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Neréididos (Polychaeta: Nereididae) Del Gran Caribe

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Neréididos (Polychaeta: Nereididae) Del Gran Caribe El Colegio de la Frontera Sur Neréididos (Polychaeta: Nereididae) del Gran Caribe TESIS Presentada como requisito parcial para optar al grado de Maestría en Ciencias en Recursos Naturales y Desarrollo Rural por Víctor Manuel Conde Vela 2015 Para las familias Conde Vela y Te Gómez En especial para ti, Astrid AGRADECIMIENTOS Esta contribución es resultado del esfuerzo sumado de muchas personas, quienes aportaron de diversas maneras a su culminación. Dedicar el tiempo necesario a este trabajo no sólo costó al autor, sino también a quienes lo apoyaron y prestaron su valioso tiempo y esfuerzo. Gracias por todo, y es para ustedes. Quiero expresar mi más sincero agradecimiento a mi director de tesis Sergio I. Salazar Vallejo, ya que sin su apoyo y su vasta experiencia en el terreno de los poliquetos, esta contribución no hubiera tenido los mismos resultados. Gracias por la insistencia en hacer cada vez más y mejor, y por los valiosos comentarios y revisiones que me guiaron durante toda la maestría. A Luis F. Carrera Parra y a Jesús Á. de León González, por aceptar ser parte del comité evaluador, y proporcionar valiosos comentarios que permitieron mejorar el escrito final. A mis sinodales adicionales Eduardo Suárez Morales, Víctor H. Delgado Blas y David González Solís, por aceptar evaluar el escrito en tan apretadas situaciones, y por sus importantes sugerencias que proporcionaron al mismo. A Patricia Bardales Pastrana por las innumerables facilidades administrativas proporcionadas. A Nancy Voss (UMML) por facilitar el material base de esta contribución. Gracias a su confianza y el préstamo del material de manera incondicional, yo y varios colegas hemos podido realizar nuestras metas en maestría y doctorado. A Karen Osborn y Geoff Keel (USNM) quienes facilitaron mi estancia de investigación en el Smithsonian Institution (Washington, EUA), y a Leslie Harris por mi estancia en el Natural History Museus of Los Angeles County (LACM-AHF). A todos ellos gracias por la confianza y el acceso ilimitado a las colecciones a su cargo. A Jesús Á. de León Gónzález (UANL) y a Leslie Harris por prestar material valioso para esta contribución. A Leslie Harris y David Ocker por la cálida hospitalidad y diversas enseñanzas durante mi estancia en Los Ángeles. Agradezco enormemente a mi esposa Astrid Estefanía Te Gómez, ya que sin su completo apoyo este trabajo no hubiera sido posible. Muchas gracias por estar ahí cuando más lo necesitaba. A las familias Conde Vela y Te Gómez, quienes siempre me han apoyado de innumerables maneras, y por dar parte de su tiempo y recursos para la realización de este trabajo, sin los cuales las metas de este trabajo no se hubieran alcanzado. A los colegas del laboratorio, Emilia, Reyna, Isabel, Jani, Stephanie, Tulio y Víctor Hugo, que hicieron más amena la estadía en ECOSUR, además de compartir sus experiencias y conocimientos. De manera especial, agradezco a todos mis compañeros de la maestría, que tuvieron muchas oportunidades de convivir conmigo y hacer esta maestría la mejor que pude haber tenido. Agradezco a CONACyT por la beca que me otorgó, sin la cual no se hubiera podido dedicarme a mis estudios de maestría. Además a Sergio I. Salazar Vallejo y Luis F. Carrera Parra, quienes aportaron recursos importantes para la realización de las estancias en el extranjero. CONTENIDO Introducción …………………………………………………………………………………… 1 Capítulo 1. Redescriptions of Nereis oligohalina (Rioja, 1946) and N. garwoodi González-Escalante & Salazar-Vallejo, 2003 and description of N. confusa sp. n. (Annelida, Nereididae) ……………………………..………………………….……….……… 7 Capítulo 2. Nereididae (Polychaeta: Phyllodocida) from the Grand Caribbean and other related regions ………..…………………..………………………………………………….. 43 Conclusiones ………………………………………………………………….…………… 282 Referencias ………………………………………………………………….……………… 285 Anexos ………………………………………………………………………………………. 292 La presente tesis no debe ser considerada como una publicación en el sentido del Código Internacional de Nomenclatura Zoológica, y los nombres científicos mencionados en ésta no deben ser citados en ninguna forma, para evitar ser considerados nomina nuda. This thesis is not a publication in the sense of the International Code of Zoological Nomenclature, and scientific names mentioned in it must not be cited in any way to avoid them to become nomina nuda. INTRODUCCIÓN La familia Nereididae de Blainville, 1818 es un grupo amplio de poliquetos que se distingue por tener una faringe tubular eversible con papilas y/o paragnatos en un arreglo regular, aunque pueden carecer de ambos, así como un par de grandes mandíbulas en su extremo, además de presentar una morfología parapodial compleja, con lóbulos y lígulas. Los neréididos