Hostname: page-component-cb9f654ff-c75p9 Total loading time: 0 Render date: 2025-09-07T12:41:31.653Z Has data issue: false hasContentIssue false
  • English
  • Français

Five new species of Homoscleromorpha (Porifera) from the Caribbean Sea and re-description of Plakina jamaicensis

Published online by Cambridge University Press:  10 April 2013

Alexander V. Ereskovsky ,
Dennis V. Lavrov  and
Philippe Willenz
Alexander V. Ereskovsky*
Affiliation:
Mediterranean Institute of Biodiversity and Ecology (IMBE), UMR 7263, Aix-Marseille Université, CNRS, IRD, Station Marine d'Endoume, Marseille, France Biological and Soil Science Faculty, Saint-Petersburg State University, Russia
Dennis V. Lavrov
Affiliation:
Department of Ecology, Evolution and Organismal Biology, Iowa State University, 253 Bessey Hall, Ames, IA, USA
Philippe Willenz
Affiliation:
Department of Invertebrates, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, Belgium
*
Correspondence should be addressed to: A.V. Ereskovsky, Mediterranean Institute of Biodiversity and Ecology (IMBE), UMR 7263, Aix-Marseille Université, CNRS, IRD, Station Marine d'Endoume, Marseille, France email: alexander.ereskovsky@imbe.fr
Get access Rights & Permissions [Opens in a new window]

Abstract

Five new species of Homoscleromorpha (Porifera) of four genera, Oscarella, Plakortis, Plakina and Corticium, are described from vertical walls of reef caves at depths ranging from 23 to 28 m in the Caribbean Sea. Oscarella nathaliae sp. nov. has a leaf-like thinly encrusting, flat body, loosely attached to the substrate and a perforated, not lobate surface. Oscarella nathaliae sp. nov. contains two bacterial morphotypes and is characterized by two mesohylar cell types with inclusions. Plakortis myrae sp. nov. has diods of two categories: abundant large ones (83–119 μm long) and rare small ones (67–71 μm long) with sinuous, S-bent centres; triods Y- or T-shaped (18–5  μm long), and abundant microrhabds (5–12 μm long). Plakortis edwardsi sp. nov. has diods of one category with thick, sinuous, S-bent centres (110 to 128 μm long); triods T-shaped (actines 28–59 μm long). It is the only species of this genus showing small diods (22–31 μm long). Plakortis dariae sp. nov. has diods of two categories: large ones (67–112 μm long) and small, rare, irregular ones, slightly curved, often deformed with one end blunt (30–59 μm long); triods rare and regular (actines 20–44 μm long). Corticium diamantense sp. nov. has oscula situated near its border, regular non-lophose calthrops of one size-class, very rare tetralophose calthrops and candelabra with the fourth actine ramified basally in 4–5 microspined rays. In addition, a re-description of Plakina jamaicensis is based on newly collected material and the type specimen. Plakortis jamaicensis has a convoluted brain-like surface; well developed sub-ectosomal cavities; irregular sinuous diods, triods, calthrops, rare monolophose calthrops, rare dilophose calthrops, rare trilophose calthrops and common tetralophose calthrops. Molecular ‘barcoding’ sequences for mitochondrial cob are given for Plakortis edwardsi sp. nov., P. dariae sp. nov., Plakina jamaicensis and Corticium diamantense sp. nov. An identification key for all western Atlantic Homoscleromorpha is provided.

