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Molecular and morphological evidence that Phymactis papillosa from Argentina is, in fact, a new species of the genus Bunodosoma (Cnidaria: Actiniidae)

Published online by Cambridge University Press:  14 December 2011

Paula Braga Gomes ,
Renata Schama  and
Antônio Mateo Solé-Cava
Paula Braga Gomes
Affiliation:
Grupo de Pesquisa em Antozoários (GPA), Departamento de Biologia, Universidade Federal Rural de Pernambuco, R. Don Manoel de Medeiros, s/n, Dois Irmãos, Recife, PE, 52.171-900, Brazil
Renata Schama
Affiliation:
Laboratório de Biodiversidade Molecular, Instituto de Biologia, CCS, Bloco A, Universidade Federal do Rio de Janeiro, Cidade Universitária, Rio de Janeiro, RJ, 21941-590, Brazil
Antônio Mateo Solé-Cava*
Affiliation:
Laboratório de Biodiversidade Molecular, Instituto de Biologia, CCS, Bloco A, Universidade Federal do Rio de Janeiro, Cidade Universitária, Rio de Janeiro, RJ, 21941-590, Brazil
*
Correspondence should be addressed to: A.M. Solé-Cava, Laboratório de Biodiversidade Molecular, Instituto de Biologia, CCS, Bloco A, Universidade Federal do Rio de Janeiro, Cidade Universitária, Rio de Janeiro, RJ, 21941-590, Brazil email: [email protected]
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Abstract

Phymactis papillosa is a rocky shore sea anemone that is commonly found in the Pacific Ocean, from the Gulf of California to Tierra del Fuego, and in the Mar del Plata region, Argentina. The genus Phymactis is closely related to Bunodosoma and, due to character plasticity, a number of misidentifications have occurred. Therefore, the presence of P. papillosa in Argentina has been doubted but the matter had not been investigated in detail. Here we analyse P. papillosa specimens from Argentina and compare them, using molecular and morphological markers, to specimens from the species' type locality. In a phylogenetic analysis using 19 allozyme markers and ribosomal internal transcribed spacers sequences of different sea anemone genera, including all West Atlantic Bunodosoma species, we have found that the specimens from Argentina were genetically divergent from P. papillosa from Chile and closely related to West Atlantic Bunodosoma species. The genetic and morphological analyses indicate that those specimens belong to a new species of the genus Bunodosoma, described here as B. zamponii sp. nov.

Keywords

allozymesribosomalinternal transcribed spacerphylogenyspecific statusActiniarianew species
Type
Research Article
Information
Journal of the Marine Biological Association of the United Kingdom , Volume 92 , Issue 5 , August 2012 , pp. 895 - 910
Copyright
Copyright © Marine Biological Association of the United Kingdom 2011

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References

REFERENCES

Acuña, F.H. (1997) Ecología trófica de actiniarios (Cnidaria, Anthozoa) intermareales: selección de la talla de las presas. Physis—A 53, 1–5.Google Scholar
Acuña, F.H. and Zamponi, M.O. (1995) Ecology of intertidal sea anemones. Density, dispersion and autoecology of Phymactis clematis Dana,1849 (Anthozoa: Actiniaria). Ciencias Marinas 21, 1–12.CrossRefGoogle Scholar
Acuña, F.H. and Zamponi, M.O. (1996) Ecología trófica de las anémonas intermareales Phymactis clematis Dana, 1849, Aulactinia marplatensis (Zamponi, 1977) y A. reynaudi (Milne-Edwards, 1857) (Actiniaria: Actiniidae): relaciones entre las anémonas y sus presas. Ciencia Marina 22, 397–413.CrossRefGoogle Scholar
Acuña, F.H. and Zamponi, M.O. (1997) The use of cnidocysts for ecological races identification from sea anemones populations (Anthozoa, Actiniidae). Iheringia—Série Zoologia 82, 9–18.Google Scholar
