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Morphological and Molecular Evidence for Osmundea Coelenterata Comb

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Morphological and Molecular Evidence for Osmundea Coelenterata Comb Hidrobiológica 2016, 26 (2): 203-211 Morphological and molecular evidence for Osmundea coelenterata comb. nov. (Ceramiales, Rhodophyta) from the tropical Atlantic Ocean Evidencias morfológicas y moleculares para Osmundea coelenterata comb. nov. (Ceramiales, Rhodophyta) del Océano Atlántico tropical Mutue T. Fujii1, Abel Sentíes2, Abdiel Jover C.3, Jhoana Díaz-Larrea2 and Arsenio Areces4 1Núcleo de Pesquisa em Ficologia, Instituto de Botânica, Av. Miguel Estéfano, 3687 - 04301-902 São Paulo. Brasil 2Departamento de Hidrobiología, Universidad Autónoma Metropolitana, Unidad Iztapalapa, Apdo. Postal 55-535, CDMX, 09340. México 3Departamento de Biología, Universidad de Oriente, Patricio Lumumba s/n. CP: 90500 Santiago de Cuba. Cuba 4Instituto de Oceanología, Primera no. 18406, Rpto. Flores, Playa, Habana. Cuba e-mail: [email protected] Fujii M. T., A. Sentíes, C. A. Jover, J. Díaz-Larrea and A. Areces. 2016. Morphological and molecular evidence for Osmundea coelenterata comb. nov. (Ceramiales, Rhodophyta) from the tropical Atlantic Ocean. Hidrobiológica 26 (2): 203-211. ABSTRACT Background. As part of the monographic studies on the Laurencia complex (Rhodomelaceae, Rhodophyta) from Cuba, detailed taxonomic research was carried out on Laurencia coelenterata, a diminutive species collected in Santiago de Cuba, in the eastern Cuban archipelago. The vegetative and reproductive features of the species clearly belong to the genus Osmundea, a member of the Laurencia complex: two pericentral cells per each vegetative axial segment, tetrasporangia cut off randomly from cortical cells, and filament-type spermatangial branches.Goals . In this study the morphological characteristics previously unknown to Laurencia coelenterata are shown, and in order that their taxonomic position within the Laurencia complex is discussed. Furthermore, the global geographic distribution of the Osmundea species is analysed. Methods. Morphological and molecular analysis of the rbcL gene sequences were performed allied with the preparation and interpretation of phylogenetic hypotheses. Results. The new morphological analysis of the paratype specimen (# 4451) housed in the Puertorriqueño Marine Herbarium showed that Laurencia coelenterata was incorrectly placed in the genus Laurencia. The phylogenetic position of the present species was inferred by the analysis of the chloroplast-encoded rbcL gene sequences, and the range of the genetic variation found in this analysis also supports the transfer of Laurencia coelenterata to the genus Osmundea, as O. coelenterata (D. L. Ballantine et Aponte) M.T. Fujii, Sentíes et Areces. Conclu- sions. The morphological and molecular approach in algal taxonomy is allowing consolidate the nomenclatural status of the species, and the new combination for Laurencia coelenterata as Osmundea coelenterata is not an exception. Key words: Laurencia sensu lato, Osmundea, rbcL, Santiago de Cuba, taxonomy. RESUMEN Antecedentes. Como parte de los estudios monográficos sobre el complejoLaurencia (Rhodomelaceae, Rhodophyta) en Cuba, se llevó a cabo la recolecta e investigación taxonómica detallada de una pequeña especie Laurencia coelenterata, presente en Santiago de Cuba, región oriental del archipiélago cubano. Los caracteres vegetativos y reproductivos pre- sentes en esta especie y que son exclusivos del género Osmundea (miembro del complejo Laurencia) son: dos células periacentrales por cada segmento axial vegetativo, los tetrasporangios nacen al azar de las células corticales y las ramas espermatangiales son de tipo “filamento”.Objetivos . En este trabajo se describen caracteres morfológicos previamente no conocidos para Laurencia coelenterata y se discute su posición taxonómica dentro del complejo Laurencia. Además se analiza la distribución geográfica global de las especies deOsmundea. Métodos. Se realizaron análisis morfológicos y moleculares de secuencias del gen rbcL. Complementándose con la elaboración e interpretación de las hipótesis filo- genéticas. Resultados. El análisis morfológico actual del paratipo (# 4451) resguardado en el Herbario Marino Puertorri- queño mostró que L. coelenterata fue ubicado de manera incorrecta dentro del género Laurencia. La posición filogenética de esta especie fue inferida por secuencias del gene cloroplástico rbcL y el intervalo de variación genética encontrado, también soporta la transferencia de Laurencia coelenterata al género Osmundea, como O. coelenterata (D. L. Ballantine et Aponte) M.T. Fujii, Sentíes et Areces. Conclusiones. La aproximación morfológica y molecular en la taxonomia algal esta permitiendo consolidar el status nomenclatural de las especies, este estudio no es la excepción