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Type: Article
Published: 2016-12-21
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Integrative taxonomic re-description of Halisarca magellanica and description of a new species of Halisarca (Porifera, Demospongiae) from Chilean Patagonia 

Royal Belgian Institute of Natural Sciences, Taxonomy and Phylogeny, Rue Vautier 29, B–1000, Bruxelles, Belgium Université Libre de Bruxelles, Laboratoire de Biologie Marine, Avenue F.D. Roosevelt 50, B–1050, Bruxelles, Belgium
Aix Marseille University, Avignon University, CNRS, IRD, IMBE, Marseille, France, Chemin de la Batterie des Lions, 13007, Marseille, France Saint-Petersburg State University, Faculty of Biology, Department of Embryology, Saint-Petersburg, Russia
Iowa State University, Department of Ecology, Evolution, and Organismal Biology, 251 Bessey Hall, 2200 Osborn Dr., Ames, IA 50011-4009, USA
Porifera Chilean Patagonia Chondrillida cytology Halisarcidae new species SEM TEM ultrastructure Verongiomorpha

Abstract

A series of recent expeditions in fjords and canals of Southern Chilean Patagonia allowed the re-collection of Halisarca magellanica Topsent, 1901 and the discovery of a new species, Halisarca desqueyrouxae sp. nov. The material studied was collected at depths ranging from 3 to 30 m at latitudes comprised between 42° and 49°S. Both species share the same habitat and show a morphological plasticity, but differ in their colour. Halisarca magellanica is bright pink to whitish with three morphs whereas H. desqueyrouxae sp. nov. is light brown to beige with two morphs. An extensive investigation in TEM and SEM reveals several differences among cell types with inclusions between both species. Three distinct spherulous cells occur. Type 1 is shared by both species, Type 2 is occasional in H. magellanica but absent from H. desqueyrouxae sp. nov. Type 3 is rare in H. magellanica and occurs abundantly in half of the specimens of H. desqueyrouxae sp. nov. Granular cells are shared by both species but do not occur in all specimens. Microgranular cells are characteristic of H. magellanica. Both species also clearly differ by their endobiotic bacteria. Phylogenetic analysis of cox1 sequences places H. magellanica as a sister group to all other previously published Halisarca species sequences (9.1–9.7% difference) except H. harmelini, while H. desqueyrouxae sp. nov. is placed as a sister group to H. dujardini (2.3% difference).

 

References

  1. Alvizu, A., Díaz, M.C., Bastidas, C., Rützler, K., Thacker, R.W. & Márquez, L.M. (2013) A skeleton-less sponge of Caribbean mangroves: invasive or undescribed? Invertebrate Biology, 132 (2), 81–94.
    http://dx.doi.org/10.1111/ivb.12015

    Azevedo, F., Hajdu, E., Willenz, Ph. & Klautau, M. (2009) New records of Calcareous sponges (Porifera, Calcarea) from the Chilean coast. Zootaxa, 2072, 1–30.

    Bagby, R.M. (1970) The Fine Structure of Pinacocytes in the Marine Sponge Microciona prolifera (Ellis and Solander). Zeitschrift für Zellforschung, 105, 579–594.
    http://dx.doi.org/10.1007/BF00335430

    Bergquist, P.R. (1996) The Marine Fauna of New Zealand: Porifera: Demospongiae. Part 5. Dendroceratida and Halisarcida. New Zealand Oceanographic Institute Memoir, 107, 7–53.

    Bergquist, P.R. & Kelly, M. (2004) Taxonomy of some Halisarcida and Homosclerophorida (Porifera: Demospongiae) from the Indo-Pacific. New Zealand Journal of Marine and Freshwater Research, 38, 51–66.
    http://dx.doi.org/10.1080/00288330.2004.9517217

    Borojevic, R. & Lévi, C. (1964) Étude au microscope électronique des cellules de l'éponge Ophlitaspongia seriata (Grant), au cours de la réorganisation après dissociation. Zeitschrift für Zellforschung, 64, 708–725.
    http://dx.doi.org/10.1007/BF01258544

    Boury-Esnault, N., De Vos, L., Donadey, C. & Vacelet, J. (1990) Ultrastructure of Choanosome and Sponge Classification. In: Rützler, K.E. (Ed.), New Perspectives in Sponge Biology. Smithsonian Institution Press, Woods Hole, pp. 237–244.

