Atlantic (Fiala, 2006), though there is still a sufficient level of nucleotide differences to define it as a new species. Myxidium coryphaenoidium was originally described by Noble (1966) from ��� Coryphaenoides sp.��� caught in the Pacific off Mexico, and has since been reported from 21 species of deepwater marine fish worldwide, most of them belonging to the family Macrouridae, but also including two morid species and one member of the family Synaphobranchidae (Yoshino & Noble, 1973; Yoshino & Moser, 1974; Moser et al., 1976; Kovaleva & Gaevskaya, 1982; Threlfall & Khan, 1990). Such large host and geographical ranges raise the possibility that M. coryphaenoidium may not be a single cosmopolitan species but a complex of sibling species. Moser et al. (1976) described two forms of spore in M. coryphaenoidium, which they called ���long��� and ���stubby���, and the occurrence of which they interpreted as polymorphism within the one species rather than mixed infections of two species. Kovaleva & Gaevskaya (1982) considered their new species M. baueri to be identical to the ���stubby��� form of M. coryphaenoideum. They also described and figured shorter forms of spore in M. baueri, which they described as ���anomalous���. We occasionally observed longer spores similar to those of the ���long��� form of M. coryphaenoideum in our material. This suggests that polymorphism may be a common feature of those species of Myxidium infecting deepwater marine fishes. It is clear that the taxonomy of the species identified as M. coryphaenoidium and M. baueri requires further investigation, using both morphological and molecular methods. Table 2 compares our material from M. magellanicus with the morphological descriptions of both M. baueri and M. coryphaenoidium., Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on pages 547-551, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Family Alatasporidae Shulman, Kovaleva & Dubina, 1979

Family Myxidiidae Th��lohan, 1892, Published as part of Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy & Brickle, Paul, 2013, Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus L��nnberg, 1907 (Teleostei: Merlucciidae), pp. 541-554 in Zootaxa 3647 (4) on page 547, DOI: 10.11646/zootaxa.3647.4.4, http://zenodo.org/record/223001

Hymedesmia (Hymedesmia) barnesi sp. nov. (Figure 10) Type material: Holotype: BELUM Mc 7627. Sample in 95% ethanol, tissue section and spicule preparation on slides; Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. Paratype: BELUM Mc 7677. Sample in 95% ethanol, tissue section and spicule preparation on slides; Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Etymology: Named after Dr David Barnes of British Antarctic Survey, project leader of the ‘Mapping the Benthic Biodiversity of South Georgia’ Darwin Initiative, in recognition of his support of this work. External morphology: In situ appearance: Thinly encrusting (Preserved appearance: Thin white crust. Skeleton: Typical hymedesmoiid skeleton with a dense basal layer of primary and echinating acanthostyles with ascending columns of the ectosomal spicules 3–6 spicules thick. Thick ectosomal layer of chelae (Fig. 10b). Spicules: Measurements from Mc7627. Primary acanthostyles: 272(317)392 by 22(29)39µm at head. Tylote head bearing short rounded spines. Spined up to 1/3 of the shaft (Fig. 10c). Echinating acanthostyles: 102(138)161 by 9.3(17.2)24.5µm at head. Entirely spined with conical pointed spines (Fig. 10d). Ectosomal spicules: styles/tornotes 188(249)276 by 5.1(6.8)9.5µm. Fusiform with the ends variable in form: the majority are styles with one rounded, sometimes slightly tylote, and one pointed end but in some spicules the rounded end is modified into a point (Fig. 10e,f). Chelae: 23.6(27.7)30.7µm (Fig. 10g). Remarks: The majority of Hymedesmia (Hymedesmia) species occurring in the Antarctic have much bigger spicules or they possess sigma microscleres (Table 5). The size range of the spicules in this species is similar to H. gaussiana Hentschel, 1914. Unfortunately the type specimen of this species (ZMH collection) was destroyed in the Second World War and so is not available for examination. However, it differs in having strongyles as ectosomal spicules and entirely spined large acanthostyles., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 17-19, {"references":["Rios, P. (2006) Esponjas del orden Poecilosclerida de las campanas espanolas de bentos Antartico. Unpublished PhD Thesis, University of Santiago de Compostela, Santiago de Compostela, pp. 1 - 527.","Burton, M. (1934) Sponges. Further Zoological Results of the Swedish Antarctic Expedition 1901 - 03. (S Bock, Ed.). P. A. Norstedt & Soner, Stockholm, pp. 58.","Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","Calcinai, B. & Pansini, M. (2000) Four new demosponge species from Terra Nova Bay (Ross Sea, Antarctica). Zoosystema, 22, 369 - 381.","Ridley, S. O. & Dendy, A. (1886) Preliminary Report on the Monaxonida collected by H. M. S. \" Challenger \". Annals and Magazine of Natural History, 18, 325 - 351.","Topsent, E. (1916) Diagnoses d'eponges recueillies dans l'Antarctique par le Pourquoi - Pas? Bulletin du Museum national d'histoire naturelle, 22, 163 - 172."]}

Lissodendoryx (Ectyodoryx) collinsi sp. nov. (Figure 9) Type material: Holotype: BELUM Mc 7676. Sample in 95% ethanol, tissue section and spicule preparation on slides; Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Paratype: BELUM Mc 7681. Sample in 95% ethanol, tissue section and spicule preparation on slides; Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Etymology: Named after Dr Martin Collins, current South Georgia Chief Executive Officer and Director of Fisheries, and member of the Shallow Marine Surveys Group, who generously provided accommodation to researchers pre- and post survey. External morphology: In situ appearance: Massively encrusting peach sponge with irregular lobed surface. Texture of sponge surface smooth (Fig. 9a). Preserved appearance: White sponge, firm but compressible. Smooth ectosomal layer. Skeleton: The choanosomal skeleton is formed of a loose, confused, reticulation of columns of styles echinated by acanthostyles. Ascending columns up to 15 spicules thick joined by bundles of 1–3 spicules. Columns anastomise frequently. The ectosomal skeleton consists of brushes of tornotes (Fig. 9b). Spicules: Measurements from Mc7676. Choanosomal styles: 301(342)396 by 9.1 (12.3) 15.2µm. Smooth styles which come to an abrupt point (Fig. 9c). Echinating acanthostyles: 95(121)178 by 3.8(7.3)13.9µm. Parallel sided then coming to an abrupt point. Entirely spined with small, neat conical spines (Fig. 9d). Ectosomal tornotes: 216(256)295 by 5.4 (7.8)10.9µm (Fig. 9e, f). Microscleres: absent. Remarks: These specimens have been assigned to the genus Lissodendoryx on the basis of the presence of a choanosomal skeleton of styles and acanthostyles and the presence of ectosomal tornotes. The presence of echinating acanthostyles in the skeletal tracts assigns this to the subgenus Lissodendoryx (Ectyodoryx) (van Soest 2002a). All other Antarctic or South Atlantic species have chelae or sigma microscleres (Table 4).

