Abstract
The sponge Tethya leysae sp. nov. (Porifera, Demospongiae, Hadromerida, Tethyidae) is a new species from the Canadian Northeast Pacific. Its type locality is the infralittoral around Ohiat Islet, Barkely Sound, Vancouver Island in Canada. Tethya leysae sp. nov. is a medium-sized spherical sponge with a verrucose surface, orange-yellow to light red alive and white with a greyish core in ethanol. The highly variable oxyspherasters (25-115 µm in size, R/C 0.34-0.69) are densely scattered almost throughout the entire cortex. The micrasters comprise acanthoxyspherasters, acanthostrongylasters, acanthotylasters and small oxyspherasters, which are present at low densities throughout the sponge and form a dense layer associated with the exopinacoderm. The megascleres and the auxiliary megascleres comprise oxeas and strongyloxeas. The new species is clearly distinguishable from the closely related T. californiana by the absence of spherules among the micrasters, the lack of an alveolar cortex and the extremely high density of megasters in the cortex. In addition, the R/C values of the megasters differ between the two species and the oxyspherasters of T. leysae sp. nov. rarely display bent rays. The morphological differences between the species are confirmed by nucleotide and amino acid substitutions within the cytochrome oxidase subunit I (COI) mtDNA gene. Phylogenetic analyses group T. leysae sp. nov. with T. californiana, T. actinia and T. minuta, which together form a sister group to a MediterraneanNorth Atlantic species cluster. Morphological analyses of the skeleton included x-ray microtomography (µCT) and virtual 3D reconstruction, which was used for the first time in conjunction with the description of a new sponge species. Microtomography permitted the visualization and analysis of spicules within the skeletal context or isolated in silico. The method represents a valuable extension to the sponge taxonomist’s toolbox since it allows morphometric measurements in 3D. µCT will thus supplement classical morphological methods such as light and scanning electron microscopy.References
Austin, B., Ott, B., McDaniel, N. & Romagosa, P. (1999–2007) An annotated checklist of marine invertebrates in the cold temperate northeast Pacific, Khoyatan Marine Laboratory. Available from: http://www.mareco.org/KML/Projects/NEsponges.asp (12.6.2009).
Beckmann, F. (2008) Neutron and Synchrotron-radiation based imaging for applications in materials science – from Macro- to Nanotomography. In: Reimers, W., Pyzalla, A.R., Schreyer, A. & Clemens, H. (Eds.) Neutrons and synchrotron radiation in engineering materials science. Wiley-VCH, Weinheim, pp. 287–370.
Beckmann, F., Donath, T., Dose, T., Lippmann, T., Martins, R.V., Metge, J. & Schreyer, A. (2004) Microtomography using synchrotron radiation at DESY: current status and future developments. Proceedings of SPIE, 5535, 1–10.
Bell, J.J. & Barnes, D.K.A. (2001) Sponge morphological diversity: a qualitative preditor of species diversity? Aquatic Conservation: Marine And Freshwater Ecosystems, 11, 109–121.
Bergquist, P.R. & Bedford, J.J. (1978) The incidence of antibacterial activity in marine Demospongiae; Systematic and geographic considerations. Marine Biology, 46, 215–221.
Bergquist, P.R., Sinclair, M.M. & Hogg, J.J. (1970) Adaptation to intertidal existance, p. 247–271. In: Reproduction cycles and larval behaviour in Demospongia. Biology of Porifera. Fray, W.G. (ed.). Academic Press, London
Betz, O., Wegst, U., Weid, D., Heethoff, M., Helfen, L., Lee, W.-K. & Cloethens, P. (2007) Imaging applications of synchrotron X-ray phase-contrast microtomography in biological morphology and biomaterials science. I. General aspects of the technique and its advantages in the analysis of millimetre-sized arthropod structure. Journal of Microscopy, 227, 51–71.
Beutel, R.G., Ge, S. & Hörnschemeyer, T. (2008) On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics, 24, 270–298.