para la nueva combina- ción de Osmundea coelenterata. Palabras clave: Laurencia sensu lato, Osmundea, rbcL, Santiago de Cuba, taxonomía. Vol. 26 No. 2 • 2016 204 Fujii, M. T. et al. INTRODUCTION Molecular study. Samples for molecular analysis were cleaned, dried, and preserved in silica gel. Total DNA was extracted using a Laurencia coelenterata D. L. Ballantine and Aponte (Ballantine et Apon- DNeasy Plant Mini Kit (QIAGEN, Valencia, CA, USA), according to the te, 1995) is a diminutive species, originally collected from the Dry Tor- manufacturer’s instructions. The rbcL gene was amplified in three over- tugas, Florida, at 9 m depth. According to the authors, the name given lapping fragments with the primer pairs suggested by Freshwater & to it was due to the superficial resemblance of juvenile plants to an Rueness (1994), using the Taq polymerase chain reaction (PCR) Core anemone. Despite this characteristic, the species is not easily found in Kit (QIAGEN). All PCR products were analyzed by electrophoresis in 1% the natural environment because of its small thalli. agarose to confirm product size. The PCR products were purified with The taxonomy of the red algal genus Laurencia J. V. Lamouroux the QIAquick Purification Kit (QIAGEN) according to the manufacturer’s sensu lato is extremely complicated because of the large degree of recommendations. Cycle sequencing was carried out on an ABI PRISM morphological plasticity and worldwide distribution from temperate to 3100 Genetic Analyzer (Applied Biosystems, Carlsbad, CA, USA), using tropical oceans. Consequently, the taxonomic position of the species in the BigDye Terminator Cycle Sequencing Reaction Kit (Applied Biosys- this group has changed rapidly as new morphological and molecular tems). Primers were used for PCR amplification and cycle sequencing. data are processed. In the roughly two hundred years since the creation Sequences were analyzed with Sequence Navigator software, version of the genus Laurencia (Lamouroux, 1813), many changes have been 1.0.1 (Applied Biosystems). proposed in what we currently refer to as the Laurencia complex, made Phylogenetic analyses. Phylogenetic relationships were inferred up of seven formally proposed genera: Laurencia sensu stricto, Osmun- with PAUP* version 4.0b10 (Swofford 2002) and MrBayes v.3.0 beta 4 dea Stackhouse (Stackhouse, 1809), Chondrophycus (Tokida et Saito) (Huelsenbeck & Ronquist, 2001). Maximum parsimony (MP) trees were Garbary et J. T. Harper (Garbary & Harper, 1998), Palisada (Yamada) K. W. constructed by applying the heuristic search option, tree-bisection- Nam (Nam, 2007), Laurenciella Gil-Rodríguez, Sentíes, Díaz-Larrea, Cas- reconnection (TBR) branch swapping, with unordered and unweighted sano et M. T. Fujii (Cassano et al., 2012), and most recently Coronaphy- characters, and gaps treated as missing data. Modeltest software, cus Metti, (Metti et al., 2015). version 3.7 (Posada & Crandall, 1998), was used to find the model of In the Cuban archipelago, the Laurencia complex has 17 species, sequence evolution least rejected in each data set by a hierarchical distributed into 3 genera: Laurencia sensu stricto (11), Palisada (5), and likelihood ratio test. Once the evolution model had determined a Maxi- Yuzurua (1 with two varieties) (Suárez et al., 2015). mum Likelihood (ML), searches were performed by applying the esti- mated parameters (substitution model, gamma distribution, proportion During the Laurencia complex expedition to the eastern coast of the of invariant sites, and frequencies of the bases). Maximum likelihood Cuban archipelago, Laurencia coelenterata was found growing on the analysis was then employed to construct the most likely tree from the consolidated substrata in a biogenic sand beach, with limestone and data set. Maximum Likelihood and MP branch supports were calculated beds of the seagrass Thallasia testudinum Banks ex König at the bottom. by nonparametric bootstrapping analysis (Felsenstein, 1985), as imple- The detailed morphological studies carried out on this material revealed mented in PAUP*. vegetative and reproductive characteristics belonging to the genus Os- mundea, and not to the genus Laurencia (Ballantine & Aponte, 1995). In The general time-reversible model of nucleotide substitution with this study, previously unknown morphological features of L. coelenterata invariant sites and gamma-distributed rates for the variable sites (GTR are revealed that justify its transfer to Osmundea. Emendation to the + I + G) were used for Bayesian analysis (BI). This model was selected species’s original description is provided. The new combination is corro- based on an ML ratio test implemented by the software Modeltest, ver- borated by chloroplast-encoded rbcL gene-sequence analysis. sion 3.06 (Posada & Crandall, 1998), with a significance level of 0.01. For Bayesian analysis,
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