Tedania (Tedaniopsis) wellsae sp. nov. (Figure 16) Type material: Holotype: BELUM Mc 7578. Sample in 95% ethanol, tissue section and spicule preparation on slides. Prion Island Site 1, South Georgia (54°001.590’S, 37°15.178’W); depth 17.6m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. Paratypes: Samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7637. Bird Sound Site 2, South Georgia (54°01.149’S, 38° 01.026’W); depth 18m; collected by C. Goodwin, S. Cartwright and P. Brickle, 22 nd November 2010. BELUM Mc 7642. Bird Sound, Site 4, South Georgia (54°02.095’S, 38° 00.187’W); depth 6m; collected by J. Brown and S. Brown, 22 nd November 2010. Comparative material examined: BMNH 79.12.27.12 Tedania tenuicapitata Ridley, 1881, Holotype. Tissue section and spicule preparation slides. Etymology: Named after Dr Emma Wells, the algal taxonomist on this expedition. External morphology: In situ appearance: Custard yellow thin (Preserved appearance: Cream crust. Firm, not compressible. Surface in brain-like folds (Fig. 16a). Skeleton: The choanosomal skeleton is formed of ascending columns of 4–6 styles irregularly branched and connected with small columns 2–3 styles wide. The onychaetes combined into the choanosomal fibres and brushes but many free onychaetes also present in the choanosomal tissue. The ectosomal skeleton conststs of a palisade of tornotes (Fig. 16b). Spicules: Measurements from Mc7578. Styles: 301(335)382 by 9.6(13.6)17.3µm. Some faintly curved (Fig. 16c). Ectosomal tornotes: 241(309)278 by 6.7(9.4)11.7µm. Anisotornotes with mucronate ends (Fig. 16d). Onychaetes: 63(74)86 and 159(190)206 by 1.4(2.2)2.9µm. The large are pointed at both ends, the small have one pointed and one rounded end (Fig. 16e, f, g). Remarks: This species can be separated from others in the subgenus by the size range of its spicules (Table 8). They are similar in size range to Tedania (Tedaniopsis) tenuicapitata Ridley, 1881 which has similarly sized styles (296–387µm) and tornotes (185–270 µm), but much longer large onychaetes (132–327 and 52–75µm). However, comparison with the type specimen shows that all spicules are much less robust and it does not have the pronounced ascending skeletal columns found in this species. It differs from Tedania (Tedaniopsis) aurantiaca sp. nov. in having an encrusting rather than massive external form and the smaller size of its styles, ectosomal tornotes and onychaetes., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 29-31, {"references":["Koltun, V. M. (1964) Sponges of the Antarctic I. Tetraxonida and Cornacuspongida. In: Pavilovskii, E. P., Andriyashev, A. P. & Ushakov, P. V., (Eds.), Biological reports of the Soviet Antarctic expedition (1955 - 1958). Volume 2. Akademya Nauk SSSR, Moscow, Leningrad, pp. 6 - 133.","Bertolino, M., Schejter, L., Calcinai, B., Cerrano, C. & Bremec, C. (2007) Sponges from a submarine canyon of the Argentine. In: Custodio, M., Lobo - Hajdu, G., Hajdu, E. & Muricy, G. (Eds), Porifera Research: Biodiversity, Innovation and Sustainability. Serie Livros 28. Rio de Janeiro: Universidad Nacional de Rio de Janeiro, pp. 189 - 201.","Dendy, A. 1924. Porifera. Part I. Non - Antarctic sponges. Natural History Report. British Antarctic (Terra Nova) Expedition, 1910 (Zoology) 6: 269 - 392.","van Soest, R. W. M. (2002 b) Family Tedaniidae Ridley and Dendy 1886. 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. 625 - 632.","Burton, M. (1934) Sponges. Further Zoological Results of the Swedish Antarctic Expedition 1901 - 03. (S Bock, Ed.). P. A. Norstedt & Soner, Stockholm, pp. 58.","Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","Ridley, S. O. & Dendy, A. (1886) Preliminary Report on the Monaxonida collected by H. M. S. \" Challenger \". Annals and Magazine of Natural History, 18, 325 - 351.","Burton, M. (1929) British Antarctic (' Terra Nova') Expedition, 1910. Natural History Report. Porifera: Part II: Antarctic Sponges. Zoology, 6, 393 - 458.","Topsent, E. (1916) Diagnoses d'eponges recueillies dans l'Antarctique par le Pourquoi - Pas? Bulletin du Museum national d'histoire naturelle, 22, 163 - 172.","Rios, P. (2006) Esponjas del orden Poecilosclerida de las campanas espanolas de bentos Antartico. Unpublished PhD Thesis, University of Santiago de Compostela, Santiago de Compostela, pp. 1 - 527.","Kirkpatrick, R. (1907) Preliminary Report on the Monaxonellida of the National Antarctic Expedition. Annals and Magazine of Natural History 20, 271 - 291.","Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392."]}

Mycale (Mycale) brownorum sp. nov. (Figure 17) Type material: Holotype: Sample in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7588. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. Paratypes: Samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7586. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7593. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7621. Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. Etymology: Named after Dr Judith Brown, Diving Officer for the expedition and her husband, Steve Brown, member of the expedition dive team. External morphology: In situ appearance: Yellow crust, up to 10mm thick. On the surface of the sponge the ectosomal spicule mesh is visible, giving a honeycomb appearance. Three specimens were growing over algae attached to bedrock. One specimen (Mc7621) is a thicker crust in which the surface has developed into a series of small lumps (Fig. 17a). Preserved appearance: Fragile white specimen which breaks easily into strands formed by the columns of the choanosomal skeleton. The ectosomal layer is more solid and glassily smooth. Skeleton: The choanosomal skeleton is formed of thick (up to 15 spicules wide) ascending columns of mycalostyles which divide towards the ectosome in a dendritic pattern. The ectosome is a tangential confused layer of mycalostyles. Chelae present throughout skeleton, the larger category are in rosettes (Fig. 17b). Spicules: Measurements from Mc7588. Mycalostyles: 448(601)537 by 11.4(15.6)20.3µm base with an oval tylote, other end coming to a rounded point (Fig. 17c). Anisochelae: two categories 30.5(44.4)52.6 and 67.2(81.7)88.0µm (Fig. 17d, e). Remarks: This species can be distinguished from most species the subgenus Mycale (Mycale) by its smaller mycalostyle size or the categories of microscleres, which are present (Table 9). It has a similar size range of mycalostyles to M. doellojuradoi Burton, 1940 but this has three categories of chelae, the smallest being 18µm. Descriptions of other species in the genus were examined in case of taxonomic confusions; these could be distinguished by differences in the size of mycalostyles or the categories of microscleres present.