Beutel, R.G. & Haas, A. (1998) Larval head morphology of Hydroscapha natans (Coleoptera, Myxophaga) with reference to miniaturization and the systematic position of Hydroscaphidae. Zoomorphology, 118, 103–116.
Blanquer, A. & Uriz, M.J. (2007) Cryptic speciation in marine sponges evidenced by mitochondrial and nuclear genes: A phylogenetic approach. Molecular Phylogenetics and Evolution, 45, 392–397.
Blanquer, A. & Uriz, M.J. (2008) 'A posteriori' searching for phenotypic characters to describe new cryptic species of sponges revealed by molecular markers (Dictyonellidae: Scopalina). Invertebrate Systematics, 22, 489–502.
Blanquer, A., Uriz, M.J. & Agell, G. (2008) Hidden diversity in sympatric sponges: adjusting life-history dynamics to share substrate. Marine Ecology Progress Series, 371, 109–115.
Bonse, U. & Busch, F. (1996) X-Ray Computed Microtomography (µCT) using Synchrotron Radiation (SR). Progress in Biophysics and Molecular Biology, 65, 133–169.
Boury-Esnault, N., Klautau, M., Bézac, C., Wulff, J. & Solé-Cava, A.M. (1999) Comparative study of putative conspecific sponge populations from both sides of the Isthmus of Panama. Journal of the Marine Biological Association of the United Kingdom, 79, 39–50.
Boury-Esnault, N., Marschal, C., Kornprobst, J.M. & Barnathan, G. (2002) A new species of Axinyssa Lendenfeld, 1897 (Porifera, Demospongiae, Halichondrida) from the Senegalese coast. Zootaxa, 117, 1–8.
Boury-Esnault, N., Pansini, M. & Uriz, M.J. (1993) Cosmopolitism in sponges: The "complex" Guitarra fimbriata with description of a new species of Guitarra from the northeast Atlantic. Scientia Marina, 57, 367–373.
Bowerbank, J.S. (1873) Contributions to a General History of the Spongiadae. Part IV. Proceedings of the Zoological Society of London, 1873, 3–25, pls. I–IV.
Burton, M. (1924) A revision of the sponge family Donatiidae. Proceeding of the zoological Society of London, 4, 1033–1046.
Corriero, G., Balduzzi, A. & Sara, M. (1989) Ecological differences in the distribution of two Tethya (Porifera, Demospongiae) species coexisting in a Mediterranean coastal lagoon. Marine Ecology, 10, 303–315.
Costeloe, J., Keegan, B.F. & Könnecker, G.F. (1986) Rocky subtidal assemblages on the west coast of Ireland. Hydrobiologia, 142, 97–111.
de Laubenfels, M.W. (1932) The marine and fresh-water sponges of California. Proceedings of the United States National Museum, 81, 1–140.
Desqueyroux-Faúndez, R. & van Soest, R.W.M. (1997) Shallow water demosponges of the Galápagos Islands. Revue suisse de Zoologie, 104, 379–467.
Drummond, A., Ashton, B., Cheung, M., Heled, J., Kearse, M., Moir, R., Stones-Havas, S., Thierer, T. & Wilson, A. (2009) Geneious 4.5 available from www.geneious.com
Duran, S. & Rützler, K. (2006) Ecological speciation in a Caribbean marine sponge. Molecular Phylogenetics and Evolution, 40, 292–297.
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucl. Acids Res., 32, 1792–1797.
Erpenbeck, D., Voigt, O., Gültas, M. & Wörheide, G. (2008) The Sponge Genetree Server - poviding a phylogenetic backbone for poriferan evolutionary studies. Zootaxa, 1939, 58–60.
Fletcher, G. (2009) Racerocks Ecological Reserve, Lester B. Pearson College of the Pacific. Available from: http://www.racerocks.com (1.7.2009).
Folmer, O., Black, M., Hoch, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Forster, G.R. (1958) Underwater observations on the fauna of shallow rocky areas in the neighbourhood of Plymouth. Journal of the Marine Biological Association of the United Kingdom, 37, 473–482.