    Boury-Esnault, N. & Rützler, K. (1997) Thesaurus of sponge morphology. Smithsonian Contributions to Zoology, 596, 1–55.
    http://dx.doi.org/10.5479/si.00810282.596

    Boury-Esnault, N., Solé-Cava, A.M. & Thorpe, J.P. (1992) Genetic and cytological divergence between colour morphs of the Mediterranean sponge Oscarella lobularis Schmidt (Porifera, Demospongiae, Oscarellidae). Journal of Natural History, 26, 271–284.
    http://dx.doi.org/10.1080/00222939200770131

    Burgsdorf, I, Erwin, PM, Lopez-Legentil, S, Cerrano, C, Haber, M, Frenk, S. & Steindler, L. (2014) Biogeography rather than association with cyanobacteria structures symbiotic microbial communities in the marine sponge Petrosia ficiformis. Frontiers in microbiology, 5, 529–529.
    http://dx.doi.org/10.3389/fmicb.2014.00529

    Burton, M. (1929) Porifera. Part II. Antarctic sponges. British Antarctic ("Terra Nova") Expedition, 1910. Natural History Report. London, British Museum (Natural History). Zoology, 6, 393–458, pls I–V.

    Burton, M. (1932) Sponges. Discovery Reports, pp. 237–392, pls 248–257.
    http://dx.doi.org/10.5962/bhl.part.24379

    Carvalho, M.S., Desqueyroux-Faúndez, R. & Hajdu, E. (2007) Phakellia sur sp. nov. (Demospongiae, Halichondrida, Axinellidae) from the lower slope off Cape Horn (South America), with a revision of the genus. Marine Biology Research, 3 (2), 109–116.
    http://dx.doi.org/10.1080/17451000701276063

    Carvalho, M.S., Desqueyroux-Faúndez, R. & Hajdu, E. (2011) Taxonomic notes on Poecillastra sponges (Astrophorida: Pachastrellidae), with the description of three new bathyal southeastern Pacific species. Scientia Marina, 75 (3), 477–492.
    http://dx.doi.org/10.3989/scimar.2011.75n3477

    Chen, W.T. (1976) Reproduction and speciation in Halisarca. In: Harrison, F.W. & Cowden, R.R. (Eds), Aspects of Sponge Biology. Academic Press. New York & London, pp. 113–139.
    http://dx.doi.org/10.1016/b978-0-12-327950-7.50014-2

    Cronquist, A. (1978) Once again, what is a species? In: Symposium on Biosystematics in Agriculture, Beltsville, MD (USA) 8–11 May 1977.

    D'Amico, F. (2005) A polychromatic staining method for epoxy embedded tissue: a new combination of methylene blue and basic fuchsine for light microscopy. Biotechnic & Histochemistry, 80 (5–6), 207–210.
    http://dx.doi.org/10.1080/10520290600560897

    DeBiasse, M.B., Richards, V.P. & Shivji, M.S. (2010) Genetic assessment of connectivity in the common reef sponge, Callyspongia vaginalis (Demospongiae: Haplosclerida) reveals high population structure along the Florida reef tract. Coral Reefs, 29 (1), 47–55.
    http://dx.doi.org/10.1007/s00338-009-0554-0

    Desqueyroux, R. & Moyano, H. (1987) Zoogeografia de Demospongias Chilenas—Zoogeography of Chilean Demospongiae. Boletin de la Sociedad de Biologia de Concepcion, 58, 39–66.

    Donadey, C. & Vacelet, J. (1977) Les cellules à inclusions de l'éponge Pleraplysilla spinifera (Schulze) (Démosponges, Dendroceratides). Archives de Zoologie expérimentale et générale, 118 (3), 273–284.