representan una de las familias más comunes en diversos sustratos; tienen una amplia distribución, debido a que son frecuentes han sido extensamente estudiados (Wilson, 2000). Actualmente, la familia tiene 44 géneros y unas 460 especies (de León-González, 2009), comprendidos en tres subfamilias: Nereidinae de Blainville, 1818, Namanereidinae Hartman, 1959, y Gymnonereidinae Banse, 1977 (Fitzhugh, 1987; Glasby, 1991). Bakken y Wilson (2005) y Santos et al. (2005) notaron algunos problemas en la clasificación actual a distintos niveles, como el bajo soporte filogenético de las subfamilias actuales, el posible restablecimiento de dos subfamilias sinonimizadas, Dendronereidinae Pillai, 1961 y Notophycinae Knox & Cameron, 1970. En América tropical se han citado 25 géneros y 104 especies de neréididos (de León- González, 2009). Sin embargo persisten problemas taxonómicos en el grupo, principalmente la supuesta existencia de un gran número de especies consideradas cosmopolitas o de amplia distribución; de especial interés son las que se consideran como anfiamericanas, es decir, que viven en las costas del Pacífico y Atlántico americano, donde al menos 18 están en esa categoría (de León-González, 2009). 1 En el Gran Caribe, que comprende el Golfo de México, el Mar Caribe, Bermudas y las costas nortes de Brasil (Salazar-Vallejo, 2000), el estudio exhaustivo de la poliquetofauna del Gran Caribe tuvo su origen en las campañas oceanográficas puntuales o de alcance mundial, como las expediciones del Challenger por el escocés Charles W. Thomson (McIntosh, 1885), las realizadas por el estadounidense Alexander Agassiz en el Mar Caribe y el Golfo de México (Ehlers, 1887; Augener, 1906), y por los daneses Anders S. Ørsted y Henrik Kröyer en el Mar Caribe, incluyendo América Central y Sudamérica (Grube, 1857, 1858, 1859). Otras campañas importantes han sido las Expediciones del Atlantis a las Indias Occidentales (Hartman, 1942), la Expedición de Barbados-Antigua de la Universidad de Iowa en 1918 (Treadwell, 1924), las Expediciones Smithsonian-Bredin en el Mar Caribe (de León-González, Solís-Weiss y Ochoa-Rivera, 1999), y las Expediciones del Mar profundo de la Universidad de Miami, realizadas por Gilbert L. Voss, Richard Robins y C. Staiger en los años 1962 a 1975. Esta última es la principal fuente de especímenes para esta investigación. Además, otras expediciones danesas sirvieron para conocer y describir varios neréididos de agua dulce y de depósitos de agua continentales, principalmente las dirigidas por P. W. Hummelinck en gran parte de las Antillas (Wesenberg-Lund, 1958; Hummelinck, 1981). A pesar de que nuevas especies fueron descritas a partir del material colectado en aquellas expediciones y en otros trabajos puntuales, muchas de éstas fueron sinonimizadas con especies de otras partes del mundo. La falta de descripciones actualizadas en varias especies, basadas en la revisión de materiales originales o tipo de las especies involucradas, ocasionó la introducción de nombres foráneos en listados 2 regionales. Debido a la tendencia contemporánea a la conservación del nombre más antiguo, especialmente después del esfuerzo de Olga Hartman (Hartman, 1959a, b) en su catálogo mundial de poliquetos, los nombres regionales del Gran Caribe fueron considerados como variantes o subespecies, esto aunado a que muchas descripciones originales son breves, con la consecuente pérdida de caracteres diagnósticos. Además algunas especies fueron sinonimizadas con otras, a pesar de que fueron parte de una revisión más profunda (e.g. Pettibone, 1971). Diversos autores han abordado el problema en varios grupos para el Gran Caribe con estudios morfológicos finos, herramientas moleculares o una combinación de ambas, y han demostrado que la conclusión de la supuesta amplia distribución se trata de una mala taxonomía o de estudios sistemáticos conservadores, es decir, con poca introducción de caracteres novedosos que permitieran separar a las especies (e.g. Klautau et al., 1999; Carrera-Parra y Salazar-Vallejo, 2011; Yáñez-Rivera y Carrera-Parra, 2012; Yáñez- Rivera y Salazar-Vallejo, 2011). En el caso de los neréididos, se han realizado varios esfuerzos para describir las especies de neréididos presentes a nivel local, principalmente de las costas caribeñas de México (de León-González y Solís-Weiss, 1997, 1998; de León-González, Solís-Weiss y Ochoa- Rivera, 1999; de León-González, Solís-Weiss y Valadez-Rocha, 2001; Salazar-Vallejo y Jiménez-Cueto, 1997), Venezuela (e.g. Liñero-Arana y Reyes-Vásquez, 1979; Liñero- Arana y Díaz-Díaz, 2007; Vanegas-Espinosa, Díaz-Díaz y Liñero-Arana, 2007) y Brasil (Santos y Lana, 2000, 2001, 2003; Santos, 2007). Amaral et al. (2013) realizaron una compilación de las especies de poliquetos registrados para Brasil, junto con la gran cantidad de literatura generada. Sin embargo, son pocos los trabajos
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