Francis, R.C., Hare, S.R., Hollowed, A.B. & Wooster, W.S. (1998) Effects of interdecadal climate variability on the oceanic ecosystems of the NE Pacific. Fisheries Oceanography, 7, 1–21.
GBIF. (2009) Tethya californiana de Laubenfels, 1932; In database: Gwaii Haanas Invertebrates. Biodiversity occurrence data provided by: OBIS Canada (Accessed through GBIF Data Portal, www.gbif.net, 2009-06-16),
Heim, I., Nickel, M. & Brümmer, F. (2007a) Phylogeny of the genus Tethya (Tethyidae: Hadromerida: Porifera): molecular and morphological aspects. Journal of the Marine Biological Association of the UK, 87, 1615–1627.
Heim, I., Nickel, M., Picton, B. & Brümmer, F. (2007b) Description and molecular phylogeny of Tethya hibernica sp. nov. (Porifera, Demospongiae) from Northern Ireland with remarks on the European species of the genus Tethya. Zootaxa, 1595, 1–15.
Hooper, J.N.A. & Van Soest, R.W.M. (2002a) Class Demospongiae Sollas,1885. In: Hooper, J.N.A. & van Soest, R.M.W. (Eds.) Systema Porifera: A Guide to the Classification of Sponges. Kluwer Academic/Plenum Publishers, New York, pp. 15–18.
Hooper, J.N.A. & Van Soest, R.W.M. (2002b) Order Hadromerida Topsent, 1894. In: Hooper, J.N.A. & van Soest, R.M.W. (Eds.) Systema Porifera: A Guide to the Classification of Sponges. Kluwer Academic/Plenum Publishers, New York, pp. 169–172.
Huesman, R., Gullberg, G., Greenberg, W. & TF, B. (1977) RECLBL Library users manual: Donner algorithms for reconstruction tomography. Lawrence Berkeley Laboratory, University of California, Livermore, pp.
Kelly-Borges, M. & Pomponi, S. (1992) The simple fool's guide to sponge taxonomy. Harbor Branch Oceanographic Institution, Fort Pierce, Florida
Kennett, D.J., Ingram, B.L., Erlandson, J.M. & Walker, P. (1997) Evidence for temporal fluctuations in marine radiocarbon reservoir ages in the Santa Barbara Channel, Southern California. Journal of Archaeological Science, 24, 1051–1059.
Klautau, M., Solé-Cava, A.M. & Borojecic, R. (1994) Biochemical systematics of sibling sympatric species of Clathrina (Porifera: Calcarea). Biochemical Systematics And Ecology, 22, 367–375.
Kleinteich, T., Beckmann, F., Herzen, J., Summers, A.P. & Haas, A. (2008) Applying x-ray tomography in the field of vertebrate biology: form, function, and evolution of the skull of caecilians (Lissamphibia: Gymnophiona), p. 70780D-10. In: Developments in X-Ray Tomography VI. Vol. 7078. SPIE, San Diego, CA, USA
Kumar, A. & Patterson, R.T. (2002) Dinoflagellate cyst assemblages from Effingham Inlet, Vancouver Island, British Columbia, Canada. Palaeogeography, Palaeoclimatology, Palaeecology, 180, 187–206.
McLain, D.R. & Thomas, D.H. (1983) Year-to-year fluctuations of the California Countercurrent and effects on marine organisms. California Cooperative Oceanic Fischeries Investigations, 24, 165–181.
Miller, K., Alvarez, B., Battershill, C., Northcote, P. & Parthasarathy, H. (2001) Genetic, morphological, and chemical divergence in the sponge genus Latrunculia (Porifera: Demospongiae) from New Zealand. Marine Biology, 139, 235–250.
Murch, A. (2009) The BC Marinelife Index, Available at http://elasmodiver.com/BCMarinelife/bc_marinelife_index.htm (accessed 2009-06-16).