    Duran, S., Pascual, M. & Turon, X. (2004) Low levels of genetic variation in mtDNA sequences over the western Mediterranean and Atlantic range of the sponge Crambe crambe (Poecilosclerida). Marine Biology, 144 (1), 31–35.
    http://dx.doi.org/10.1007/s00227-003-1178-5

    Duran, S. & Rützler, K. (2006) Ecological speciation in a Caribbean marine sponge. Molecular Phylogenetics and Evolution, 40, 292–297.

    Ereskovsky, A.V. (2006) A new species of Oscarella (Demospongiae: Plakinidae) from the Western Sea of Japan. Zootaxa, 1376, 37–51.

    Ereskovsky, A.V. (2007) A new species of Halisarca (Demospongiae: Halisarcida) from the Sea of Okhotsk, North Pacific. Zootaxa, 1432, 57–66.

    Ereskovsky, A.V., Gonobobleva, E. & Vishnyakov, A. (2005) Morphological evidence for vertical transmission of symbiotic bacteria in the viviparous sponge Halisarca dujardini Johnston (Porifera, Demospongiae, Halisarcida). Marine Biology, 146 (5), 869–875.
    http://dx.doi.org/10.1007/s00227-004-1489-1

    Ereskovsky, A.V., Lavrov, D.V., Boury-Esnault, N. & Vacelet, J. (2011) Molecular and morphological description of a new species of Halisarca (Demospongiae: Halisarcida) from Mediterranean Sea and a redescription of the type species Halisarca dujardini. Zootaxa, 2768, 5–31.

    Ereskovsky, A.V., Lavrov, D.V. & Willenz, Ph. (2014) Five new species of Homoscleromorpha (Porifera) from the Caribbean Sea and re-description of Plakina jamaicensis. Journal of the Marine Biological Association of the United Kingdom, 94 (02), 285–307.
    http://dx.doi.org/10.1017/S0025315413000295

    Ereskovsky, A.V., Sanamyan, K. & Vishnyakov, A.E. (2009) A new species of the genus Oscarella (Porifera: Homosclerophorida: Plakinidae) from the North-West Pacific. Cahiers de Biologie Marine, 50, 369–381.

    Erpenbeck, D., Sutcliffe, P., Cook, S., Dietzel, A., Maldonado, M., van Soest, R.W.M, Hooper, J.N.A., Wo¨rheide, G. (2012) Horny sponges and their affairs: on the phylogenetic relationships of keratose sponges. Molecular Phylogenetics and Evolution, 63 (3), 809–816.
    http://dx.doi.org/10.1016/j.ympev.2012.02.024

    Esteves, E.L., Lôbo-Hajdu, G. & Hajdu, E. (2007) Three new species of Crambe (Crambeidae: Poecilosclerida: Demospongiae) from the south-eastern Pacific, with a review of morphological characters for the genus. Journal of the Marine Biological Association of the United Kingdom, 87 (06), 1367–1378.
    http://dx.doi.org/10.1017/s0025315407058110

    Flot, J.-F. (2015) Species Delimitation's Coming of Age. Systematic Biology, 64 (6), 897–899.
    http://dx.doi.org/10.1093/sysbio/syv071

    Gaino, E., Burlando, B. & Buffa, P. (1986) The vacuolar cells of Oscarella lobularis (Porifera, Demospongiae): Ultrastructural organization, origin, and function. Journal of Morphology, 188, 29–37.
    http://dx.doi.org/10.1002/jmor.1051880104

    Gazave, E., Lavrov, D.V., Cabrol, J., Renard, E., Rocher, C., Vacelet, J., Adamska, M., Borchiellini, C. & Ereskovsky, A.V. (2013) Systematics and molecular phylogeny of the family Oscarellidae (homoscleromorpha) with description of two new Oscarella species. PLoS ONE, 8 (5), e63976.
    http://dx.doi.org/10.1371/journal.pone.0063976

    Gloeckner, V., Hentschel, U., Ereskovsky, A.V. & Schmitt, S. (2013) Unique and species-specific microbial communities in Oscarella lobularis and other Mediterranean Oscarella species (Porifera: Homoscleromorpha). Marine Biology, 160 (4), 781–791.
    http://dx.doi.org/10.1007/s00227-012-2133-0