Clathria (Microciona) matthewsi sp. nov. (Figure 8) Type material: Holotype: BELUM Mc 7606. Sample in 95% ethanol, tissue section and spicule preparation on slides; Rosita Harbour Site 2, South Georgia (54°00.649’S, 37° 25.618’W); depth 11.5m; collected by C. Goodwin, J. Brown, and S. Brown, 20 th November 2010. Paratypes: Samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7625. Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. BELUM Mc 7631 and BELUM Mc 7632. Bird Sound Site 1, South Georgia (54°02.058’S, 38° 00.242’W); depth 18m; collected by C. Goodwin, S. Cartwright and P. Brickle, 22 nd November 2010. BELUM Mc 7667. Husvik, South Georgia (54°10.285’S, 36° 40.412’W); depth 18m; collected by C. Goodwin, D. Poncet and P. Brewin, 26 th November 2010. BELUM Mc 7678. Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Etymology: Named for marine mammal biologist Dr Leonard Harrison Matthews who worked on South Georgia during the Discovery Antarctic Investigations. External morphology: In situ appearance: Lemon to custard yellow thinly encrusting (Preserved appearance: Thin cream crust with a smooth surface. Skeleton: The choanosome is plumose with ascending choanosomal columns of primary acanthostyles very strongly echinated by secondary acanthostyles. Columns placed closely so that the ends of the echinating acanthostyles intermesh. The ectosomal skeleton consists of brushes of a separate category of ectosomal styles (Fig. 8b). Spicules: Measurements from Mc7606. Primary acanthostyles: 244(354)432 by 15.8(18.3)24.5µm. Head not tylote. Spined only basally, to about 1/8 up shaft from the head, with small spines (Fig. 8c). Secondary acanthostyles: 85(108)196 by 9.3(13.8)20.8µm. Entirely spined with large conical spines along their entire length, head not tylote (Fig. 8d). Ectosomal styles: 168(217)254 by 7.6(9.2)11.5µm. Microspined on head (Fig. 8e). Toxa: 59(86)121µm with spined ends (Fig. 8f). Chelae: 9(10)11µm very abundant (Fig. 8g). Remarks: We have assigned these specimens to the subgenus Clathria (Microciona) on the basis of their encrusting growth form and plumose skeletal architecture (Hooper 2002). There are four species of Clathria (Microciona) which have been recorded from the Antarctic and South Atlantic: C.antarctica (Topsent, 1917), C. basispinosa (Burton, 1934), C. tuberculata (Burton, 1934), and C. sigmoidea (Cuartas, 1992). However, none of these possess chelae and therefore can be readily distinguished from our specimens., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 15-16, {"references":["Topsent, E. (1917) Spongiaires. In: Joubin, L. (Ed.), Deuxieme Expedition Antarctique Francaise (1908 - 1910) Commandee par le Dr. Jean Charcot. Sciences Physiques: Documents Scientifiques. Masson & Cie, Paris, pp. 1 - 88.","Burton, M. (1934) Sponges. Further Zoological Results of the Swedish Antarctic Expedition 1901 - 03. (S Bock, Ed.). P. A. Norstedt & Soner, Stockholm, pp. 58.","Cuartas, E. I. (1992) Poriferos de la provincia biogeografica Argentina. III. Poecilosclerida (Demospongiae), del litoral marplatense. Physis (Buenos Aires) (A), 47, 73 - 88."]}

Myxilla (Ectyomyxilla) kerguelensis (Hentschel, 1914) (Figure 14) Synonymy: Ectyomyxilla kerguelensis Hentschel, 1914. Myxilla kerguelensis (Hentschel, 1914). Crellomyxilla intermedia Dendy, 1924. Not Myxilla tornotata Brøndsted, 1924. Material: Samples in 95% ethanol, tissue section and spicule preparation on slides; BELUM Mc 7589, BELUM Mc 7591, BELUM Mc 7595 and BELUM Mc 7596. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7684. Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Comparative material examined: ZMH S2325 Ectomyxilla kerguelensis, S2319 Acanthoxa werthii External morphology: In situ appearance: Thickly encrusting orange sponge, some specimens very thick and mounded. Scattered large oscules up to 0.5cm in diameter. Surface with a honeycomb appearance due to patches of ostia (Fig. 14a). Preserved appearance: Very firm, pale yellow, crust with some inclusions of grey sediment grains. Spaces visible in choanosome. Ectosome is glassily smooth but is not easily detachable. Skeleton: Choanosome: Dense reticulation of bundles of 2–3 acanthostyles, predominantly large acanthostyles but some small acanthostyles are scattered through the bundles. Sigmas and chelae scattered abundantly through the tissue. Ectosome: Palisade of tornotes covered with tangential crust of the small acanthostyles (Fig. 14b). Spicules: Measurements from Mc7589. Choanosomal acanthostyles: 203(226)245 by 15(19)25µm. Parallel sided acanthostyles with large, conical spines. End terminates in an abrupt, unspined point (Fig. 14c). Ectosomal acanthostyles: 74(91)109 by 7(10)13µm. Similar in form to the choanosomal acanthostyles but with much larger spines relative to the shaft (Fig. 14d). Ectosomal tornotes: 174(190)210 by 7(9)12µm. Fusiform tornotes with mucronate points (Fig. 14e). Chelae: 18(20)23µm (Fig. 14f). Sigmas: 18(24)29µm (Fig. 14g). Remarks: The specimen appears to be a good match for the type description. There is a slight difference in that the type specimens are reported as massive and ‘mostly in rounded form’ with the largest piece 11.5cm wide and 7cm high, rather than thickly encrusting. However, it seems that specimens might be quite variable in form, as samples collected in this survey range from thickly encrusting to massive mounds (Mc7596 and Mc7684 were approximately 10cm high). Hentschel (1914) reports that some of his specimens were found on whale bones which may indicate that they were actually very thickly encrusting. In terms of speculation, the chelae (12.5–19µm), small acanthostyles (56–75µm) and sigmas (17–22µm) reported from the type are slightly smaller than those of our specimens. However, Boury-Esnault and Van Beveren (1982) described a larger size range in their specimens, similar to that found in ours. It was not possible to examine the type as the type specimen jar was found to contain a specimen of Acanthoxa werthii Hetschel, 1914 (now reassigned to Spanioplon werthi (Hentschel, 1914). The species Crellomyxilla intermedia Dendy, 1924, described from New Zealand (Dendy 1924), is currently regarded as a synonym. This has similar sizes spicules and ectosomal acanthostyles to the type