Keywords

taxonomyPoriferaHomoscleromorphabiodiversityOscarellaPlakortisPlakinaCorticiumJamaicaMartinique

Information

Type
Research Article
Information
Journal of the Marine Biological Association of the United Kingdom , Volume 94 , Issue 2 , March 2014 , pp. 285 - 307
Copyright
Copyright © Marine Biological Association of the United Kingdom 2013 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

Article purchase

Temporarily unavailable

References

REFERENCES

Aiello, A., Fattorusso, E., Magno, S., Mayol, L. and Menna, M. (1990) Isolation of two novel 5 alpha, 6 alpha-epoxy-7-ketosterols from the encrusting Demospongia Oscarella lobularis. Journal of Natural Products 53, 487–491.Google Scholar
Bergquist, P.R. (1978) Sponges. London: Hutchinson.Google Scholar
Berrué, F., Thomas, O.P., Bon, C.F.-L., Reyes, F. and Amade, P. (2005) New bioactive cyclic peroxides from the Caribbean marine sponge Plakortis zyggompha. Tetrahedron 61, 11843–11849.Google Scholar
Blanquer, A. and 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.Google Scholar
Borbone, N., De Marino, S., Iorizzi, M., Zollo, F., Debitus, C., Esposito, G. and Iuvone, T. (2002) Minor steroidal alkaloids from the marine sponge Corticium sp. Journal of Natural Products 65, 1206–1209.Google Scholar
Borchiellini, C., Chombard, C., Manuel, M., Alivon, E., Vacelet, J. and Boury-Esnault, N. (2004) Molecular phylogeny of Demospongiae: implications for classification and scenarios of character evolution. Molecular Phylogenetics and Evolution 32, 823–837.Google Scholar
Boury-Esnault, N. (1973) Résultats scientifiques des campagnes de la ‘Calypso'. Campagne de la Calypso au large des côtes atlantiques de l'Amérique du Sud (1961–1962). I. 29. Spongiaires. Annales de l'Institut Océanographique 49 (Supplement 10), 263–295.Google Scholar
Boury-Esnault, N., Muricy, G., Gallissian, M.-F. and Vacelet, J. (1995) Sponges without skeleton: a new Mediterranean genus of Homoscleromorpha (Porifera, Demospongiae). Ophelia 43, 25–43.Google Scholar
Boury-Esnault, N., Solé-Cava, A.M. and 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.Google Scholar
Carter, H.J. (1879) Contributions to our knowledge of the Spongida. Annals and Magazine of Natural History (5) 3, 284–304, 343–360.Google Scholar
Cruz-Barraza, J.A. and Carballo, J.L. (2005) First record of Plakortis Schulze (Porifera: Homosclerophorida) from the Northeast Pacific coast, with the description of Plakortis albicans sp. nov. Zootaxa 868, 1–12.Google Scholar
De Marino, S., Iorizzi, M., Zollo, F., Roussakis, C. and Debitus, C. (1999) Plakinamines C and D and three other new steroidal alkaloids from the sponge Corticium sp. European Journal of Organic Chemistry 1999, 697–701.Google Scholar
De Marino, S., Zollo, F., Iorizzi, M. and Debitus, C. (1998) A new steroidal alkaloid from a marine sponge Corticium sp. Tetrahedron Letters 39, 7611–7614.Google Scholar
Dendy, A. (1905) Report on the sponges collected by Professor Herdman at Ceylon, in 1902. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Reports 18, 57–246, pls I–XVI.Google Scholar
Desqueyroux-Faúndez, R. and van Soest, R.W.M. (1997) Shallow water Demosponges of the Galápagos Islands. Revue Suisse de Zoologie 104, 379–467.Google Scholar
Díaz, M.C. (2005) Common sponges from shallow marine habitats from Bocas del Toro region, Panama. Caribbean Journal of Science 41, 465–475.Google Scholar