Acuña, F.H., Zamponi, M.O. and Perez, C.D. (1996) Metodología para la cuantificación de algunas estructuras taxonómicas en Actiniaria (Cnidaria, Anthozoa). Iheringia—Série Zoologia 80, 27–31.Google Scholar
Acuña, F.H., Excoffon, A.C., McKinstry, S.R. and Martinez, D.E. (2007) Characterization of Aulactinia (Actiniaria: Actiniidae) species from Mar del Plata (Argentina) using morphological and molecular data. Hydrobiologia 592, 249–256.CrossRefGoogle Scholar
Ayre, D.J., Minchinton, T.E. and Perrin, C. (2009) Does life history predict past and current connectivity for rocky intertidal invertebrates across a marine biogeographic barrier? Molecular Ecology 18, 1887–1903.CrossRefGoogle ScholarPubMed
Belém, M.J.C. (1987) Aspectos da biologia de Bunodosoma caissarum Corrêa, 1964 (Cnidaria, Anthozoa, Actiniidae) do litoral do Rio de Janeiro, com ênfase na estimativa de seu comportamento reprodutivo. PhD thesis. Universidade de São Paulo, São Paulo, Brazil.Google Scholar
Belém, M.J.C. (1988) Anatomy and biology of Bunodosoma caissarum Corrêa, 1964 (Cnidaria, Anthozoa, Actiniidae). 1. Systematic position and revision of morphology and microanatomy. Anais da Academia Brasileira de Ciencias 60, 365–375.Google Scholar
Belém, M.J.C. and Monteiro, D.C. (1981) Fauna de cnidários do Rio de Janeiro. III. Anthopleura varioarmata Watzl, 1922 (Actiniaria, Endomyaria), uma nova ocorrência de Actiniidae. Seminários de Biologia Marinha–Academia Brasileira de Ciências, pp. 193–203.Google Scholar
Belkhir, K., Borsa, P., Chikhi, L., Raufaste, N. and Bonhomme, F. (2002) GENETIX 4.04, logiciel sous Windows TM pour la génétique des populations. Laboratoire Génome, Populations, Interactions, CNRS UMR 5000, Université de Montpellier II, Montpellier, France.Google Scholar
Billingham, M. and Ayre, D.J. (1996) Genetic subdivision in the subtidal, clonal sea anemone Anthothoe albocincta. Marine Biology 125, 153–163.CrossRefGoogle Scholar
Brattstrom, H. and Johanssen, A. (1983) Ecological and regional zoogeography of marine benthic fauna of Chile. Report No. 49 of the Lund University Chile Expedition, 1948–59. Sarsia 68, 289–339.CrossRefGoogle Scholar
Bucklin, A. and Hedgecock, D. (1982) Biochemical genetic evidence for a third species of Metridium (Coelenterata: Actiniaria). Marine Biology 66, 1–7.CrossRefGoogle Scholar
Carlgren, O. (1899) Zoantharien. Hamburger Magalhaensische Sammelreise 4, 1–48.Google Scholar
Carlgren, O. (1920) Förteckning över de i skånska resan omnämnda djurarterna. In Linnés Skånska Resa med anmärkningar utgiven av Jöran Sahlgren. Stockholm: P.A. Norstedt and Söners, pp. 19–25.Google Scholar
Carlgren, O. (1922) Actiniaria und Zoantharia von Juan Fernandez und der Osterinsel. In Skottsberg, C. (ed.) The natural history of Juan Fernandez and Easter Island. 3 Part 2. Uppsala: Almquist & Wiksells Boktryckeri, pp. 145–160.Google Scholar
Carlgren, O. (1924) On Boloceroides, Bunodeopsis and their supposed allied genera. Arkiv förZoologi 17A, 1–20.Google Scholar
Carlgren, O. (1934) Zur revision der Actiniarien. Arkiv för Zoologi 26A, 1–36.Google Scholar
Carlgren, O. (1939) Actiniaria and Zoantharia of the Scottish National Antarctic Expedition, 1902–1904. Transactions of the Royal Society of Edinburgh 49, 791–800.CrossRefGoogle Scholar
Carlgren, O. (1945) Further contributions to the knowledge of the cnidom in the Anthozoa especially in the Actiniaria. Kungliga Fysiografiska Sällskapets Handlingar 56 NF, 1–24.Google Scholar
Carlgren, O. (1949) A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. Kungliga Svenska Vetenskapsakademiens Handlingar 4, 1–121.Google Scholar