Nichols, S.A. & Barnes, P.A.G. (2005) A molecular phylogeny and historical biogeography of the marine sponge genus Placospongia (Phylum Porifera) indicate low dispersal capabilities and widespread crypsis. Journal of Experimental Marine Biology and Ecology, 323, 1–15.
Nickel, M., Bullinger, E. & Beckmann, F. (2006a) Functional morphology of Tethya species (Porifera): 2. Three-dimensional morphometrics on spicules and skeleton superstructures of T. minuta. Zoomorphology, 125, 225–239.
Nickel, M., Donath, T., Schweikert, M. & Beckmann, F. (2006b) Functional morphology of Tethya species (Porifera): 1. Quantitative 3D-analysis of Tethya wilhelma by synchrotron radiation based X-ray microtomography. Zoomorphology, 125, 209–223.
Nickel, M., Hammel, J., Herzen, J., Bullinger, E. & Beckmann, F. (2008) High density resolution synchrotron radiation based x-ray microtomography (SR μCT) for quantitative 3D-morphometrics in zoological sciences. Proceedings of SPIE, 7078, 7078W.
Nylander, J.A.A., Ronquist, F., Huelsenbeck, J.P. & Aldrey, J.L.N. (2004) Bayesian phylogenetic analysis of combined data. Systematic Biology, 53, 47–67.
Page, R.D.M. (1996) TreeView: An application to display phylogenetic trees on personal computers. Computer Applications in the Biosciences, 12, 357–358.
Rasband, W.S. (1997–2009) ImageJ. National Institutes of Health, Bethesda, Maryland, USA; http://rsb.info.nih.gov/ij/
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Sarà, M. (1998) A Biogeographic and Evolutionary Survey of the Genus Tethya (Porifera, Demospongiae). In: Watanabe, Y. & Fusetani, N. (Eds.) Sponge Sciences. Multidisciplinary Perspectives. Springer, Tokyo, pp. 83–94.
Sarà, M. (2002) Family Tethyidae Gray 1848. In: Hooper, J.N.A. & van Soest, R.M.W. (Eds.) Systema Porifera: A Guide to the Classification of Sponges. Kluwer Academic/Plenum Publishers, New York, pp. 245–265.
Sarà, M., Bavestrello, G. & Mensi, P. (1992) Redescription of Tethya norvegica Bowerbank (Porifera, Demospongiae), with remarks on the genus Tethya in the North East Atlantic. Zoologica Scripta, 21, 211–216.
Sarà, M. & Corriero, G. (1993) Redescription of Tethya californiana De Laubenfels as a valid species for Tethya aurantia var. californiana (Porifera, Demospongiae). Ophelia, 37, 203–211.
Sara, M. & Gaino, E. (1987) Interspecific variation in arrangement and morphology of micrasters of Tethya species (Porifera, Demospongiae). Zoomorphology, 107, 313–317.
Sarà, M., Gómez, P. & Sarà, A. (2001) East Pacific Mexican Tethya (Porifera: Demospongiae) with descriptions of five new species. Proceedings of the Biological Society of Washington, 114, 794–821.
Sarà, M. & Melone, N. (1965) Una nuova specie del genere Tethya, T. citrina sp. n. dal Mediterraneo (Porifera, Demospongiae). Atti della Società Peloritana di Scienze fisiche matematiche e naturali, 11, 123–138.
Sarà, M., Mensi, P., Manconi, R., Bavestrello, G. & Balletto, E. (1989) Genetic variability in Mediterranean Sea populations of Tethya (Porifera: Demospongiae). In: Ryland, J.S. & Tyler, P.A. (Eds.) Reproduction, genetics and distributions of marine organisms 293–298. Olsen & Olsen, Fredensborg, pp. 293–298.
Sarà, M. & Sarà, A. (2004) A revision of Australian and New Zealand Tethya (Porifera:Demospongiae) with a preliminary analysis of species grouping. Invertebrate Systematics, 18, 117–156.
Schaefer, S.A. (2003) Relationships of Lithogenes villosus Eigenmann, 1909 (Siluriformes, Loricariidae): Evidence from High-Resolution Computed Microtomography. American Museum Novitates, 1–55.
