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Relaciones Evolutivas De Las Especies Del Genéro Lucifuga (Ophidiiformes: Bythitidae) En Cuba

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Relaciones Evolutivas De Las Especies Del Genéro Lucifuga (Ophidiiformes: Bythitidae) En Cuba Relaciones Evolutivas de las Especies del Genéro Lucifuga (Ophidiiformes: Bythitidae) en Cuba Item Type Theses and Dissertations Authors Centro de Investigaciones Marinas Universidad de La Habana; Hernández Martínez, D. Download date 26/09/2021 05:16:41 Item License http://creativecommons.org/licenses/by-nc/3.0/ Link to Item http://hdl.handle.net/1834/5530 Centro de Investigaciones Marinas Universidad de la Habana Relaciones Evolutivas de las Especies del Genéro Lucifuga (Ophidiiformes: Bythitidae) en Cuba Trabajo presentado en opción al Título Académico de Maestro en Biología Marina y Acuicultura con mención en Ecología Marina Autor: Lic. Damir Abel Hernández Martínez Tutor: Dr. Erik García Machado La Habana 2005 Resumen Resumen. En el presente trabajo se realizó un estudio de las relaciones evolutivas de las especies representativas de peces del género Lucifuga en Cuba mediante el análisis de sus variaciones genéticas y moleculares. Se estudiaron 11 sistemas proteicos y la secuencia parcial del gen mitocondrial citocromo b (cytb). Los animales fueron colectados en diferentes localidades de la zona occidental de Cuba. La filogenia reconstruida a partir de los caracteres moleculares indica la existencia de dos linajes evolutivos fundamentales cuya divergencia está sustentada por 11 loci proteicos, 73 sinapomorfías mitocondriales y un uso diferencial de los codones que codifican para el aminoácido valina. Uno de los linajes está constituido por las especies L. dentatus, L. dentatus var. holguinensis y L. simile y el otro por las especies L. subterraneus y L. teresinarum. La distribución de las especies en los linajes evolutivos se corresponde con la asignación de las mismas a los dos subgéneros definidos para este género. El análisis filogenético sostiene que las especies L. simile, L. dentatus y L. subterraneus constituyen linajes evolutivos independientes caracterizados por múltiples sinapomorfías, coincidiendo con clasificación taxonómica actual. L. teresinarum no se sustenta como clado monofilético y apunta a ser una sinonimia de L. subterraneus, aunque los resultados al respecto no son concluyentes. Los ejemplares de la denominada L. dentatus var. holguinensis capturados en dos localidades del norte de Matanzas constituyen un clado bien soportado definido por caracteres moleculares, bioquímicos y morfológicos. Esto sugiere la revisión de su estatus taxonómico. Índice Introducción...............................................................................................................................i 1 Revisión Bibliográfica......................................................................................................1 1.1 Clasificación taxonómica del género Lucifuga. .............................................................1 1.2 Características generales del género Lucifuga. ..............................................................1 1.3 Consideraciones sobre las especies que conforman el género. ......................................3 1.3.1 Descripción de las especies. ...................................................................................3 1.3.2 Distribución geográfica de las especies cubanas....................................................5 1.4 Criterios sobre la evolución del género. .........................................................................6 1.5 Filogenia. ........................................................................................................................7 1.6 Caracteres moleculares...................................................................................................8 1.6.1 Proteínas. ................................................................................................................9 1.6.1.1 Métodos electroforéticos. .................................................................................10 1.6.2 ADN mitocondrial. ...............................................................................................12 1.6.2.1 ADN mitocondrial como carácter molecular....................................................13 2 Materiales y Métodos.....................................................................................................15 2.1 Material biológico........................................................................................................15 2.2 Procesamiento de las muestras. ....................................................................................16 2.2.1 Análisis Proteico...................................................................................................16 2.2.1.1 Preparación de los extractos. ............................................................................16 2.2.1.2 Electroforesis....................................................................................................17 2.2.1.3 Características y visualización de los sistemas proteicos estudiados...............17 2.2.2 Análisis de la variación de la secuencia del gen mitocondrial citocromo b (cytb)..18 2.2.2.1 Obtención del ADN total..................................................................................18 2.2.2.2 Reacción en Cadena de la Polimerasa (PCR) y Secuenciación........................19 2.3 Procesamiento Matemático. .........................................................................................20 2.3.1 Proteínas. ..............................................................................................................20 2.3.2 ADN mitocondrial. ...............................................................................................21 3 Resultados. ......................................................................................................................23 3.1 Variación de loci proteicos en tres taxa........................................................................23 3.1.1 Análisis Filogenético. ...........................................................................................26 3.2 Análisis del gen mitocondrial cytb. ..............................................................................27 3.2.1 Análisis de la variación. .......................................................................................27 3.2.2 Saturación de las secuencias.................................................................................32 3.2.3 Análisis Filogenético. ...........................................................................................33 3.2.3.1 Comparación de Topologías.............................................................................34 4 Discusión..........................................................................................................................39 4.1 Relaciones evolutivas...................................................................................................39 4.2 Consideraciones taxonómicas.......................................................................................42 5 Conclusiones....................................................................................................................51 6 Recomendaciones............................................................................................................52 Bibliografía..............................................................................................................................53 Índice Anexos. ................................................................................................................................... 60 Introducción Introducción. El género Lucifuga de la familia Bythitidae, subfamilia Brosmophycinae, está constituido por pequeños peces de cuerpo prolongado y comprimido lateralmente, que carecen de visión y que habitan en cuevas, cacimbas y grietas en aguas desde totalmente dulce hasta marinas (Nielsen et al., 1999). El género está conformado por 6 especies, de éstas, 4 son endémicas de Cuba. Estos peces constituyen los únicos vertebrados estrictamente troglobios que existen en el archipiélago cubano y juegan un papel importante en los ecosistemas donde habitan al ser, generalmente, el eslabón más alto de la cadena trófica (García-Debrás y Pérez, 1999). Al suponerse que el tamaño de las poblaciones es pequeño y producto de la fragilidad del hábitat que ocupan, todas las especies han sido consignadas como vulnerables en el libro rojo de especies amenazadas de la unión internacional para la protección de la naturaleza y los recursos naturales (IUCN Red List, 2000). Aunque el descubrimiento de las primeras especies del género data de más de un siglo, el interés que estos peces han despertado en la comunidad científica ha sido esporádico. Una parte importante de los estudios (Poey, 1858; Gill, 1863; Cohen y Robins, 1970; Nalbant, 1980; Díaz et al., 1987a; Díaz, 1988; Cohen y McKoster, 1998) se han centrado en la discusión de los caracteres morfológicos que definen el género y las especies, y unos pocos trabajos han abordado las relaciones filogenéticas de las especies sobre la base de caracteres morfológicos (Vergara, 1980, 1981). Los criterios más generalizados reconocen a la especie de la Bahamas, L. speleotes, como la más ancestral (Cohen y Robins, 1970; Vergara, 1980; Díaz, 1988) y a las especies cubanas más recientes siguiendo un gradiente de especiación desde la más plesiomórfica (L. dentatus), hasta las más apomórficas y recientes (L. subterraneus y L. teresinarum) (Vergara, 1980, 1981; Díaz, 1988). Sin embargo, algunos de los caracteres morfológicos utilizados para la definición de las especies y la inferencia de sus relaciones se comportan
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