Carlgren, O. (1951) The actinian fauna of the Gulf of California. Proceedings of the United States National Museum 101, 415–449.CrossRefGoogle Scholar
Carlgren, O. (1959) Corallimorpharia and Actiniaria with description of new genus and species from Peru. Reports of the Lund University Chile Expedition 1948–49. Lund Universitets Arsskrift 38, 1–39.Google Scholar
Carter, M.A. and Thorpe, J.P. (1981) Reproductive, genetic and ecological evidence that Actinia equina var. mesembrianthemum and var. fragacea are not conspecific. Journal of the Marine Biological Association of the United Kingdom 61, 79–93.CrossRefGoogle Scholar
Carter-Verdeilhan, C.D. (1965) Actinias de Montemar, Valparaiso. Revista de Biologia Marina 12, 129–157.Google Scholar
Corrêa, D.D. (1964) Corallimorpharia e Actiniaria do Atlântico Oeste Tropical. PhD thesis. Universidade de São Paulo, Brazil.Google Scholar
Currie, D.R. and Small, K.J. (2006) The influence of dry-season conditions on the bottom dwelling fauna of an east Australian sub-tropical estuary. Hydrobiologia 560, 345–361.CrossRefGoogle Scholar
Daly, M. (2003) On the anatomy, terminology and homology of acrorhagi and pseudoacrorhagi. Zoologische Mededelingen 345, 89–102.Google Scholar
Daly, M. (2004) Anatomy and taxonomy of three species of sea anemones (Cnidaria: Anthozoa: Actiniidae) from the Gulf of California, including Isoaulactinia hespervolita Daly, n. sp. Pacific Science 58, 377–390.CrossRefGoogle Scholar
Daly, M., Chaudhuri, A., Gusmão, L. and Rodríguez, E. (2008) Phylogenetic relationships among sea anemones (Cnidaria: Anthozoa: Actiniaria). Molecular Phylogenetics and Evolution 48, 292–301.CrossRefGoogle ScholarPubMed
Damato, M.E. and Corach, D. (1996) Genetic diversity of populations of the fresh-water shrimp Macrobrachium borellii (Caridea, Palaemonidae) evaluated by RAPD analysis. Journal of Crustacean Biology 16, 650–655.CrossRefGoogle Scholar
Douek, J., Barki, Y., Gateño, D. and Rinkevich, B. (2002) Possible cryptic speciation within the sea anemone Actinia equina complex detected by AFLP markers. Zoological Journal of the Linnaean Society 136, 315–320.CrossRefGoogle Scholar
Excoffon, A.C. and Zamponi, M.O. (1991) La biologia reproductiva de Phymactis papillosa Dana, 1849 (Actiniaria: Actiniidae); gametogénesis, períodos reproductivos, desarrollo embrionario y larval. Spheniscus 9, 25–39.Google Scholar
Excoffon, A.C., Genzano, G.N. and Zamponi, M.O. (1999) Macrobenthos associated with a population of Anthothoe chilensis (Lesson, 1830) (Cnidaria, Actiniaria) in Mar del Plata Harbor, Argentina. Ciencias Marinas 25, 177–191.CrossRefGoogle Scholar
Farris, J.S., Kallersjo, M., Kluge, A.G. and Bult, C. (1995) Constructing a significance test for incongruence. Systematic Biology 44, 570–572.CrossRefGoogle Scholar
Fautin, D.G. (2011) Hexacorallins of the World. Available at: http://geoporbal.kgs.ku.edu/hexacoral/anenone2/index.cfm (accessed 28 October 2011).Google Scholar
Fautin, D.G., Hickman, C.P. Jr, Daly, M. and Molodtsova, T. (2007) Shallow-water sea anemones (Cnidaria: Anthozoa: Actiniaria) and tube anemones (Cnidaria: Anthozoa: Ceriantharia) of the Galápagos Islands. Pacific Science 61, 548–573.CrossRefGoogle Scholar
Felsenstein, J. (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular Evolution 17, 368–376.CrossRefGoogle ScholarPubMed
Forsman, Z.H., Barshis, D.J., Hunter, C.L. and Toonen, R.J. (2009) Shape-shifting corals: molecular markers show morphology is evolutionarily plastic in Porites. BMC Evolutionary Biology 9, 45.CrossRefGoogle ScholarPubMed
Forsman, Z.H., Guzman, H.M., Chen, C.A., Fox, G.E. and Wellington, G.M. (2005) An ITS region phylogeny of Siderastrea (Cnidaria: Anthozoa): is S. glynni endangered or introduced? Coral Reefs 24, 343–347.CrossRefGoogle Scholar