Scripps Institution of Oceanography. (1981) Surface water temperatures at shore stations. United States West Coast 1980, p. 45. Marine Life Research Committee and Marine Life Research Group, University of California, Scripps Institution of Oceanography, La Jolla, California
Sim, C.J. (1981) A taxonomic study on the marine sponges in Korea [R.], 2: Hadromerida. Korean Journal of Zoology, 24, 9–17.
Solé-Cava, A.M., Klautau, M., Boury-Esnault, N., Borojecic, R. & Thorpe, J.P. (1991) Genetic evidence for cryptic speciation in allopatric populations of two cosmopolitan species of the calcareous sponge genus Clathrina. Marine Biology, 111, 381–386.
Solé-Cava, A.M. & Thorpe, J.P. (1986) Genetic differentiation between morphotypes of the marine sponges Suberites ficus (Demospongiae: Hadromerida). Marine Biology, 93, 247–253.
Solé-Cava, A.M. & Thorpe, J.P. (1987) The use of electrophoresis in sponge taxonomy. NATO ASI Series - Taxonomy of Porifera, G 13, 243–258.
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A Rapid Bootstrap Algorithm for the RAxML Web Servers. Systematic Biology, 57, 758 – 771.
Tatusov, T. & Tatusov, R. (1988–2009) ORF Finder (Open Reading Frame Finder), National Center for Biotechnology Information (NCBI) Available from: http://www.ncbi.nlm.nih.gov/gorf/gorf.html
Thiele, J. (1905) Die Kiesel- und Hornschwämme der Sammlung Plate. Zoologische Jahrbücher, Supplement 6, 107–496.
Topsent, E. (1918) Eponges de San Thome. Essai sul les genres Spirastrella, Donatia et Chondrilla. Archives de zoologie experimentale et generale, 57, 535–618.
Van Der Meijden, A., Boistel, R., Gerlach, J., Ohler, A., Vences, M. & Meyer, A. (2007) Molecular phylogenetic evidence for paraphyly of the genus Sooglossus, with the description of a new genus of Seychellean frogs. Biological Journal of the Linnean Society, 91, 347–359.
Van Soest, R.W.M., Boury-Esnault, N., Hooper, J.N.A., Rützler, K., de Voogd, N.J., Alvarez, B., Hajdu, E., Pisera, A.B., Vacelet, J., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R. & Klautau, M. (2009) World Porifera database. Available online at http://www.marinespecies.org/porifera. Consulted on 2009-06-19. Available from:
Wirkner, C.S. & Prendini, L. (2007) Comparative morphology of the hemolymph vascular system in scorpions-A survey using corrosion casting, MicroCT, and 3D-reconstruction. Journal of Morphology, 268, 401–413.
Wörheide, G., Degnan, B.M., Hooper, J.N.A. & Reitner, J. (2000) Phylogeography and taxonomy of the Indo-Pacific reef cave dwelling coralline demosponge Astrosclera 'willeyana': new data from nuclear internal transcribed spacer sequences, p. 339–346. In: 9th International Coral Reef Symposium. Vol. 1. Moosa, K. (ed.), Bali
Wulff, J.L. (2006) Sponge systematics by starfish: predators distinguish cryptic sympatric species of Caribbean fire sponges, Tedania ignis and Tedania klausi n. sp. (Demospongiae, Poecilosclerida). Biological Bulletin, 211, 83–94.
Xavier, J.R., Rachello-Dolmen, P.G., Parra-Velandia, F., Schöneberge, C.H.L., Breeuwer, J.A.J. & Van Soest, R.W.M. (2009) Molecular evidence of cryptic speciation in the "cosmopolitan"excavating sponge Cliona celata (Porifera, Clionaidae). under review.
Zollikofer, C. & Ponce de León, M. (2005) Virtual Reconstruction. A Primer in Computer-Assisted Paleontology and Biomedicine. Wiley & Sons, New York, pp.