specimen of M. kerguelensis. Dendy describes two categories of chelae, but Burton (1929) on re-examination of the type found only one category. Burton (1929, 1934) also considered Myxilla tornotata Brøndstedt, 1924 a synonym, despite noting it differed in having microspined ends to the tornotes. However, the two species can be separated by the size of the sigmas and the size and form of the ectosomal spicules (Lévi 1956; Boury-Esnault and Van Beveren 1982) and have been recorded sympatrically from the Kerguelen Islands (Boury-Esnault and Van Beveren 1982). Distribution: Other than the type locality, Kerguelen (Lévi 1956; Boury-Esnault & Van Beveren 1982), this species has been recorded from McMurdo sound (Burton, 1929), although these specimens differ from the type description in that they are flabellate specimens with chelae up to 33µm, and New Zealand (as Crellomyxilla intermedia) (Dendy 1924). There are no previous records from South Georgia., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 26-27, {"references":["Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","Dendy, A. 1924. Porifera. Part I. Non - Antarctic sponges. Natural History Report. British Antarctic (Terra Nova) Expedition, 1910 (Zoology) 6: 269 - 392.","Burton, M. (1929) British Antarctic (' Terra Nova') Expedition, 1910. Natural History Report. Porifera: Part II: Antarctic Sponges. Zoology, 6, 393 - 458.","Burton, M. (1934) Sponges. Further Zoological Results of the Swedish Antarctic Expedition 1901 - 03. (S Bock, Ed.). P. A. Norstedt & Soner, Stockholm, pp. 58.","Levi, C. (1956) Eponges littorales des Iles Kerguelen recoltees par M. Angot. Memoires de l'Institut Scientifique de Madagascar (A), 10, 25 - 34."]}

Mycale (Mycale) cartwrighti sp. nov. (Figure 18) Type material: Holotype: Sample in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7590. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. Paratypes: Sample in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7638. Bird Sound Site 2, South Georgia (54°01.149’S, 38° 01.026’W); depth 18m; collected by C. Goodwin, S. Cartwright and P. Brickle, 22 nd November 2010. Etymology: Named after Steve Cartwright, member of the expedition dive team. External morphology: In situ appearance: Thickly encrusting lemon yellow sponge (up to 10mm thick) in large patches (> 15cm) with sparsely scattered large oscules. Sub-ectosomal spaces clearly visible through ectosome giving the sponge surface a slightly lumpy appearance (Fig. 18a). Preserved appearance: Very pale yellow crust. Choanosome firm but compressible, ectosome visible as a separate, harder layer. Skeleton: The choanosomal skeleton is formed of thick (up to 20 spicules wide) ascending columns of mycalostyles which anastomise and coalesce. The ascending columns are joined by shorter, thinner columns (1–3 spicules in width). The ectosomal skeleton is formed by the choanosomal columns fanning out at the surface to form a confused tangential ectosomal layer of mycalostyles. Chelae and trichodragmata present throughout skeleton, chelae do not form rosettes (Fig. 18b). Spicules: Measurements from Mc7590. Styles: 349 (407)451 by 9.3 (12.6)16.9µ m. Mycalostyles with a neat rounded head and abruptly taped tip (Fig. 18c). Chelae: 23(35)47µm—rarely 70µ m in Mc7638. Typical mycalid anisochelae with the lower front alae coming to a blunt point (Fig. 18d). Microxea: 12 (23)40µm, very small microxea Remarks: This species can be distinguished from the majority of other Mycale (Mycale) species by the possession of small microxea/trichodragmata which is unusual in the genus Mycale (Table 9). Of those species which do possess these Mycale (Mycale) macrochela Burton, 1932 has sigmas and smaller chelae (7–35µm), Mycale (Aegogropila) meridionalis Lévi, 1963 has trichodragmata 16–17µm but also has sigmas and smaller mycalostyles, Mycale (Aegogropila) nodulosa Goodwin et al., 2011a has similar sized spicules but possesses sigmas, a second minute (5µm) category of microxea and has a clearly reticulate ectosomal skeleton which assigns it to a different subgenus, and Mycale (Carmia) diminuta Sará, 1978 has similarly sized mycalostyles but larger trichodragmata (45–60µm). Mycale (Mycale) brownorum sp. nov. can be distinguished as it also lacks microxea microscleres., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 32-33, {"references":["Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392.","Levi, C. (1963) Spongiaires d'Afrique du Sud. (1) Poecilosclerides. Transactions of the Royal Society of South Africa, 37, 1 - 72.","Sara, M. (1978) Demospongie di acque superficiali della Terra del Fuoco (Spedizioni AMF Mares - GRSTS e SAI). Bollettino dei Musei e degli Istituti Biologici della Universita di Genova, 46, 7 - 117."]}

Clathria (Clathria) stromnessa sp. nov. (Figure 6) Type material: Holotype: BELUM Mc 7690. Sample in 95% ethanol, tissue section and spicule preparation on slides; Green Island, Stromness, Site 2, South Georgia (54°09.381’S, 36° 39.852’W); depth 17.4m; collected by C. Goodwin, J. Brown, and S. Brown, 28 th November 2010. Paratype: BELUM Mc 7674. Sample in 95% ethanol, tissue section and spicule preparation on slides, Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. Etymology: Named after the type locality, Stromness, South Georgia External morphology: In situ appearance: Lobed massive (type specimen maximum diameter 20cm) rust orange sponge with large oscules on top of lobes (Fig. 6a). Preserved appearance: Grey, firm, with a slightly hispid surface. Skeleton: The choanosomal skeleton is an irregular reticulation of bundles of 4–6 smooth styles (Fig. 6b), there is no differentiation between axial and extra-axial regions. The ectosome is formed of brushes of styles. Toxa and chelae microscleres are present throughout the tissue. Spicules: Measurements from Mc7690. Choanosomal styles: 424(495)563 by 17(26)31µm. Fat smooth styles, the majority are gently curved (Fig. 6c). Ectosomal styles: 232(292)414 by 3(6)7µm. Thin styles, the heads are microspined, bearing several short blunt spines (Fig. 6d, e). Chelae: 10(12)14µm. Typical clathriid palmate isochelae (Fig. 6f). Toxa: 45(150)477µm, thin smooth toxas, very wide ranging in size. Some of the smallest have a pronounced central flexion