    Gonobobleva, E. & Maldonado, M. (2009) Choanocyte ultrastructure in Halisarca dujardini (Demospongiae, Halisarcida). Journal of Morphology, 270 (5), 615–627.
    http://dx.doi.org/10.1002/jmor.10709

    Hajdu, E. & Desqueyroux-Faúndez, R. (2008) A reassessment of the phylogeny and biogeography of Rhabderemia Topsent, 1890 (Rhabderemiidae, Poecilosclerida, Demospongiae). Revue Suisse de Zoologie, 115 (2), 377–395.

    Hajdu, E., Desqueyroux-Faúndez, R., Carvalho, M.D.S., Lôbo-Hajdu, G. & Willenz, Ph. (2013) Twelve new Demospongiae (Porifera) from Chilean fjords, with remarks upon sponge-derived biogeographic compartments in the SE Pacific. Zootaxa, 3744 (1), 1–64.
    http://dx.doi.org/10.11646/zootaxa.3744.1.1

    Hajdu, E., Desqueyroux-Faúndez, R. & Willenz, Ph. (2006) Clathria (Cornulotrocha) rosetafiordica sp. nov. from a south-east Pacifc fjord (Chilean Patagonia) (Microcionidae: Poecilosclerida: Demospongiae: Porifera). Journal of the Marine Biological Association of United Kingdom, 86, 957–961.
    http://dx.doi.org/10.1017/S0025315406013920

    Hebert, P.D.N., Ratnasingham, S. & deWaard, J.R. (2003) Barcoding Animal Life: Cytochrome c Oxidase Subunit 1 Divergences among Closely Related Species. Proceedings: Biological Sciences, Vol. 270, Supplement: Biology Letters, S96–S99.
    http://dx.doi.org/10.1098/rsbl.2003.0025

    Johnston, G. (1842) A History of British Sponges and Lithophytes. In. Lizars, W.H., Edinburgh, pp. i–xii, 1–264, pls I–XXV.
    http://dx.doi.org/10.5962/bhl.title.51495

    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 772–780.
    http://dx.doi.org/10.1093/molbev/mst010

    Koltun, V.M. (1976) Porifera–Part 1: Antarctic Sponges. Report B.A.N.Z. Antarctic Research Expedition 1929–1931 Series B (Zoology and Botany), 9 (4), 147–198.

    Korotkova, G.P. & Ermolina, N.O. (1986) The participation of specialized amoebocytes in the sexual embryogenesis of the White Sea sponge Halisarca dujardini Johnston. Biologicheskie Nauki, 1986 (5), 48–53.

    Lancellotti, D.A. & Vasquez, J.A. (2000) Zoogeografía de macroinvertebrados bentónicos de la costa de Chile: contribución para la conservación marina. Revista Chilena de Historia Natural, 73, 99–129.
    http://dx.doi.org/10.4067/S0716-078X2000000100011

    Lavrov, D.V., Maikova, O.O., Pett, W. & Belikov, S.I. (2012) Small inverted repeats drive mitochondrial genome evolution in Lake Baikal sponges. Gene, 505 (1), 91–99.
    http://dx.doi.org/10.1016/j.gene.2012.05.039

    Lavrov, D.V., Wang, X. & Kelly, M. (2008) Reconstructing ordinal relationships in the Demospongiae using mitochondrial genomic data. Molecular Phylogenetics and Evolution, 49 (1), 111–124.
    http://dx.doi.org/10.1016/j.ympev.2008.05.014

    Lévi, C. (1953) Travaux de la station Biologique de Roscoff. XVII. Halisarca metschnikovi, n. sp. éponge sans squelette des côtes de France. Ses caractères embryologiques. Archives de Zoologie expérimentale et générale (Notes et Revues), 90 (2), 87–91.

    Lévi, C. (1956) Étude des Halisarca de Roscoff. Embryologie et systématique des démosponges. Archives de Zoologie expérimentale et générale, 93 (1), 1–181.