Díaz, M.C. and Rützler, K. (2009) Biodiversity and abundance of sponges in Caribbean mangrove: Indicators of environmental quality. Smithsonian Contributions to the Marine Sciences 38, 151–172.Google Scholar
Díaz, M.C., Smith, K.P. and Rützler, K. (2004) Sponge species richness and abundance as indicators of mangrove epibenthic community health. Atoll Research Bulletin 518, 1–17.Google Scholar
Díaz, M.C. and van Soest, R.W.M. (1994) The Plakinidae: A Systematic review. In van Soest, R.W.M., van Kempen, T.M.G. and Braekman, J.-C. (eds) Sponges in time and space. Rotterdam: Balkema, pp. 93–109.Google Scholar
Ereskovsky, A.V. (2006) A new species of Oscarella (Demospongiae: Plakinidae) from the Western Sea of Japan. Zootaxa 1376, 37–51.Google Scholar
Ereskovsky, A.V. (2010) The comparative embryology of sponges. London: Springer-Verlag.Google Scholar
Ereskovsky, A., Borchiellini, C., Gazave, E., Ivanišević, J., Lapebie, P., Pérez, T., Renard, E. and Vacelet, J. (2009a) The homoscleromorph sponge Oscarella lobularis, a promising sponge model in evolutionary and developmental biology. BioEssays 31, 89–97.Google Scholar
Ereskovsky, A.V., Sanamyan, K. and Vishnyakov, A.E. (2009b) A new species of the genus Oscarella (Porifera: Homosclerophorida: Plakinidae) from the North-West Pacific. Cahiers de Biologie Marine 50, 369–381.Google Scholar
Faulkner, D.J. (2002) Marine natural products. Natural Products Reports 19, 1–49.Google Scholar
Gazave, E., Lapébie, P., Ereskovsky, A.V., Vacelet, J., Renard, E., Càrdenas, P. and Borchiellini, C. (2012) No longer Demospongiae: Homoscleromorpha formal nomination as a fourth class of Porifera. Hydrobiologia 687, 3–10.Google Scholar
Gazave, E., Lapébie, P., Renard, E., Vacelet, J., Rocher, C., Ereskovsky, A.V., Lavrov, D.V. and Borchiellini, C. (2010) Molecular phylogeny restores the supra-generic subdivision of Homoscleromorph sponges (Porifera, Homoscleromorpha). PLoS ONE 5, e14290.Google Scholar
Gloeckner, V., Hentschel, U., Ereskovsky, A.V. and Schmitt, S. (2012) Unique and species-specific microbial communities in Oscarella lobularis and other Mediterranean Oscarella species (Porifera: Homoscleromorpha). Marine Biology DOI 10.1007/s00227-012-2133-0.Google Scholar
Gochfeld, D.J. and Hamann, M.T. (2001) Isolation and biological evaluation of filiformin, plakortide F, and plakortone G from the Caribbean sponge Plakortis sp. Journal of Natural Products 64, 1477–1479.Google Scholar
Grant, R.E. (1836) Animal kingdom. In Todd, R.B. (ed.) The cyclopaedia of anatomy and physiology. Volume 1. London: Sherwood, Gilbert, and Piper, pp. 107–118.Google Scholar
Gray, J.E. (1867) Notes on the arrangement of sponges, with the descriptions of some new genera. Proceedings of the Zoological Society of London 1867, 492–558, pls XXVII–XXVIII.Google Scholar
Guindon, S., Dufayard, J., Lefort, V., Anisimova, M., Hordijk, W. and Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59, 307–321.Google Scholar
Hechtel, G.J. (1965) A systematic study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History 20, 1–103.Google Scholar
Holzwarth, M., Trendel, J.M., Albrecht, P., Maier, A. and Michaelis, W. (2005) Cyclic peroxides derived from the marine sponge Plakortis simplex. Journal of Natural Products 68, 759–761.Google Scholar
Hoshino, T. (1977) Demosponges from the Kii channel and its environs, western Japan. Proceedings of the Japanese Society of Systematic Zoology 13, 5–15, pls 3–5.Google Scholar
