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Avances En El Estudio Sistemático De La Familia Galaxiidae (Osteichthys: Salmoniformes)

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Avances En El Estudio Sistemático De La Familia Galaxiidae (Osteichthys: Salmoniformes) Arch. Biol. Med. Exper. 12:107-118, 1979 Avances en el estudio sistemático de la familia Galaxiidae (Osteichthys: Salmoniformes) Advances in the systematic study of Galaxiidae HUGO CAMPOS C. Instituto de Zoología, Universidad Austral de Chile, Casilla 567, Valdivia, Chile. (Recibido para publicación el 7 de septiembre de 1977) CAMPOS, H.C. Avances en el estudio sistemático de la familia Galaxiidae {Osteichthys: Salmo- niformes). (Advances in the systematic study of Galaxiidae (Osteichthys: Salmoniformes). Arch. Biol. Med. Exper. 12: 107-118, 1979- The principal results obtained at the systematic study of the family Galaxiidae were revised. This family is distributed on the South of continents and islands around the Antarctic (South America, South Africa, Australia and New Zealand), with more or less 40 species and 5 genus. The family, genus and species present great phylogenetic and biogeographic questions in relation to their center of origin and dispersion. One species, Galaxias maculatus is common to a great part of the family-distribution (South America, Australia and New Zealand) and its way of dispersion is being discussed. A discussion of systematic advances obtained in Galaxiids is done by characters of external and internal Morphology, Cytogenetic, Electrophoresis, Analysis of Multivariance, Ecology and Zoogeography. GALAXIIDAE SYSTEMATIC GEOGRAPHIC DISTRIBUTION Las especies de la familia Galaxiidae tienen una los avances alcanzados en esta familia y las meto­ amplia distribución en las zonas temperadas de dologías empleadas. los continentes que rodean la Antartica. Su distribución está circunscrita a las aguas con­ SISTEMÁTICA tinentales del sur de Sudamérica, Sudafrica, Australia y Nueva Zelandia. Esta distribución De acuerdo con Simpson (48) consideramos presenta problemas sistemáticos, zoogeográ- que la Sistemática es una ciencia que estudia ficos y evolutivos, lo que permite aplicar teorías las especies o diversidad de organismos esta­ de larga discusión en estas ciencias. Otros gru­ bleciendo las relaciones entre ellas. Así la dife­ pos de animales y plantas muestran una distri­ renciamos de Clasificación que es más bien un bución semejante en el Hemisferio Sur a los ordenamiento de los organismos dentro de galaxüds y el conocimiento en ellos logrados grupos y de Taxonomía que sería un estudio teó­ son un antecedente comparativo. El estudio de rico de la Clasificación estableciendo sus bases, esta Familia contribuye a ampliar los conoci­ principios, procedimientos y reglas. La diver­ mientos de la fauna originada en el Hemisferio sidad tiene como base el carácter en un amplio Sur y la importancia que ha tenido la Antartica concepto, desde lo morfológico a lo funcional. en su distribución actual. El carácter es el punto de partida para construir El objetivo del presente trabajo es revisar el Taxón entendido como un grupo de individuos 107 108 CAMPOS de alguna naturaleza o rango. Todo carácter porque proviene del concepto tipológico. Estos tiene en último término un registro genético que caracteres morfológicos externos son emplea­ está sometido a todos los intrincados caminos dos en la mayoría de los casos como únicos cri­ que nos enseñan la Genética y la Evolución. Su terios clasificatorios, por la facilidad de recono­ manifestación fenotípica es el instrumento del cer las especies a la vista o tacto y por la facilidad sistemático y depende del análisis o interpreta­ de trabajar con especímenes conservados tion que de él se haga, el nivel que le otorga. En en colecciones. Además, a muchos caracteres muchos casos los (actores bióticos o abióticos se les da un alto valor cualitativo cuando real­ configuran una diversidad de gran magnitud en mente sus rangos de variación pueden ser dudo­ grupos básicos. Esta diversidad origina confu­ sos y sea necesario aplicar criterios estadísti­ sión en la clasificación y sistemática, que llevan cos cuantitativos. (Ej.: caracteres merísticos). finalmente a falsas interpretaciones filogené- Estos caracteres pueden mostrar marcada di­ ticas y evolutivas. Un aspecto relevante es la rela­ versidad entre un grupo y otro, permitiendo ción entre el carácter y el lugar donde se encuen­ una acertada clasificación. Sin embargo, cuan­ tre el grupo, estableciéndose una estrecha unión do la diversidad es menor como sucede en grupos entre Biogeografía con Ecología y Evolución. muy parecidos, caracteres que son simples varia­ Los peces de aguas continentales están es­ ciones intraespecíficas, son elevados a un nivel trechamente unidos al origen geológico de los específico o genérico. Así, muchas especies cursos de aguas que habitan, adquiriendo por o géneros son