Försterra, G. and Haussermänn, V. (2003) First report on large scleractinian (Cnidaria: Anthozoa) accumulations in cold-temperate shallow water of South Chilean fjords. Zoologische Verhandelingen, Leiden 345, 117–128.Google Scholar
Fukami, H., Chen, C.A., Budd, A.F., Collins, A., Wallace, C., Chuang, Y., Chen, C., Dai, C., Iwao, K., Sheppard, C. and Knowlton, N. (2008) Mitochondrial and nuclear genes suggest that stony corals are monophyletic but most families of stony corals are not (Order Scleractinia, Class Anthozoa, Phylum Cnidaria). PLoS One 3, 1–9.CrossRefGoogle Scholar
Garese, A., Guzmán, H.M. and Acuña, F.H. (2009) Sea anemones (Cnidaria: Actiniaria and Corallimorpharia) from Panama. Revista de Biologia Marina y Oceanografia 44, 791–802.Google Scholar
Genzano, G.N., Acuña, F.H., Excoffon, A.C. and Perez, C.D. (1996) Cnidarios bentónicos de la provincia de Buenos Aires. Listado sistemático, distribución y estrategias de colonización. Actas Jornadas Pampeanas Ciencias Naturales 6, 113–121.Google Scholar
Gomes, P.B. (2002) Anêmonas-do-mar (Cnidaria, Actiniaria) de Pernambuco (Brasil). In Tabarelli, M. and Silva, J.M.C. (eds) Diagnóstico da biodiversidade de Pernambuco. Volume II. Recife, Brazil: SECTMA e Editora Massangana, pp. 343–364.Google Scholar
Gomes, P.B., Belém, M.J. and Schlenz, E. (1998) Distribution, abundance and adaptations of three species of Actiniidae (Cnidaria, Actiniaria) on an intertidal beach rock in Carneiros beach, Pernambuco, Brasil. Miscellanea Zoologica 21, 65–72.Google Scholar
Grohmann, P.A. (1998) Anêmonas-do-mar (Cnidaria, Anthozoa, Actiniaria) de Vitória, ES. In XII Congresso Brasileiro de Zoologia, Universidade Federal de Pernambuco, Brazil, p. 9.Google Scholar
Gusmão, L.C. (2010) Systematics and evolution of sea anemones (Cnidaria: Actiniaria: Hormathiidae) symbiotic with hermit crabs. PhD thesis. Ohio State University, United States of America.CrossRefGoogle Scholar
Haldane, J.B.S. (1954) An exact test for randomness of mating. Genetics 52, 631–635.Google Scholar
Hare, M.P., Karl, S.A. and Avise, J.C. (1996) Anonymous nuclear DNA markers in the American oyster and their implications for the heterozygote deficiency phenomenon in marine bivalves. Molecular Biology and Evolution 13, 334–345.CrossRefGoogle ScholarPubMed
den Hartog, J.C. (1987) A redescription of the sea anemone Bunodosoma biscayensis (Fischer, 1984) (Actiniaria, Actiniidae). Zoologische Mededelingen 61, 533–558.Google Scholar
Haussermänn, V. (2004) Redescription of Phymactis papillosa (Lesson, 1830) and Phymantea pluvia (Brayton in Dana, 1846) (Cnidaria: Anthozoa), two actiniid sea anemones from the Southeast Pacific with a discussion of the genera Phymacti, Bunodosoma and Phymantea. Zoologische Mededelingen, Leiden 78, 345–381.Google Scholar
Hellberg, M.F. (2009) Gene flow and isolation among populations of marine animals. Annual Review of Ecology, Evolution and Systematics 40, 291–310.CrossRefGoogle Scholar
Hillis, D.M., Moritz, C., Porter, C.A. and Baker, R.J. (1991) Evidence for biased gene conversion in concerted gene evolution of ribosomal DNA. Science 251, 308–310.CrossRefGoogle ScholarPubMed
Jukes, T.H. and Cantor, C.R. (1969) Evolution of protein molecules. In Munro, H.N. (ed.) Mammalian protein metabolism. New York: Academic Press, New York, pp. 21–123.CrossRefGoogle Scholar
Kiernan, J.A. (1990) Histological and histochemical methods: theory and practice. Oxford: Pergamon.Google Scholar
Knowlton, N. (2000) Molecular genetic analyses of species boundaries in the sea. Hydrobiologia 420, 73–90.CrossRefGoogle Scholar