    Lévi, C. & Lévi, P. (1976) Embryogenèse de Chondrosia reniformis (Nardo), démosponge ovipare, et transmission des bactéries symbiotiques. Annales des Sciences Naturelles. Zoologie et Biologie Animale, 18 (4), 367–380.

    Loh, T.-L., López-Legentil, S., Song, B. & Pawlik, J.R. (2012) Phenotypic variability in the Caribbean Orange Icing sponge Mycale laevis (Demospongiae: Poecilosclerida). Hydrobiologia, 687 (1), 205–217.
    http://dx.doi.org/10.1007/s10750-011-0806-1

    Lopes, D.A., Bravo, A. & Hajdu, E. (2011) New carnivorous sponges (Cladorhizidae: Poecilosclerida: Demospongiae) from off Diego Ramírez Archipelago (south Chile), with comments on taxonomy and biogeography of the family. Invertebrate Systematics, 25, 407–443.
    http://dx.doi.org/10.1071/IS11015

    López Gappa, J. & Landoni, N.A. (2005) Biodiversity of Porifera in the Southwest Atlantic between 35°S and 56°S. Revista dei Museo Argentino de Ciencias Naturales, 7 (2), 191–219.
    http://dx.doi.org/10.22179/REVMACN.7.328

    López-Legentil, S., Erwin, P.M., Henkel, T.P., Loh, T.-L. & Pawlik, J.R. (2010) Phenotypic plasticity in the Caribbean sponge Callyspongia vaginalis (Porifera: Haplosclerida). Scientia Marina, 74 (3), 445–453.
    http://dx.doi.org/10.3989/scimar.2010.74n3445

    Maldonado, M. (2007) Intergenerational transmission of symbiotic bacteria in oviparous and viviparous demosponges, with emphasis on intracytoplasmically-compartmented bacterial types. Journal of the Marine Biological Association of the United Kingdom, 87 (06), 1701–1713.
    http://dx.doi.org/10.1017/s0025315407058080

    Morrow, C. & Cardenas, P. (2015) Proposal for a revised classification of the Demospongiae (Porifera). Frontiers in Zoology, 12, 7–27.
    http://dx.doi.org/10.1186/s12983-015-0099-8

    Muricy, G., Boury-Esnault, N., Bézac, C. & Vacelet, J. (1996) Cytological evidence for cryptic speciation in Mediterranean Oscarella species (Porifera, Homoscleromorpha). Canadian Journal of Zoology, 74, 881–896.
    http://dx.doi.org/10.1139/z96-102

    Muricy, G. & Pearse, J.S. (2004) A New Species of Oscarella (Demospongiae: Plakinidae) from California. Proceedings of the California Academy of Sciences, Series 4, 55 (33), 598–612.

    Redmond, N.E., Morrow, C.C., Thacker, R.W., Diaz, M.D., Boury-Esnault, N., Cárdenas, P., Hajdu, E., Lobo-Hajdu, G., Picton, B.E., Pomponi, S.A., Kayal, E. & Collins, A.G. (2013) Phylogeny and systematics of demospongiae in light of new small-subunit ribosomal DNA (18S) sequences. Integrative and Comparative Biology, 53 (3), 388–415.
    http://dx.doi.org/10.1093/icb/ict078

    Reynolds, E.S. (1963) The use of lead citrate at high pH as an electron-opaque stain in electron microscopy. Journal of Cell Biology, 17 (1), 208–212.
    http://dx.doi.org/10.1083/jcb.17.1.208

    Ridley, S.O. (1881) XI Spongida. Horny and Siliceous Sponges of Magellan Straits, S.W. Chili, and Atlantic off S.W. Brazil. In: A. Günter (Ed.), Account of the Zoological Collections made during the Survey of H.M.S. ‘Alert’ in the Straits of Magellan and on the Coast of Patagonia. 1881. Proceedings of the Zoological Society of London. pp. 107–137, 140–141, pls X–XI.

