Juan Francisco Saborido Rey EL GÉNERO SEBASTES Cuvier, 1829

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Juan Francisco Saborido Rey EL GÉNERO SEBASTES Cuvier, 1829 Juan Francisco Saborido Rey EL GÉNERO SEBASTES Cuvier, 1829 (PISCES, SCORPAENIDAE) EN EL ATLÁNTICO NORTE: IDENTIFICACIÓN DE ESPECIES Y POBLACIONES MEDIANTE MÉTODOS MORFOMÉTRICOS; CRECIMIENTO Y REPRODUCCIÓN DE LAS POBLACIONES EN FLEMISH CAP. Tesis Doctoral Universidad Autónoma de Madrid Facultad de Biología Departamento de Zoología 1994 Universidad Autónoma de Madrid EL GÉNERO SEBASTES Cuvier, 1829 (PISCES, SCORPAENIDAE) EN EL ATLÁNTICO NORTE: IDENTIFICACIÓN DE ESPECIES Y POBLACIONES MEDIANTE MÉTODOS MORFOMÉTRICOS; CRECIMIENTO Y REPRODUCCIÓN DE LAS POBLACIONES EN FLEMISH CAP. Memoria de Tesis Doctoral Memoria de Tesis Doctoral que presenta Juan Francisco Saborido Rey para optar al grado de Doctor en Ciencias Biológicas Madrid, 1994 D. ANTONIO VÁZQUEZ RODRÍGUEZ, Colaborador Científico del Instituto de Investigacións Mariñas (Consejo Superior de Investigaciones Científicas) CERTIFICA: Que la presente memoria titulada “El Género Sebastes Cuvier, 1829 (Pisces, Scorpaenidae) en el Atlántico Norte: identificación de especies y poblaciones mediante métodos morfométricos; crecimiento y reproducción de las poblaciones en Flemish Cap”, que para optar al grado de Doctor presenta D. Juan Francisco Saborido Rey, ha sido realizada bajo mi dirección en el Instituto de Investigacions Mariñas (CSIC) de Vigo. Y considerando que constituye un trabajo que reúne las condiciones necesarias de una Tesis, autorizo su presentación. Y para que así conste, se expide el presente certificado en Vigo, en noviembre de mil novecientos noventa y cuatro. Dr. Antonio Vázquez Rodríguez A mis padres A mis hermanos AGRADECIMIENTOS A lo largo de los años necesarios para realizar este trabajo han sido numerosas las personas que, de una u otra forma, han colaborado a que fuese posible. Vaya de antemano a todas ellas mi más sincero agradecimiento. En especial quiero destacar: Al Dr. Antonio Vázquez, director de esta Tesis, por darme la oportunidad de iniciarla, por la minuciosa corrección del manuscrito y por sus consejos y orientaciones, sin los cuales, sin duda, este trabajo jamás habría visto la luz. A la Dra. Susana Junquera, del Instituto Español de Oceanografía, por sus valiosas sugerencias en todos los campos que abarca este trabajo, por sus consejos, siempre acertados, por su apoyo y amistad, siempre constantes; por ser, además, la persona que, por su dedicación y esfuerzo en el trabajo, me ha servido de guía todo este tiempo. Al Dr. Germán Perez-Gándaras, que desde el inicio me introdujo y orientó en el tema de morfometría y estadística. Al Dr. Kjell Nedreaas, del Havforskningsinstituttet (Instituto de Investigaciones Marinas) de Bergen, por sus consejos en la identificación de las gallinetas de Noruega, la información sobre la edad y crecimiento y por proporcionarme las muestras procedentes de Noruega, además de permitirme asistir a una de sus campañas de investigación. A mis compañeros del equipo Pesquerías Lejanas del Instituto de Investigaciones Marinas: Dr. Javier Zamarro, Dr. Javier Paz y en especial Mikel Casas, compañero de fatigas. A Carmen Alonso y a Rosa Collazo, por las largas y tediosas horas de laboratorio que pasamos juntos; sin su ayuda aún estaría haciendo este trabajo. A María González, por su inestimable ayuda en la realización del trabajo histológico, por su amistad y simpatía. i A todos mis compañeros del Instituto de Investigaciones Marinas, en especial a Marigel García, a Isabel Cunha, y a los Drs. Ángel Guerra y Roberto Munilla. La mayor parte del material analizado en este trabajo se recogió a lo largo de diez campañas de investigación. Quisiera agradecer a las tripulaciones de los buques oceanográficos: Cryos, Ignat Pavlyuchenkov, G. O. Sars, Cornide de Saavedra y Atlántida, el apoyo obtenido. Así mismo, sin la participación de mis compañeros en cada una de las campañas hubiera sido imposible recoger las muestras, gracias a todos ellos. Parte del material lo recogieron tres biólogos a bordo de otros tantos buques comerciales, quisiera agradecer a Alberto Riveiro, Santiago Cerviño y a Rafael Bañón su colaboración, especialmente a este último, compañero además en cinco campañas oceanográficas. Este estudio se realizó dentro de los proyectos: “Bottom trawl survey of the Flemish Cap” (proyecto de la Unión Europea MA 1-246) y “Analysis of exploited fish stocks on Flemish Cap” (proyecto de la Unión Europea AIR1-CT92-0558) realizados conjuntamente por el Instituto de Investigaciones Marinas (CSIC), el Instituto Español de Oceanografía y el Instituto Portugués de Investigaçáo Marítima (Lisboa) y financiado por la Unión Europea y los citados organismos; mi agradecimiento a todos los que lo hacen posible. He dejado para el final, por ser los más importantes, a mis padres y hermanos, hacia los cuales no