Ivanišević, J., Thomas, O.P., Lejeusne, C., Chevaldonné, P. and Pérez, T. (2010) Metabolic fingerprinting as an indicator of biodiversity: towards understanding inter-specific relationships among Homoscleromorpha sponges. Metabolomics 7, 289–304.Google Scholar
Katoh, K. and Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics 9, 286–298.Google Scholar
Laubenfels, M.W. de (1934) New sponges from the Puerto Rican deep. Smithsonian Miscellaneous Collections 91, 1–28.Google Scholar
Laubenfels, M.W. de (1936a) A discussion of the sponge fauna of the Dry Tortugas in particular and the West Indies in general, with material for a revision of the families and orders of the Porifera. Carnegie Institute of Washington (Tortugas Laboratory Paper N° 467) 30, 1–225.Google Scholar
Laubenfels, M.W. de (1936b) A comparison of the shallow-water sponges near the Pacific end of the Panama Canal with those at the Caribbean end. Proceedings of the United States National Museum 83, 441–466.Google Scholar
Laubenfels, M.W. de (1954) The sponges of the West-Central Pacific. Oregon State Monographs. Studies in Zoology 7, i–x, 1–306, pls I–XII.Google Scholar
Lavrov, D.V., Wang, X. and Kelly, M. (2008) Reconstructing ordinal relationships in the Demospongiae using mitochondrial genomic data. Molecular Phylogenetics and Evolution 49, 111–124.Google Scholar
Lee, H-S., Seo, Y., Rho, J-R., Shin, J. and Paul, V.J. (2001) New steroidal alkaloids from an undescribed sponge of the genus Corticium. Journal of Natural Products 64, 1474–1476.Google Scholar
Lehnert, H. and van Soest, R.W.M. (1998) Shallow water sponges of Jamaica. Beaufortia 48, 71–103.Google Scholar
von Lendenfeld, R. (1887) On the systematic position and classification of sponges. Proceedings of the Zoological Society of London 1886, 558–662.Google Scholar
von Lendenfeld, R. (1907) Die Tetraxonia. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf der Dampfer Valdivia 1898–1899 11, i–iv, 59–374, pls IX–XLVI.Google Scholar
Lévi, C. (1953) Description de Plakortis nigra nov. sp. et remarques sur les Plakinidae (Démosponges). Bulletin du Muséum National d'Histoire Naturelle 25, 320–328.Google Scholar
Lévi, C. (1958) Résultats scientifiques des Campagnes de la ‘Calypso’. Campagne 1951–1952 en Mer Rouge (suite). 11. Spongiaires de Mer Rouge recueillis par la ‘Calypso’ (1951–1952). Annales de l'Institut Océanographique 34, 3–46.Google Scholar
Lévi, C. and Lévi, P. (1983) Démosponges bathyales récoltées par le N/O‘Vauban’ au sud de la Nouvelle-Calédonie. Bulletin du Muséum National d'Histoire Naturelle 5, 931–997.Google Scholar
Moraes, F.C. and Muricy, G. (2003) Taxonomy of Plakortis and Plakinastrella (Demospongiae: Plakinidae) from oceanic islands off north-eastern Brazil, with description of three new species. Journal of the Marine Biological Association of the United Kingdom 83, 1–13.Google Scholar
Mothes, B. and Bastian, M.C.K. (1993) Esponjas do Arquipèlago de Fernando de Noronha, Brasil (Porifera, Demospongiae). Iheringia (Zoologia) 75, 15–31.Google Scholar
Muricy, G. (2011) Diversity of Indo-Australian Plakortis (Demospongiae: Plakinidae), with description of four new species. Journal of the Marine Biological Association of the United Kingdom 91, 303–319.Google Scholar
Muricy, G., Bézac, C., Gallissian, M.-F. and Boury-Esnault, N. (1999) Anatomy, cytology and symbiotic bacteria of four Mediterranean species of Plakina Schulze, 1880 (Demospongiae, Homosclerophorida). Journal of Natural History 33, 156–176.Google Scholar