Kumar, S., Tamura, K. and Nei, M. (2004) Mega 3: integrated software for molecular evolutionary genetics analysis and sequence alignment. Briefings in Bioinformatics 5, 150–163.CrossRefGoogle Scholar
Lancellotti, D.A. and Vásquez, J.A. (1999) Biogeographical patterns of benthic macroinvertebrates in the Sutheastern Pacific littoral. Journal of Biogeography 26, 1001–1006.CrossRefGoogle Scholar
Lessios, H.A. (1992) Testing electrophoretic data for agreement with Hardy–Weinberg expectations. Marine Biology 112, 517–523.CrossRefGoogle Scholar
Lima, D., Freitas, J.E.P., Araujo, M.E. and Solé-Cava, A.M. (2005) Genetic detection of cryptic species in the frill goby Bathygobius soporator. Journal of Experimental Marine Biology and Ecology 320, 211–223.CrossRefGoogle Scholar
Lôbo-Hajdu, G., Guimarães, A.C., Mendes, A.M., Lamarão, F.R.M., Vieiralves, T., Mansure, J.J. and Albano, R.M. (2004) Intragenic, intra- and interspecific variation in the rDNA ITS of Porifera revealed by PCR-single-strand comformation polymorphism (PCR-SSCP). Bolletino dei Musei e degli Instituti Biologici dell'Università di Genova 68, 413–423.Google Scholar
Manchenko, G.P. (1994) Handbook of detection of enzymes on electrophoretic gels. Ann Arbor, MI: CRC Press Inc.Google Scholar
Manchenko, G.P., Dautova, T.N. and Latypov, Y.Y. (2000) High level of genetic divergence between sympatric color morphs of the littoral sea anemone Anthopleura orientalis (Anthozoa: Actiniaria). Biochemical Systematics and Ecology 28, 737–750.CrossRefGoogle ScholarPubMed
McCommas, S.A. (1982) Biochemical genetics of the sea anemone Bunodosoma cavernata and the zoogeography of the Gulf of Mexico. Marine Biology 68, 169–173.CrossRefGoogle Scholar
McCommas, S.A. and Lester, L.J. (1980) Electrophoretic evaluation of the taxonomic status of two species of sea anemone. Biochemical Systematics and Ecology 8, 289–292.CrossRefGoogle Scholar
McFadden, C.S., Grosberg, R.K., Cameron, B.B., Karlton, D.P. and Secord, D. (1997) Genetic relationships within and between clonal and solitary forms of the sea anemone Anthopleura elegantissima revisited: evidence for the existence of two species. Marine Biology 128, 127–139.CrossRefGoogle Scholar
McManus, M.G., Place, A.R. and Zamer, W.E. (1997) Physiological variation among clonal genotypes in the sea anemone Haliplanella lineata: growth and biochemical content. Biological Bulletin. Marine Biological Laboratory, Woods Hole 192, 426–443.CrossRefGoogle ScholarPubMed
McMurrich, J.P. (1904) The Actiniae of the Plate Collection. Zoologische Jahrbuecher 6 Supplement, 215–306.Google Scholar
Miller, K.J. and Ayre, D.J. (2008) Population structure is not a simple function of reproductive mode and larval type: insights from tropical corals. Journal of Animal Ecology 77, 713–724.CrossRefGoogle Scholar
Miller, M.P. (1997) Tools for population genetic analysis. Version 1.3. Department of Biological Sciences, Northern Arizona University, Flagstaff.Google Scholar
Monteiro, F.A., Solé-Cava, A.M. and Thorpe, J.P. (1997) Extensive genetic divergence between populations of the common intertidal sea anemone Actinia equina from Britain, the Mediterranean and the Cape Verde Islands. Marine Biology 129, 425–433.CrossRefGoogle Scholar
Nei, M. (1978) Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89, 583–590.CrossRefGoogle ScholarPubMed
Olivera, E.G., Patronelli, D.L. and Zamponi, M.O. (2009) Morphological and functional study of the marginal sphincter of the sea anemones Phymactis clematis and Aulactinia marplatensis from intertidal of Mar del Plata, Argentina. Iheringia 99, 313–316.CrossRefGoogle Scholar