and upturned points (Fig. 6g). Remarks: We have assigned this species to Clathria (Clathria) rather than one of the other seven sub-genera on the basis of the lack of differentiation between the axial and extra-axial regions of the choanosome and the presence of a reticulate skeleton and only a single category of auxillary style (Hooper 2002). This species is unusual in not possessing any echinating acanthostyles, however these can be secondarily lost in this subgenus (Hooper 2002). The subgenus Clathria (Isociella) lacks echinating acanthostyles but has a regular renieroid reticulate skeleton with plumose multispicular tracts connected by paucispicular ones (Hooper 2002), whereas the skeleton of our species is irregularly reticulate. From the South Atlantic and Antarctic C. papillosa Thiele, 1905 and C. paucispicula (Burton, 1932) are the only species in Clathria (Clathria), which do not possess any echinating spicules. However, the former has strongylote ectosomal spicules and bipocoelles and should probably be reassigned to Iophon, and the latter lacks any microscleres. The large size of the toxa is also unusual; C. toxipraedita Topsent, 1913 (type locality Burdwood Bank to the south of Falkland Islands and north west of South Georgia) has toxa up to 1750µm but possesses echinating acanthostyles., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on page 12, {"references":["Thiele, J. (1905) Die Kiesel - und Hornschwamme der Sammlung Plate. Zoologische Jahrbucher, Supplement, 407 - 496.","Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392.","Topsent, E. (1913) Spongiaires de l'Expedition Antarctique Nationale Ecossaise. Transactions of the Royal Society of Edinburgh, 49, 579 - 643."]}

Polymastia invaginata Kirkpatrick, 1907 (Figure 3) Synonymy: Polymastia invaginata Kirkpatrick, 1907. Polymastia invaginata var. gaussi Hentschel, 1914 is regarded as a synonym by Plotkin and Janussen (2008). Material: All samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7610. Rosita Harbour Site 2, South Georgia (54°00.649’S, 37° 25.618’W); depth 11.5m; collected by C. Goodwin, J. Brown, and S. Brown, 20 th November 2010. BELUM Mc 7618 and BELUM Mc 7620. Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. BELUM Mc 7648. Jagged Point, Possession Bay, South Georgia (54°04.514’S, 37° 07.188’W); depth 10.4m; collected by C. Goodwin, D. Poncet and P. Brewin, 23 rd November 2010. BELUM Mc 7660. Husvik, South Georgia (54°10.285’S, 36° 40.412’W); depth 18m; collected by C. Goodwin, D. Poncet and P. Brewin, 26 th November 2010. Comparative material examined: BMNH 03.2.5.78 Polymastia invaginata. Discovery Antarctic Expedition McMurdo Bay ~20fms. W.Q.28.2.02. Specimen in alcohol. This specimen is the figured half specimen in PL XIV Fig. 5 of the description. External morphology: In situ appearance: Brown hispid mound attached to bedrock bearing single, lemon yellow, smooth surfaced, large papillae with large terminal oscule. Individuals up to 20cm in height (Fig. 3a). Often occur in clusters of several individuals. Preserved appearance: Transverse slice of basal mound of specimen. Tissue very tough. Choanosome grey, cortical layer white and 2–3mm thick. A fringe of dark grey hairs ~ 5mm long present on the surface. Skeleton: Radiate skeleton of bundles of large styles, these penetrate the ectosome and form the thick surface pile (Fig. 3b). Stellate groups of small tylostyles are present between the fibres (Fig. 3c). The ectosome is formed of a dense tylostyles, positioned vertically with their points towards the surface. This layer forms a fibrous cortex to the sponge, easily visible on slides, around 0.5–1cm thick. Spicules (Fig. 3d): Styles: 1825 (2462) 3076 by 22.1 (26.9) 42.1µm—although many broken and difficult to measure so longer spicules may be present. Tylostyles: 119 (307) 625 by 6.3 (10.1) 13.6µm. Fusiform tylostyles with a neat swelling at their head. Some forming stellate clusters between the fibres but these do not seem to represent a seperate size category. Remarks: Our specimens are a good match with the type description and specimens and correspond to the external form and spiculation of other specimens assigned to this species (Brueggeman, 1998; Hentschel 1914; Koltun 1964; Plotkin and Janussen, 2008). However, like previous authors, we did not record the sceptre-like spicules noted by Plotkin and Janussen (2008) in the cortical palisade and our styles are of a larger size than those noted by Boury-Esnault and van Beveren (1982). Polymastia invaginata can be distinguished from other Antarctic and Southern Atlantic species of Polymastia by its single inhalant papillae, densely hispid surface and single spicule layer in the cortex (Plotkin and Janussen, 2008). Kirkpatrick (1907) noted that the papillae in all of his specimens was ‘invaginated’, flush with the surface of the basal mound, this is presumably the origination of the species name. He was studying preserved material and this may have been an artifact of preservation, in all our living specimens the papillae stood proud. Distribution: Originally recorded from Winter Quarters (18–55m depth) and from off Mount Erebus (914m depth). Widespread in the Antarctic and sub-Antarctic: records from Kerguelen and Heard Islands (Boury-Esnault and Van Beveren 1982) McMurdo Sound (Burton 1929), South Georgia, South Orkneys and South Shetlands (Burton 1932) in depths of 18–1080m. Polymastia invaginata var. gaussi Hentschel, 1914 was regarded as a synonym by Burton (1932) but this is a much smaller sponge (maximum 8mm high) and has smaller spicules (styles up to 1792µm, tylostyles 120–600µm) so is a distinct species., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 6-7, {"references":["Kirkpatrick, R. (1907) Preliminary Report on the Monaxonellida of the National Antarctic Expedition. Annals and Magazine of Natural History 20, 271 - 291.","Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","Plotkin, A. S. & Janussen, D. (2008) Polymastiidae and Suberitidae (Porifera: Demospongiae: Hadromerida) of the deep Weddell Sea, Antarctic. Zootaxa, 1866, 95 - 135.","Brueggeman, P. (1998) Porifera - Demospongiae: demosponges. Underwater Field Guide to Ross Island & McMurdo Sound, Antarctica. National Science Foundation.","Koltun, V. M. (1964) Sponges of the Antarctic I. Tetraxonida and Cornacuspongida. In: Pavilovskii, E. P., Andriyashev, A. P. & Ushakov, P. V., (Eds.), Biological reports of the Soviet Antarctic expedition (1955 - 1958). Volume 2. Akademya Nauk SSSR, Moscow, Leningrad, pp. 6 - 133.","Burton, M. (1929) British Antarctic (' Terra Nova') Expedition, 1910. Natural History Report. Porifera: Part II: Antarctic Sponges. Zoology, 6, 393 - 458.","Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392."]}