Muricy, G., Boury-Esnault, N., Bézac, C. and Vacelet, J. (1996) Cytological evidence for cryptic speciation in Mediterranean Oscarella species (Porifera, Homoscleromorpha). Canadian Journal of Zoology 74, 881–896.Google Scholar
Muricy, G., Boury-Esnault, N., Bézac, C. and Vacelet, J. (1998) Taxonomic revision of the Mediterranean Plakina Schulze (Porifera, Demospongiae, Homoscleromorpha). Zoological Journal of the Linnean Society 124, 169–203.Google Scholar
Muricy, G. and Diaz, M.C. (2002) Order Homosclerophorida Dendy, 1905, Family Plakinidae Schluze, 1880. In Hooper, J.N.A. and van Soest, R.W.M. (eds) Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic/Plenum, pp. 73–81.Google Scholar
Muricy, G. and Hajdu, E. (2006) Porifera Brasilis. Guia de identificação das esponjas marinhas mais comuns do Sudeste do Brasil. Rio de Janeiro: Museu Nacional (Série Livros 17).Google Scholar
Muricy, G. and Moraes, F.C. (1998) Marine sponges of Pernambuco State, NE Brazil. Revista Brasileira de Oceanografia 46, 213–217.Google Scholar
Muricy, G. and Pearse, J.S. (2004) A new species of Oscarella (Demospongiae: Plakinidae) from California. Proceedings of the California Academy of Sciences, Series 4 55, 598–612.Google Scholar
Pérez, T., Ivanišević, J., Dubois, M., Pedel, L., Thomas, O.P., Tokina, D. and Ereskovsky, A.V. (2011) Oscarella balibaloi, a new sponge species (Homoscleromorpha: Plakinidae) from the Western Mediterranean Sea: cytological description, reproductive cycle and ecology. Marine Ecology 32, 174–187.Google Scholar
Pulitzer-Finali, G. (1986) A collection of West Indian Demospongiae (Porifera). In appendix, a list of the Demospongiae hitherto recorded from the West Indies. Annali del Museo civico di storia naturale Giacomo Doria 86, 65–216.Google Scholar
Pulitzer-Finali, G. (1993) A collection of marine sponges from East Africa. Annali del Museo civico di storia naturale Giacomo Doria 89, 247–350.Google Scholar
Pulitzer-Finali, G. (1996) Sponges from the Bismarck Sea. Bollettino dei Musei e degli Istituti Biologici della (R.) Università di Genova 60–61, 101–138.Google Scholar
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, 208–212.Google Scholar
Ridley, C.P. and Faulkner, D.J. (2003) New cytotoxic steroidal alkaloids from the Philippine sponge Corticium niger. Journal of Natural Products 66, 1536–1539.Google Scholar
Rosser, R.M. and Faulkner, D.J. (1984) Two steroidal alkaloids from a marine sponge, Plakina sp. Journal of Organic Chemistry 49, 5157–5160.Google Scholar
Rudi, A., Afanii, R., Gravalos, G.L., Aknin, M., Gaydou, E., Vacelet, J. and Kashman, Y. (2003) Three new cyclic peroxides from the marine sponge Plakortis aff. simplex. Journal of Natural Products 66, 682–685.Google Scholar
Rützler, K., Diaz, M. C., van Soest, R. W. M., Zea, S., Smith, K. P., Alvarez, B. and Wulff, J. (2000) Diversity of sponge fauna in mangrove ponds, Pelican Cays, Belize. Atoll Research Bulletin 476, 229–248.Google Scholar
Saghai-Maroof, M.A., Soliman, K.M., Jorgensen, R.A. and 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.Google Scholar
Schmidt, O. (1862) Die Spongien des adriatischen Meeres. Leipzig: Wilhelm Engelmann, i–viii, 1–88, pls 1–7.Google Scholar
Schmidt, O. (1868) Die Spongien der Küste von Algier. Mit Nachträgen zu den Spongien des Adriatischen Meeres (Drittes Supplement). Leipzig: Wilhelm Engelmann, i–iv, 1–44, pls I–V.Google Scholar
Schmidt, O. (1880) Die Spongien des Meerbusen von Mexico (Und des caraibischen Meeres). Abtheilung II. Hexactinelliden. Heft II. Reports on the dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico, by the USCSS ‘Blake'. Jena: Gustav Fischer, pp. 33–90.Google Scholar