encuentro otras palabras para expresarme más que tres: gracias, os quiero. ii Tabla de Contenido AGRADECIMIENTOS i TABLA DE CONTENIDO iii LISTA DE FIGURAS vii LISTA DE TABLAS ix INTRODUCCIÓN 13 DESCRIPCIÓN GENERAL 17 MORFOLOGÍA 17 Musculatura de la vejiga natatoria 21 BIOLOGÍA 25 Reproducción 25 Crecimiento 27 Alimentación 28 SISTEMÁTICA 28 Sistemática del género en el Atlántico Norte 29 DISTRIBUCIÓN 31 Atlántico Norte 33 Poblaciones de gallineta 35 IDENTIFICACIÓN DE ESPECIES Y DELIMITACIÓN DE POBLACIONES MEDIANTE MÉTODOS MORFOMÉTRICOS 37 INTRODUCCIÓN 39 MÉTODOS DE IDENTIFICACIÓN DE POBLACIONES 40 MATERIAL Y MÉTODOS 43 MATERIAL 43 MÉTODOS 50 Análisis morfométrico 50 Exploración previa de los datos 50 Detección de datos incoherentes 50 Normalidad 54 El problema de la talla y la forma 56 Análisis discriminante 65 Comprobación de la igualdad de grupos centroides y varianzas 69 Dimorfismo sexual 70 Análisis de caracteres merísticos 71 Correlación con la talla 71 Dimorfismo sexual 72 Comparación de especies y poblaciones 73 RESULTADOS* 75 COMPARACIÓN DE MÉTODOS PARA ELIMINAR LA INFLUENCIA DE LA TALLA 75 iii ANÁLISIS MORFOMÉTRICO 79 Dimorfismo sexual 79 Comparación de poblaciones de Sebastes en el Atlántico Norte 81 Comparación por áreas geográficas 86 Costa de Noruega 86 Banco de Flemish Cap 88 Norte del Gran Banco de Terranova 89 Banco de Saint Pierre 92 Comparación intraespecífica 93 Sebastes marinus 93 Sebastes mentella 95 Sebastes fasciatus 97 Sebastes fasciatus vs S. viviparus 99 ANÁLISIS DE CARACTERES MERÍSTICOS 100 Dimorfismo sexual 101 Comparación por áreas geográficas 101 Costa de Noruega 101 Flemish Cap 103 Norte del Gran Banco de Terranova 104 Banco de Saint Pierre 105 Comparación intraespecífica 106 Sebastes marinus 106 Sebastes mentella 111 Sebastes fasciatus 112 Sebastes fasciatus vs S. viviparus 113 DISCUSIÓN 139 FLEMISH CAP 157 INTRODUCCIÓN 159 DESCRIPCIÓN DE FLEMISH CAP 159 OCEANOGRAFÍA 159 DESCRIPCIÓN DEL ECOSISTEMA 161 OBJETIVOS 164 MATERIAL Y MÉTODOS 165 MATERIAL 165 MÉTODOS 166 Edad y crecimiento 166 Reproducción 170 EDAD Y CRECIMIENTO 175 RESULTADOS 176 Escamas vs otolitos 176 Validación del criterio de lectura 179 Comparación del crecimiento entre sexos 182 Comparación intraespecífica del crecimiento 189 DISCUSIÓN 190 REPRODUCCIÓN 199 RESULTADOS 199 Ovogénesis 199 Folículos postovulatorios 203 iv Ovocitos en atresia 203 Desarrollo embrionario 204 Ciclo reproductivo 206 Talla de primera maduración 209 Proporción de sexos 210 DISCUSIÓN 213 CONCLUSIONES 235 FOTOGRAFÍAS 241 BIBLIOGRAFÍA 253 v Lista de Figuras Figura 1. Morfología externa de la gallineta . Dibujo de S. mentella idealizado siguiendo la Tabla 1 ................................................................................................................................................... 19 Figura 2. Musculatura de la vejiga natatoria. Los números representan las vértebras; las dos primeras vértebras no tienen costillas......................................................................................... 23 Figura 3. Clasificación de las especies a estudio................................................................................. 29 Figura 4. Distribución mundial del género Sebastes. ........................................................................... 32 Figura 5. Distribución del género Sebastes en el Atlántico Norte. ...................................................... 32 Figura 6. Procedencia de las muestras analizadas. A) Archipiélago de Svalbard y costa de Noruega. B) Flemish Cap y Gran Banco de Terranova. C) Saint Pierre..................................................... 45 Figura 7. Distancias medidas en cada individuo.................................................................................. 48 Figura 8. Diagrama de los centroides de los grupos respecto a las tres primeras variables canónicas. .................................................................................................................................. 83 Figura 9. Puntuaciones discriminantes individuales de las once poblaciones consideradas................. 84 Figura 10. Matriz de amalgamiento..................................................................................................... 85 Figura 11. Puntuaciones discriminantes individuales de las poblaciones de Sebastes en la costa de Noruega. Las letras M, T y V representan la posición de los centroides de S. marinus, S. mentella y S. viviparus respectivamente...................................................................................... 88 Figura 12. Puntuaciones discriminantes individuales de las poblaciones de Sebastes en Flemish Cap Las letras M, T y F representan la posición de los centroides de S. marinus, S. mentella y S. fasciatus respectivamente...........................................................................................................
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