Haliclona (Soestella) crowtheri sp. nov. (Figure 19) Type material: Holotype: Sample in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7579. Prion Island Site 1, South Georgia (54°001.590’S, 37°15.178’W); depth 17.6m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. Paratypes: Sample in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7587 and BELUM Mc 7594. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7612. Rosita Harbour Site 2, South Georgia (54°00.649’S, 37° 25.618’W); depth 11.5m; collected by C. Goodwin, J. Brown, and S. Brown, 20 th November 2010. BELUM Mc 7623. Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. BELUM Mc 7633. Bird Sound Site 1, South Georgia (54°02.058’S, 38° 00.242’W); depth 18m; collected by C. Goodwin, S. Cartwright and P. Brickle, 22 nd November 2010. Etymology: Named for the former Head of the Department of Natural Sciences National Museums Northern Ireland, Dr Peter Crowther, who retired this year after many years of service to the museum; in recognition of his support of this work when in post. External morphology: In situ appearance: Thickly encrusting white sponge (up to 15mm thick) forming large patches (up to 20cm in diameter) on bedrock. Smooth surface bearing numerous large oscules (up to 1cm in diameter) (Fig. 19a). Preserved appearance: Thick crust with very hard texture. Choanosome brick red in Mc7612 but white in some specimens and patchily red in others. The ectosome is a white, easily detachable, smooth layer. Skeleton: Confused choanosomal skeleton with primary columns of 4–7 spicules joined by unispicular secondary lines. Some rounded meshes present (Fig. 19b). Detachable white ectosome formed of hexagonal meshes of oxea, each side composed of single bundle of oxea 2–3 spicules thick (Fig. 19c). Spicules: Measurements from Mc7579. Oxea: 266(299)321 by 11(17)20µm—occasional very thin (Remarks: This species is assigned to Haliclona as it is a Chalindae with unispicular secondary lines (de Weerdt 2002). The presence of a specialised ectosomal skeleton with rounded meshes assigns it to the subgenus Haliclona (Soestella) (de Weerdt 2002). Two other species of Haliclona (Soestella) have been recorded from this region: H. auletta (Thiele, 1905) from Calbuco, Chile and H. chilensis (Thiele, 1905). However, these both possess much smaller oxeas (150 and 130–200µm respectively). The taxonomy of this genus is still confused so species from other sub-genera were considered. Although descriptions of other species from different genera in the family occurring in the region have been examined none can be found in which such a distinctive ectosomal skeleton is described., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 33-36, {"references":["Levi, C. (1963) Spongiaires d'Afrique du Sud. (1) Poecilosclerides. Transactions of the Royal Society of South Africa, 37, 1 - 72.","Burton, M. (1940) Las Esponjas marinas del Museo Argentino de Ciencias Naturales. (Parte 1). Anales del Museo argentino de ciencias naturales \" Bernardino Rivadavia \", 40, 95 - 121.","Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392.","Willenz, P., Hajdu, E., Desqueyroux - Faundez, R., Lobo - Hajdu, G. & de Souza Carvalho, M. (2009) Class Demospongiae. In: Haussermann, V. & Forsterra, G., (Eds.) Marine Benthic Fauna of Chilean Patagonia. Nature in Focus, Puerto Montt, pp. 94 - 170.","Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","Thiele, J. (1905) Die Kiesel - und Hornschwamme der Sammlung Plate. Zoologische Jahrbucher, Supplement, 407 - 496."]}

Myxilla (Burtonanchora) ponceti sp. nov. (Figure 13) Type material: Holotype: BELUM Mc 7572. Sample in 95% ethanol, tissue section and spicule preparation on slides; Floating Dock, Grytviken, South Georgia (54º 16.937´S, 36º 30.475´W); depth range: 0–6.6m; collected by C. Goodwin and S. Cartwright, 18 th November 2010. Paratypes: Samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7573. Floating Dock, Grytviken, South Georgia (54º 16.937´S, 36º 30.475´W); depth range: 0–6.6m; collected by C. Goodwin and S. Cartwright, 18 th November 2010. BELUM Mc 7600. Prion Island Site 2, South Georgia (54°001.862’S, 37° 15.032’W); depth 18m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7655. Prince Olav Harbour, South Georgia (54°03.579’S, 37° 08.203’W); depth 18m; collected by C. Goodwin, D. Poncet, and S. Brown, 23 rd November 2010. Etymology: Named for Dion Poncet, member of the expedition dive team, who alerted us to this ‘probable sponge’ under Grytviken jetty, and whose knowledge of South Georgia was invaluable to this survey. External morphology: In situ appearance: Specimens are large (10–20cm maximum dimension), smooth surfaced, lobed sponges, carrot orange in colour, bearing large terminal oscules on the ends of the lobes. The two largest specimens are fan shaped and bear the oscules in a line along the edge of the fan. They are attached by the base of the specimen to the substrate; the attachment point is quite broad in some specimens but in the fan shaped specimens is smaller and might be considered a stalk (Fig. 13a). Preserved appearance: Tissue firm but compressible. Ectosome is a dark brown, choanosome a paler brown. Skeleton: Choanosomal skeleton consists of ascending fibres of 4–10 styles joined by smaller, shorter, fibres of 1–3 styles. There is an ectosomal palisade of tylotes. Chelae scattered throughout skeleton and not forming rosettes (Fig. 13b). Spicules: Measurements from Mc7572. Choanosomal subtylostyles: 281(325)369 by 12.2(14.3)18.7µm. Smooth styles, often slightly curved. Some have a faint swelling at the head (Fig. 13c). Ectosomal tylotes: 197(276)324 by 6.9(10.3)14.5µm. Anisotylotes in which one or both ends are swollen. In some one end has a mucronate point (Fig. 