Schulze, F.E. (1880) Untersuchungen über den Bau und die Entwicklung der Spongien. Neunte Mittheilung. Die Plakiniden. Zeitschrift für Wissenschaftliche Zoologie 34, 407–451.Google Scholar
SoestR.W.M., van R.W.M., van, Boury-Esnault, N., Hooper, J.N.A., 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. and Díaz, M.C. (2013) World Porifera database. Available online at: http://www.marinespecies.org/porifera (accessed 25 February 2013).Google Scholar
Spurr, A.R. (1969) A low-viscosity epoxy resin embedding medium for electron microscopy. Journal of Ultrastructure Research 26, 31–43.Google Scholar
Thomas, P.A. (1968) Studies on Indian sponges–IV. Additions to the genus Corticum Schmidt with notes on the distribution of C. candelabrum Schmidt. Journal of Marine Biological Association of India 10, 260–263.Google Scholar
Topsent, E. (1890) Sur la distribution géographique de quelques Microsclerophora. Bulletin de la Société Zoologique de France 15, 231–233.Google Scholar
Topsent, E. (1895) Étude monographique des spongiaires de France. II. Carnosa. Archives de Zoologie Expérimentale et Générale 3, 493–590.Google Scholar
Topsent, E. (1897) Spongiaires de la Baie d'Amboine. (Voyage de MM. M. Bedot et C. Pictet dans l'Archipel Malais). Revue Suisse de Zoologie 4, 421–487, pls 18–21.Google Scholar
Topsent, E. (1923) Spongiaires du Musée Zoologique de Strasbourg. Choristides. Bulletin de l'Institut Océanographique, Monaco 435, 1–16.Google Scholar
Topsent, E. (1927) Diagnoses d'éponges nouvelles recueillies par le Prince Albert ler de Monaco. Bulletin de l'Institut Océanographique, Monaco 502, 1–19.Google Scholar
Topsent, E. (1937) Notes diverses sur les éponges. Bulletin de l'Institut Océanographique de Monaco 722, 1–15.Google Scholar
Tuzet, O. and Paris, J. (1963) Octavella galangaui n.g. n.sp. Demospongiae, Oscarellidae de Banyuls-sur-Mer. Vie et Milieu 14, 71–89.Google Scholar
Vishnyakov, A.E. and Ereskovsky, A.V. (2009) Bacterial symbionts as an additional cytological marker for identification of sponges without a skeleton. Marine Biology 156, 1625–1632.Google Scholar
Vosmaer, G.C.J. (1881) Ueber Leucandra aspera, H. Nebst allgemeinen Bemerkungen ueber das Canalsystem der Spongien. Tijdschrift der Nederlandsche Dierkundige Vereeniging 5, 144–164, pls III–IV.Google Scholar
Vosmaer, G.C.J. (1887) Klassen und Ordnungen der Spongien (Porifera). Die Klassen und Ordnungen des Thierreichs. Leipzig: Winter'sche Verlaghandlung.Google Scholar
Wiedenmayer, F. (1977) Shallow-water sponges of the western Bahamas. Experientia Supplementum 28, 1–287, pls 281–243.Google Scholar
Wilson, H.V. (1902) The sponges collected in Porto Rico in 1889 by the U.S. Fish Commission Steamer Fish Hawk. Bulletin of the United States Fish Commission 2, 375–411.Google Scholar
Zampella, A., D'Orsi, R., Sepe, V., De Marino, S., Borbone, N., Valentin, A., Debitus, C., Zollo, F. and D'Auria, M.V. (2005) Isolation of plakinamine I: a new steroidal alkaloid from the marine sponge Corticium sp. and synthesis of an analogue model compound. European Journal of Organic Chemistry 2005, 4359–4363.Google Scholar
Zea, S. (1987) Esponjas del Caribe Colombiano. Bogota: Editorial Catálogo Científico.Google Scholar
Zea, S., Henkel, T.P. and Pawlik, J.R. (2009) The sponge guide: a picture guide to Caribbean sponges. Available at: http://www.spongeguide.org (accessed 25 February 2013).Google Scholar