    Ridley, S.O. & Dendy, D. (1887) Preliminary report on the Monaxonida collected by H.M.S. “Challenger” during the years 1873–1876. Report on the Scientific Results of the Voyage of H.M.S. “Challenger” 1873–1876 —Zoology, 20 (59), i–lxvii, 1–275, pls I–LI.

    Riesgo, A., Maldonado, M. & Durfort, M. (2008) Occurrence of somatic cells within the spermatic cysts of demosponges: A discussion of their role. Tissue & Cell, 40 (6), 387–396.
    http://dx.doi.org/10.1016/j.tice.2008.03.005

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Hohna, S. Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542.
    http://dx.doi.org/10.1093/sysbio/sys029

    Ruiz, C., Ivanišević, J., Chevaldonné, P., Ereskovsky, A.V., Boury-Esnault, N., Vacelet, J., Thomas, O.P. & Pérez, T. (2015) Integrative taxonomic description of Plakina kanaky, a new polychromatic sponge species from New Caledonia (Porifera: Homoscleromorpha). Marine Ecology, 36 (4). 1129–1143.
    http://dx.doi: 10.1111/maec.1220

    Saghai-Maroof, M.A., Soliman, K.M., Jorgensen, R.A. & Allard, R.W. (1984) Ribosomal DNA spacer-length polymorphisms in barley: Mendelian inheritance, chromosomal location, and population dynamics. Proceedings of the National Academy of Sciences of the United States of America, 81, 8014–8018.
    http://dx.doi.org/10.1073/pnas.81.24.8014

    Sarà, M., Balduzzi, A., Barbieri, M., Bavestrello, G. & Burlando, B. (1992) Biogeographic traits and checklist of Antarctic demosponges. Polar Biology, 12, 559–585.
    http://dx.doi.org/10.1007/BF00236980

    Schmidt, O. (1862) Die Spongien des adriatischen Meeres. In. Wilhelm Engelmann, Leipzig, pp. i–viii, 1–88, pls 81–87.

    Setiawan, E., de Voogd, N.J., Swierts, T., Hooper, J.N.A., Wörheide, G. & Erpenbeck, D. (2016) MtDNA diversity of the Indonesian giant barrel sponge Xestospongia testudinaria (Porifera: Haplosclerida)—implications from partial cytochrome oxidase 1 sequences. Journal of the Marine Biological Association of the United Kingdom, 96 (02), 323–332.
    http://dx.doi.org/10.1017/S0025315415001149

    Simpson, T.L. (1963) The Biology of the Marine Sponge Microciona prolifera (Ellis and Solander). I. A Study of Cellular Function and Differentiation. Journal of Experimental Zoology, 154 (1), 135–151.
    http://dx.doi.org/10.1002/jez.1401540109

    Simpson, T.L. (1968) The Structure and Function of Sponge Cells: New Criteria for the Taxonomy of Poecilosclerid Sponges (Demospongiae). Peabody Museum of Natural History Yale University Bulletin, 25, 1–125, pls 121–117.

    Simpson, T.L. (1984) The Cell Biology of Sponges. Springer-Verlag, New York, Berlin, Heidelberg, 662 pp.
    http://dx.doi.org/10.1007/978-1-4612-5214-6

    Sollas, W.J. (1885) A Classification of the Sponges. Annals and Magazine of Natural History (5), 16 (95), 395–395.
    http://dx.doi.org/10.1080/00222938509459901

    Spurr, A.R. (1969) A low-viscosity epoxy resin embedding medium for electron microscopy. Journal of Ultrastructure Research, 26, 31–43.
    http://dx.doi.org/10.1016/S0022-5320(69)90033-1

    Sukhodolskaya, A.N. & Krasukevitch, T.N. (1984) The effect of inozine on the development of Halisarca dujardini from small fragments of the body. Vestnik of Leningrad University, 3, 56–62.

    Thiele, J. (1905) Die Kiesel- und Hornschwämme der Sammlung Plate. Supplement 6 (Fauna Chiliensis III). Zoologische Jahrbücher, 1905, 407–496, pls 427–433.