13d). Chelae: In two categories 31(40)49 and 57(66)72µm (Fig. 13e). Remarks: Myxilla (Burtonanchora) is defined by possession of smooth rather than spined choanosomal styles. There are several species of Antarctic Myxilla (Burtonanchora) species (Table 7). However, all of these can be distinguished by having much larger choanosomal styles or the presence of sigmata. Myxilla (Burtonanchora) pistillaris Topsent, 1916 is most similar but has larger styles (480–500µm) and possesses raphides which are not present in this species., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 24-26, {"references":["Rios, P. & Cristobo, F. J. (2007) Sponges of genus Myxilla Schmidt, 1862, collected in Antarctic waters by Spanish Antarctic expeditions. In: Custodio, M., Lobo - Hajdu, G., Hajdu, E. & Muricy, G. (Eds), Porifera Research: Biodiversity, Innovation and Sustainability. Serie Livros 28. Rio de Janeiro: Universidad Nacional de Rio de Janeiro, pp. 525 - 546.","Topsent, E. (1916) Diagnoses d'eponges recueillies dans l'Antarctique par le Pourquoi - Pas? Bulletin du Museum national d'histoire naturelle, 22, 163 - 172.","Ridley, S. O. & Dendy, A. (1886) Preliminary Report on the Monaxonida collected by H. M. S. \" Challenger \". Annals and Magazine of Natural History, 18, 325 - 351."]}

Phorbas glaberrimus (Topsent, 1917) (Figure 12) Synonymy: Clathrissa glaberrima Topsent, 1917. Anchinoe glaberrima (Topsent, 1917). Material: Samples in 95% ethanol, tissue section and spicule preparation on slides; BELUM Mc 7651. Jagged Point, Possession Bay, South Georgia (54°04.514’S, 37° 07.188’W); depth 10.4m; collected by C. Goodwin, D. Poncet and P. Brewin, 23 rd November 2010. BELUM Mc 7671. Husvik, South Georgia (54°10.285’S, 36° 40.412’W); depth 18m; collected by C. Goodwin, D. Poncet and P. Brewin, 26 th November 2010. BELUM Mc 7675. Green Island, Stromness, Site 1, South Georgia (54°09.448’S, 36° 39.752’W); depth 17.4m; collected by C. Goodwin, P. Brickle and S. Cartwright, 27 th November 2010. BELUM Mc 7687 and BELUM Mc 7689. Green Island, Stromness, Site 2, South Georgia (54°09.381’S, 36° 39.852’W); depth 17.4m; collected by C. Goodwin, J. Brown, and S. Brown, 28 th November 2010. Comparative material examined: NMHN DT734 Phorbas glaberrimus (Topsent, 1916) Holotype. From near Terre Alexandre. Spicule preparation and tissue section on slides. External morphology: In situ appearance: Massively thickly encrusting sponge with the surface rising up into one or more mounds. Specimens very variable in size but can be quite large with diameters of greater than 30cm. Sponge encrusting on bedrock and are frequently overgrown by ascidians and bryozoans. Surface covered with large pore sieves (up to 1.5cm in diameter) and large, irregularly scattered, oscules, which can reach 2cm in diameter, are present. Sponge white to peach in colour but with a patchily dark chocolate brown surface layer present in some specimens, possibly formed of algae. The pore sieves often have a slightly lighter rim than the main sponge body. (Fig. 12a). Preserved appearance: Firm but compressible grey sponge with pore sieves clearly visible on the tough ectosomal layer. Skeleton: Choanosomal skeleton composed of a basal layer of acanthostyles from which thick ascending columns of up to 20 oxea ascend. Acanthostyles echinate the lower parts of these columns. In the ectosome the ends of the columns of oxea fan out to form a continuous surface layer. Chelae are very abundant and scattered throughout tissue (Fig. 12b). Spicules: Measurements from Mc7689. Acanthostyles: 216(264)313 by 9.5(14.1)19.7µm. Parallel sided with an abrupt point. Head not tylote. Strongly spined along whole length (Fig. 12c). Ectosomal oxea: 365(428)476 by 9.9(13.0)17.9µm. Fusiform with abrupt points (Fig. 12d). Chelae: 18.9(22.4)25.4µm (Fig. 12e). Remarks: Phorbas glaberrimus (Topsent, 1917) was originally described from 297m in the Antarctic and appears to be a good match for these specimens, although Topsent reports slightly larger oxea (530–600 by 20–22µm). Rios (2006) reports smaller oxea of 420–590µm, more similar to the range in our specimens, and Koltun (1964) also gives a wider range (382–600µm). Rios (2006) reports two categories of acanthostyles (200–260 and 300–350µm), the acanthostyles of Mc7689 could be sub-divided into two length categories 216–233µm and 271–313µm but this division is not apparent in the other specimens and not obvious when viewed down the microscope. Distribution: This species is widely distributed in the Antarctic: Alexander I land (Topsent 1917), Wilheim II coast, Banzare coast, Wilkes Land, Victoria Land, Princess Astrid Coast (Koltun 1964), MacRobertson Coast (Koltun 1976), Weddell Sea (Barthel et al. 1990; Gutt and Koltun 1995), Ross Sea (Pansini et al. 1994), South Trinidad Island and Bransfield Strait (Rios 2006) from depths 90–1370m. Burton (1929) also erroneously reported it as a synonym of Pyloderma latrunculioides (Ridley and Dendy, 1886) from the Antarctic but it is not clear which of his specimens are this species. There are no previous published records from South Georgia., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 23-24, {"references":["Topsent, E. (1917) Spongiaires. In: Joubin, L. (Ed.), Deuxieme Expedition Antarctique Francaise (1908 - 1910) Commandee par le Dr. Jean Charcot. Sciences Physiques: Documents Scientifiques. Masson & Cie, Paris, pp. 1 - 88.","Topsent, E. (1916) Diagnoses d'eponges recueillies dans l'Antarctique par le Pourquoi - Pas? Bulletin du Museum national d'histoire naturelle, 22, 163 - 172.","Rios, P. (2006) Esponjas del orden Poecilosclerida de las campanas espanolas de bentos Antartico. Unpublished PhD Thesis, University of Santiago de Compostela, Santiago de Compostela, pp. 1 - 527.","Koltun, V. M. (1964) Sponges of the Antarctic I. Tetraxonida and Cornacuspongida. In: Pavilovskii, E. P., Andriyashev, A. P. & Ushakov, P. V., (Eds.), Biological reports of the Soviet Antarctic expedition (1955 - 1958). Volume 2. Akademya Nauk SSSR, Moscow, Leningrad, pp. 6 - 133.","Koltun, V. M. (1976) Porifera - Part 1: Antarctic Sponges. Report B. A. N. Z. Antarctic Research Expedition 1929 - 1931 (B, Zoology and Botany), 5, 163 - 198.","Barthel, D., Tendal, O. S. & Panzer, K. (1990) Ecology and taxonomy of sponges in the eastern Weddell Sea shelf and slope communities. Reports on polar research, 68, 120 - 131.","Gutt, J. & Koltun, V. M. (1995) Sponges of the Lazarev and Weddell Sea, Antarctica: explanations for their patchy occurrence. Antarctic Science, 7, 227 - 234.","Pansini, M., Calcinai, B., Cattaneo-Vietti, R. & Sara, M. (1994) Demosponges from Terra Nova Bay (Ross Sea, Antarctica): 1987 / 1988 and 1989 / 90. Programma Nazionale di Ricerche in Antartide, Expeditions. National Scientific Commission for Antarctica Oceanographic Campaign 1987 / 88 and 1989 / 90, 3, 67 - 100.","Burton, M. (1929) British Antarctic (' Terra Nova') Expedition, 1910. Natural History Report. Porifera: Part II: Antarctic Sponges. Zoology, 6, 393 - 458.","Ridley, S. O. & Dendy, A. (1886) Preliminary Report on the Monaxonida collected by H. M. S. \" Challenger \". Annals and Magazine of Natural History, 18, 325 - 351."]}