Paranhos, J.D.N., Pinto, S.L., Silva, C.B., Amaral, L.F.B. and Farias, R.R.S. (1999) Levantamento preliminar dos Cnidaria da praia de Barra Grande—PI. In XII Encontro de Zoologia do Nordeste, Feira de Santana, Brazil, p. 184.Google Scholar
Patronelli, D., Olivera, D. and Zamponi, M.O. (2005) Influence of intertidal environment on muscle activity in different species of sea anemones (Actiniaria). Animal Biology 55, 101–109.CrossRefGoogle Scholar
Patronelli, D., Olivera, E.G., Zamponi, M.O. and Crupkin, M. (2008) Caracterización morfológica, funcional y bioquímica del esfínter marginal de la anémona de mar Phymactis clematis (Cnidaria). Investigaciones Marinas 29, 73–77.Google Scholar
Patronelli, D., Zamponi, M.O., Bustos, A. and Vega, F. (1987) Morphological adaptations in the marginal sphincter of anemone Phymactis clematis Dana, 1849 from different environment. Biochemical Physiology 88a, 337–340.CrossRefGoogle Scholar
Pax, F. (1924) Actiniarien, Zoantharien und Ceriantharien von Curaçao. Kungliga Zoologisch Genootschap Natura Artis Magistra 23, 93–122.Google Scholar
Pollero, R.J. (1983) Lipid and fatty acid characterization and metabolism in the sea anemone Phymactis clematis (Dana). Lipids 18, 12–17.CrossRefGoogle Scholar
Posada, D. and Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics 14, 817–818.CrossRefGoogle ScholarPubMed
Raymond, M. and Rousset, F. (1995) An exact test for population differentiation. Evolution 49, 1280–1283.CrossRefGoogle ScholarPubMed
Rivadeneira, M.M. and Oliva, E. (2001) Patrones asociados a la conducta de desplaziamiento local en Phymactis papillosa Drayton (Anthozoa: Actiniidae). Revista Chilena de Historia Natural 74, 855–863.CrossRefGoogle Scholar
Rocha, L.A., Bass, A.L, Robertson, D.R. and Bowen, B.W. (2002) Adult habitat preferences, larval dispersal, and the comparative phylogeography of three Atlantic surgeonfishes (Teleostei: Acanthuridae) Molecular Ecology 11, 243–251.CrossRefGoogle ScholarPubMed
Russo, C.A.M. and Solé-Cava, A.M. (1991) Genetic variation and differentiation between populations of a tropical sea-anemone (Bunodosoma caissarum Corrêa). Revista de Biologia Tropical 39, 41–46.Google Scholar
Russo, C.A.M., Solé-Cava, A.M. and Thorpe, J.P. (1994) Population structure and genetic variation in two tropical sea anemones (Cnidaria, Actinidae) with different reproductive strategies. Marine Biology 119, 267–276.CrossRefGoogle Scholar
Saitou, N. and Nei, M. (1987) The neighbor joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4, 406–425.Google Scholar
Schama, R., Solé-Cava, A.M. and Thorpe, J.P. (2005) Genetic divergence between East and West Atlantic populations of Actinia sea anemones (Cnidaria: Actiniidae). Marine Biology 146, 435–443.CrossRefGoogle Scholar
Schmidt, H. (1969) Die nesselkapseln der Aktinien und ihre differentialdiagnostische Bedeutung. Helgoländer Wissenschaftliche Meeresuntersuchungen 19, 284–317.CrossRefGoogle Scholar
Schmidt, H. (1972) Prodromus zu einer Monographie der mediterranen Aktinien. Zoologica 42, 1–120.Google Scholar
Schmidt, H. (1974) On evolution in the Anthozoa. Proceedings of the Second International Coral Reef Symposium 1, 533–560.Google Scholar
Sebens, K. and Paine, R. (1978) Biogeography of anthozoans along the West coast of South America: habitat, disturbance, and prey availability. Proceedings of the International Symposium of Marine Biogeography and Evolution of South Hemisphere 1, 219–237.Google Scholar
Sneath, P.H.A. and Sokal, R.R. (1973) Numerical taxonomy. San Francisco, CA: W.H. Freeman Press.Google Scholar
Solé-Cava, A.M. and Thorpe, J.P. (1989) Biochemical correlates of genetic variation in marine lower invertebrates. Biochemical Genetics 27, 303–312.CrossRefGoogle ScholarPubMed