    Topsent, E. (1901) Spongiaires. Résultats du voyage du S.Y. Belgica en 1897–99 sous le commandement de A. de Gerlache de Gomery. Expédition antarctique belge. Zoologie, 4, 1–54.

    Uriz, M.J. & Turon, X. (2012) Sponge ecology in the molecular era. Advances in Marine Biology, 61, 345–410.

    Vacelet, J. (1970) Description de cellules à bactéries intranucléaires chez des éponges Verongia. Journal de Microscopie, 9 (3), 333–346.

    Vacelet, J., Boury-Esnault, N., De Vos, L. & Donadey, C. (1989) Comparative Study of the Choanosome of Porifera: 2. The Keratose Sponges. Journal of Morphology, 201 (2), 119–129.

    Vacelet, J. & Donadey, C. (1987) A new species of Halisarca (Porifera, Demospongiae) from the Caribbean, with remarks on the cytology and affinities of the genus. In: Jones, W.C. (Ed.), European Contributions to the Taxonomy of Sponges. Sherkin Island Marine Station, Sherkin Island, pp. 5–12.

    Van Soest, R.W.M., Boury-Esnault, N., Vacelet, J., Dohrmann, M., Erpenbeck, D., De Voogd, N.J., Santodomingo, N., Vanhoorne, B., Kelly, M., Hooper, J.N.A (2012) Global Diversity of Sponges (Porifera). PLoS ONE, 7 (4), 1–23.

    Van Soest, R.W.M., Boury-Esnault, N., Hooper, J., Rützler, K., de Voogd, N.J., Alvarez de Glasby, B., Hajdu, E., Pisera, A.B., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R., Klautau, M., Picton, B., Kelly, M., Vacelet, J., Dohrmann, M., Díaz, M.-C., Cárdenas, P. & Carballo, J.L (2016) World Porifera database. Accessed at http://www.marinespecies.org/porifera (2016-10-07)

    Vargas, S., Schuster, A., Sacher, K., Büttner, G., Schätzle, S., Läuchli, B., Hall,K., Hooper, J.N.A., Erpenbeck, D. & Wörheide, G. (2012) Barcoding sponges: an overview based on comprehensive sampling. PLoS ONE, 7 (7), e39345.
    https://doi.org/10.1371/journal.pone.0039345

    Vishnyakov, A.E. & Ereskovsky, A.V. (2009) Bacterial symbionts as an additional cytological marker for identification of sponges without a skeleton. Marine Biology, 156 (8), 1625–1632.

    Volkova, M.A. & Zolotareva, G.A. (1981) The development of Halisarca dujardini Johnston from conglomerates of somatic cells. In: G. P. Korotkova (Ed), Morphogenesis in sponges. Leningrad University Press. Leningrad, pp. 74–92.

    Whalan, S., de Nys, R., Smith-Keune, C., Evans, B.S., Battershill, C. & Jerry, D.R. (2008) Low genetic variability within and among populations of the brooding sponge Rhopaloeides odorabile on the central Great Barrier Reef. Aquatic Biology, 3, 111–119.

    Willenz, Ph., Hajdu, E., Desqueyroux-Faúndez, R., Lôbo-Hajdu, G. & Carvalho, M.S. (2009) Class Demospongiae. In: V.F. Häussermann, G. (Ed.), Marine Benthic Fauna of Chilean Patagonia. Nature in Focus. Santiago de Chile, pp. 107–157.

    Wilson, H.V. & Penney, J.T. (1930) The regeneration of sponges (Microciona) from dissociated cells. Journal of Experimental Zoology, 56 (1), 73–137, pls 131–135.

    Wörheide, G. (1998) The reef cave dwelling ultraconservative coralline demosponge Astrosclera willeyana Lister 1900 from the Indo-Pacific. Micromorphology, ultrastructure, biocalcification, isotope record, taxonomy, biogeography, phylogeny. Facies, 38, 1–88.

    Wörheide, G. (2006) Low variation in partial cytochrome oxidase subunit I (COI) mitochondrial sequences in the coralline demosponge Astrosclera willeyana across the Indo-Pacific. Marine Biology, 148, 907–912.