Tedania (Tedaniopsis) aurantiaca sp. nov. (Figure 15) Type material: Holotype: BELUM Mc 7661. Husvik, South Georgia (54°10.285’S, 36° 40.412’W); depth 18m; collected by C. Goodwin, D. Poncet and P. Brewin, 26 th November 2010. Paratypes: Samples in 95% ethanol, tissue section and spicule preparation on slides. BELUM Mc 7583. Prion Island Site 1, South Georgia (54°001.590’S, 37°15.178’W); depth 17.6m; collected by C. Goodwin, D. Poncet, and P. Brewin, 19 th November 2010. BELUM Mc 7624. Right Whale Bay, South Georgia (54°00.173’S, 37° 40.856’W); depth 18m; collected by C. Goodwin, J. Brown and S. Brown, 21 st November 2010. BELUM Mc 7652. Jagged Point, Possession Bay, South Georgia (54°04.514’S, 37° 07.188’W); depth 10.4m; collected by C. Goodwin, D.Poncet and P. Brewin, 23 rd November 2010. BELUM Mc 7659. Husvik, South Georgia (54°10.285’S, 36° 40.412’W); depth 18m; collected by C. Goodwin, D. Poncet and P. Brewin, 26 th November 2010. BELUM Mc 7670. Husvik, South Georgia (54°10.150’S, 36° 39.322’W); depth 18m; collected by D. Poncet, P. Brewin, C. Goodwin 26 th November 2010. Comparative material examined: MNHN DT1667 Tedania (Tedaniopsis) charcoti Topsent, 1908 Holotype. Ile Booth-Wandel ‘ Le Francais’. Microscope preparations. ZMB S2315 Tedania vanhöffeni var. gracilis Hentschel, 1914 (Holotype T. gracilis (Hetschel, 1914)), specimen and spicule preparation and tissue section prepared from specimen. BMNH 79.12.27.12 Tedania tenuicapitata Ridley, 1881, Holotype. Tissue section and spicule preparation slides. Etymology: From the Latin aurantiacus, meaning orange-coloured. External morphology: In situ appearance: Mustard yellow to cadmium orange lobed crust with large oscules. In thicker, larger, specimens the oscules may be raised on mammiform processes (Fig. 15a). Preserved appearance: Cream sponge composed of rounded lobes, firm, not compressible. Ectosome very smooth. Skeleton: The choanosomal skeleton is a loose reticulation of bundles of 2–3 styles. The ectosomal skeleton consists of bundles of tornotes, which fan out to form a tangential surface layer. Onychaetes are scattered abundantly throughout the choanosome and ectosome (Fig. 15b). Spicules: Measurements from Mc7661. Styles: 356(407)447 by 17(21)25µm. Often slightly curved, parallel sided then tapering at the end to a sharp point (Fig. 15c). Ectosomal tornotes: 335(359)403 by 10(13)16µm. Fusiform anisotornotes (Fig. 15d). Onychaetes: two categories 74(90)125 and 235(261)283 by 1.7(2.1)2.8µm a few intermediates (172,206µm). The largest are pointed at both ends, the smaller with one pointed and one rounded end (Figs 15 e, f, g). Remarks: Desqueyroux-Faúndez and van Soest (1996) reassessed the genera Tedania and Trachytedania, and reclassified the species present into three sub-genera: Tedania (Tedaniopsis) with long styles 300–700 µm, Tedania (Tedania) with short styles 150–300 µm and mucronate tornotes, and Tedania (Trachytedania) with smooth or spined short styles 150–300 µm and oxeote or mucronate tornotes (see also van Soest 2002b). The redescription of Tedania (Trachytedania) was based on the fact that no basal acanthostyles, the characterizing feature of the genus, could be found in the type species. Trachytedania has since been re-established as a valid genus by Cristobo and Urgorri (2001) who re-examined the type and located basal acanthostyles. This species confirms to the current definition of Tedania (Tedaniopsis) as it possesses styles longer than 350µm (Desqueyroux-Faúndez and van Soest 1996). There are many Southern Ocean species within this subgenus but the majority have spicules that are much larger (Table 8). Three species have spicules of a similar range: T. gracilis (Hentschel, 1914) is the most similar in spicule size but has larger styles and shorter tornotes (Table 8). Comparison with the specimen designated as the holotype showed this has much thinner, longer styles (552(591)653 by 12(15)16µm) and thinner ectosomal spicules (397(437)490 by 6(9)13µm), however, this specimen may not be the one described in type description as it appears to differ in spiculation and is a thinly encrusting species on a bryozoan rather a sea urchin. T. charcoti Topsent, 1908 has much thinner styles (420–450 by 13µm in description, 401(434)466 by 9(13)16 µm from our measurements of type) and smaller tornotes (305–340 by10µm from description, 280–350 by 8(10)14µm from our measurements). Tedania tenuicapitata Ridley, 1881 has much smaller styles (296–387µm), tornotes (185–270µm), and onychaetes (132–327, 52–75µm) (Table 8)., Published as part of Goodwin, Claire & Brickle, Paul, 2012, Sponge biodiversity of South Georgia island with descriptions of fifteen new species, pp. 1-48 in Zootaxa 3542 on pages 28-29, {"references":["Hentschel, E. (1914) Monaxone Kieselschwamme und Hornschwamme der Deutschen Sudpolar - Expedition 1901 - 1903. Deutsche Sudpolar - Expedition, 1901 - 03, 15, 35 - 141.","van Soest, R. W. M. (2002 b) Family Tedaniidae Ridley and Dendy 1886. 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. 625 - 632.","Cristobo, F. J. & Urgorri, V. (2001) Revision of the genus Trachytedania (Porifera: Poecilosclerida) with a description of Trachytedania ferrolensis sp. nov. from the north - east Atlantic. Journal of the Marine Biological Association of the United Kingdom, 81, 569 - 579."]}