Solé-Cava, A.M., Thorpe, J.P. and Kaye, J.G. (1985) Reproductive isolation with little genetic divergence between Urticina (= Tealia) felina and U. eques (Anthozoa: Actiniaria). Marine Biology 85, 279–284.CrossRefGoogle Scholar
Stephenson, T.A. (1935) The British sea anemones. Volume 2. London: The Ray Society.Google Scholar
Stoletzki, N. and Schierwater, B. (2005) Genetic and color morphs differentiations in the Caribbean sea anemone Condylactis gigantea. Marine Biology 147, 747–754.CrossRefGoogle Scholar
Stotz, W.B. (1979) Functional morphology and zonation of three species of sea anemones from rocky shores in Southern Chile. Marine Biology 50, 181–188.CrossRefGoogle Scholar
Swofford, D.L. (2000) PAUP*: phylogenetic analysis using parsimony (*and other methods). Sunderland, MA: Sinauer Associates.Google Scholar
Swofford, D.L. and Selander, R.B. (1981) BIOSYS-1, a FORTRAN programme for the comprehensive analysis of electrophoretic data in population genetics and systematics. Journal of Heredity 72, 281–283.CrossRefGoogle Scholar
Thompson, J.D., Higgins, D.G. and Gibson, T.J. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22, 4673–4680.CrossRefGoogle ScholarPubMed
Thorpe, J.P. and Solé-Cava, A.M. (1994) The use of allozyme electrophoresis in invertebrate systematics. Zoologica Scripta 23, 3–18.CrossRefGoogle Scholar
Valentin, J.L. (2000) Ecologia numérica: uma introdução à análise multivariada de dados ecológicos. Rio de Janeiro: Interciência LTDA.Google Scholar
Vianna, P., Schama, R. and Russo, C.A.M. (2003) Genetic divergence and isolation by distance in the West Atlantic sea anemone Actinia bermudensis (McMurrich, 1889). Journal of Experimental Marine Biology and Ecology 297, 19–30.CrossRefGoogle Scholar
Ward, R.D. and Beardmore, J.A. (1977) Protein variation in the plaice (Pleuronectes platessa). Genetic Research 30, 45–62.CrossRefGoogle ScholarPubMed
Watzl, O. (1922) Die Actiniarien der Bahamainseln. Arkiv för Zoologi 14, 1–89.Google Scholar
Zamponi, M.O. (1977) La anemofauna de Mar del Plata y localidades vecinas. I. Las anemonas Boloceroidaria y Endomyaria (Coelentherata: Actiniaria). Acta Neotrópica 23, 137–153.Google Scholar
Zamponi, M.O. (1989) Los Cnidaria y su interaccion pelagico-bentonica. PhD thesis. Universidad Nacional de Mar del Plata, Mar del Plata, Argentina.Google Scholar
Zamponi, M.O. (1993) El ambiente intermareal subtemplado frio como un posible «pool» de tipos reproductivos. Physis—A 51, 13–15.Google Scholar
Zamponi, M.O. (2000) El estuario del Río de La Plata: una barrera geográfica para los cnidarios bentónicos marinos? Biociências 8, 127–136.Google Scholar
Zamponi, M.O. (2005) Estudio de la reproducción sexual de las anémonas de mar (Actiniaria) y la estrategia del hombre pobre (the poor man's game). Revista Real Academia Galega de Ciencias 24, 5–28.Google Scholar
Zamponi, M.O. and Perez, C.D. (1996) Comparative morphological study of different species of Actiniaria between the intertidal zone from Mar del Plata and Santa Clara del Mar (Argentine). I. Phymactis papillosa Dana, 1849 (Anthozoa, Actiniidae). Biociências 4, 91–102.Google Scholar
Zamponi, M.O., Belém, M.J.C., Schlenz, E. and Acuña, F.H. (1998a) Distribution and some ecological aspects of Corallimorpharia and Actiniaria from shallow waters of the South American Atlantic Coast. Physis—A 55, 31–45.Google Scholar
Zamponi, M.O., Genzano, G.N., Acuña, F.H. and Excoffon, A.C. (1998b) Studies of benthic cnidarian populations along a transect off Mar del Plata (Argentina). Russian Journal of Marine Biology 24, 7–13.Google Scholar
Zouros, E. and Foltz, D.W. (1984) Possible explanations of heterozygote deficiency in bivalve molluscs. Malacologia 25, 583–591.Google Scholar