producto de la experiencia del in­ lo tanto gran valor su distribución geográfica, vestigador basado en criterios subjetivos o apre­ en relación con sus procesos de especiación ciaciones comparativas. Las especies de la fami­ (Ej.: aislamiento geográfico). Conociendo la lia Galaxiidae son un buen ejemplo de esta situa­ edad de algunos lagos se ha podido suponer la ción. edad actual de algunas especies. También el nú­ mero de especies está correlacionado con la ma­ Desde la descripción de la primera especie yor o menor diversidad del medio de acuerdo de galaxiid hecha por Gmelin, en 1789, en base con algunas reglas ecológicas (55, 56). a un ejemplar colectado por el capitán Cook en La familia Galaxiidae tiene alrededor de 40 1772 (Nueva Zelandia), hasta las monografías especies en cinco géneros. El valor dado a los realizadas (25, 26, 28) para Nueva Zelandia, caracteres para determinar sus especies han si­ Sud América y Sud África, hay una larga histo­ do diversos, originando una gran confusión en ria sistemática. Muchos géneros se crearon la creación de muchas especies y géneros en (Mesistes, Austrocobitis, Agalaxias, Queri- los distintos lugares de su amplia distribución. galaxias, Lyragalaxias y otros) que posterior­ Esta situación ha sido en parte paulatinamente mente pasaron a sinonimia, sucediendo lo mismo superada con el empleo de métodos cuyas apli­ con las especies (cerca de 44 en Australia, 16 en caciones han dependido de la complejidad feno- Sud América, 17 en Nueva Zelandia, 4 en Sud típica del grupo. Sin embargo, muchos proble­ África). Actualmente se reconocen los géneros mas de interpretación zoogeográficas y evoluti­ Galaxias, Brachygalaxias, Xeochana, Para- vas son motivo de controversias. Es ilustrativo galaxias, Saxilaga y Nesogalaxias. Las especies para el estudio de estos peces analizar los caminos se han reducido a cerca de 1~ en Australia. 1 en seguidos en la sistemática de esta familia. Estos Nueva Caledonia, 13 en Nueva Zelandia, 4 en caminos los hemos resumido en Morfología Sud América y 1 en Sud África. En general externa, Morfología interna, Citogenética, las galaxnds se caracterizan por un cuerpo des­ Electroforesis y Análisis de Multivarianza. nudo (sin escamas) teniendo muy desplazado hacia la región posterior del cuerpo las aletas Anal v Dorsal. El Género Galaxias CUVKT (10) Morfología extema cuenta con el mayor número de especies, alre­ dedor de 32, Brachygalaxias Eigenmann (11) La diversidad de las estructuras anatómicas ex­ con 3 especies, Neochana Gúnther (15) 3 espe­ ternas de los galaxuds han sido consideradas cies, Paragalaxias Scott (44) con 1 especie, Saxila­ principalmente como caracteres para clasificar ga Scott (45) con 2 especies y Nesogalaxias los individuos en especies. Este método es lo que Whitley (59) con 1 especie. Entre las muchas podríamos llamar clásico en la Sistemática AVANCES EN SISTEMÁTICA DE GALAXIIDAE 109 causas que motivaron esta gran confusión en la Paragalaxias shannonensis Scott Familia, ha sido la semejanza morfológica de las Saxilaga cleaveri Scott especies y presencia de un estado juvenil Saxilaga upcheri Scott. (Whitebait) que varias veces fue descrito como especie. Morfología interna Lista preliminar de las especies de galaxiids en sus lugares de distribución. Los caracteres basados en la anatomía interna parecen presentar mayor constancia en las Nueva Zelandia (32): poblaciones, lo que permite generalmente ca­ Galaxiasfasciatus Gray racterizar Taxa superiores. Los caracteres os­ Galaxias argénteus Gmelin teológicos son los que han aportado mayores Galaxias postvectis Clarke antecedentes para diferenciar géneros, familias Galaxias gracilis McDowall u otros Taxa y son los referidos a la cabeza, cintu­ Galaxias maculatus Jenyns ras pectorales, cinturas pélvicas, el armazón de la Galaxias brevipinnis Günther aleta caudal y la columna vertebral. Galaxias vulgaris Stokell En las especies de la familia Galaxiidae, poco Galaxias divergens Stokell se consideró este tipo de caracteres, lo que mo­ Galaxiaspaucispondylus Stokell tivó especialmente la confusión en las descrip­ Galaxias prognathus Stokell ciones genéricas. Desde las primeras descrip­ Xcachaña apoda Günther ciones se hicieron algunas observaciones aisla­ Neochana diversa Stokell das en las características óseas (42), sin embar­ Neochana burrowsia Phillipps go faltaba un trabajo comparativo que estudiara la relación de la familia Galaxiidae entre las fa­ Sudamérica (26) milias del orden Salmoniformes. Este trabajo Galaxias maculatus Jenyns lo realizó Me Dowall (24) con una comparación Galaxias platel Steindachner entre las familias e interesantes conclusiones Galaxiasglobiceps Eigenmann sobre su origen, concluyendo que Galaxiidae Brachygalaxias bullocki Regan y Retropinidae son unidades filéticas natura­ les y la familia Aplochitonidae se divide en Pro- Sudáfrica (28) totroctidae y Aplochitonidae. Rosen (43)
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