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Relaciones Biogeográficas De La Avifauna Cubana. Ii Biogeografía Descriptiva1

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Relaciones Biogeográficas De La Avifauna Cubana. Ii Biogeografía Descriptiva1 ORNITOLOGIA NEOTROPICAL 14: 441–467, 2003 © The Neotropical Ornithological Society RELACIONES BIOGEOGRÁFICAS DE LA AVIFAUNA CUBANA. II BIOGEOGRAFÍA DESCRIPTIVA1 Ricardo Vergara R. Apartado Postal 4126, Z. P. Habana 10400, La Habana, Cuba. Abstract. – Biological relationships of Cuban avifauna. II. Descriptive biogeography. – The main general properties that characterize the Cuban avifauna are discussed, chiefly regarding its similarities and differences with those of the other Antilles. The most outstanding biogeographical patterns concerning its composition are reviewed, emphasizing the evolutionary implications. Two different avian biogeographical divisions for the Antilles are proposed on the basis of their tectonical delimitation (on both sides of the Cayman Trough): the North American and the Caribbean; the first one with lower carrying capacity, which account for the likewise lower taxonomical diversity. The concepts of evolutionary valence (of an area), index of species density, and semiharmonic fauna are introduced. It is concluded that in the Cuban avi- fauna: 1) the North American elements are broadly predominant; 2) the generalized species outnumber the specialized forms due to the fact that the stochastic environmental factors are more intense than are those that are deterministic; and 3) has the lower index of species density and interesting biogeographical charac- teristics such as territorial exclusion, and discontinuous distribution of some species as well as relicts in restricted areas, the presence of predatory and large-sized species, which are not uncommon, in spite of the insular conditions with not completed ecological harmony. Accepted 19 April 2003. Resumen. – Se describen las principales características de la avifauna cubana actual, sus relaciones coroló- gicas y factores causales de su evolución. Se establecen dos divisiones ornitogeográficas para las Antillas: la norteamericana y la caribeña, separadas por la depresión de Caimán. La primera, integrada por el Archipié- lago Cubano y las islas Bahamas y Caimán, se caracteriza por su menor capacidad limitativa ambiental y consiguiente pobreza de la diversidad taxonómica. Dentro de la avifauna cubana, el grupo corogénico nor- teamericano tiene mayor representatividad que el sudamericano, y la mayoría de sus especies son altamente generalistas en un ambiente estocástico. Se introducen los conceptos de valencia evolutiva (de un área) y fauna semiarmónica. Se concluye que el Archipiélago cubano: 1) presenta un número absoluto de especies mayor que el de las otras Antillas, pero menor en lo relativo a la densidad de especies por área; 2) fue un centro de evolución ornítica en atención al número de fósiles, endémicos y semiendémicos satélites en las áreas próximas; y 3) presenta interesantes características biogeográficas, tales como: exclusión territorial y distribución discontinua de algunos especies constituyentes, así como de relictos en áreas muy restringidas. Key words: Biogeography, birds, Cuban avifauna, Antillean avifauna, Cuban Archipelago, Antilles. INTRODUCCIÓN referente a la distribución geográfica de sus especies constituyentes con las de las otras En esta segunda contribución acerca de la avi- Antillas, a fin de conocer su delimitación fauna cubana se compara la información regional y propiedades diferenciales generales. ______________ Los objetivos son: 1) determinar los factores 1Dedicado a la memoria de mi profesor, el Dr. causales de su diversidad; 2) discernir, con el Abelardo Moreno Bonilla. mayor grado de precisión posible, sus relacio- 441 VERGARA nes corológicas; y 3) establecer su caracteriza- ricano, de edad cretácico-terciaria (“área atá- ción, especialmente con relación la avifauna vica”; Croizat-Chaley 1975, Rosen 1985), antillana. aquí llamado área primordial centroameri- Se tiene en cuenta la importancia primor- cana, comprendida desde el sur de los Esta- dial de: 1) los eventos geohistóricos, principal- dos Unidos hasta el nordeste de Sudamé- mente en su relación con la saturación rica. Los procesos tectónicos (acreciones de taxonómica y con las tasas de renovación, de placas y subducciones) y las fluctuaciones acuerdo con las conclusiones del estudio glacioeustáticas, con las consiguientes expan- precedente (Vergara 1988a); 2) la localización siones y contracciones de los biótopos, ocu- del potencial genético (cf. Fundamentación rridas dentro de dicha área, dieron lugar a y terminología) sobre las cuestiones relativas los procesos de vicarianza y progresión coro- a las extensiones de las áreas y distancias lógica ordenada (sensu Vergara 1988b), deter- existentes entre las mismas; y 3) la influ- minativos de la formación de la avifauna encia de los factores climáticos, determinati- cubana. vos de la estabilidad ambiental y consiguiente viabilidad de las especies. Se introducen las FUNDAMENTACIÓN Y TERMINO- subespecies solamente cuando las mismas LOGÍA presentan gran importancia biogeográfica, como endémicas o indicadoras de relaciones En el curso de este estudio se emplearán, en corológicas. gran medida, las iniciales de los conceptos y El estudio de la avifauna cubana reviste definiciones aparecen a continuación: gran interés para la cuestiones descriptivas de la biogeografía (comparativas y de estableci- Archipiélago cubano. El conjunto integrado por miento de delimitaciones regionales; Brundin la isla de Cuba, la Isla de la Juventud (antigua 1975). A causa de que después de la ultima Isla de Pinos) y las islas e islotes (“cayos”) glaciación, sus especies constituyentes se adyacentes. presentan con: 1) un número relativamente bajo en un área insular tropical y extensa; Área de concentración de potencial genético. De 2) una adecuación genética a condiciones acuerdo con los planteamientos de Croizat- de incertidumbre ambiental, con el desarrollo Chaley (1975) es un centro de diferenciación de características comunes que permiten dis- taxonómica (“nudo”) donde se encuentra el tinguirlas regionalmente; 3) bastante frecuen- grueso de los descendientes de formas ances- cia, en áreas periféricas restringidas y trales de alta potencialidad evolutiva y que ampliamente separadas; además, algunas de determina la distribución actual de los mis- ellas tienen condición de endémicas que, mos. por sus diferenciaciones taxonómicas y tem- porales, son indicadores de relaciones coroló- Valencia evolutiva (de un área). Es la concordan- gicas. cia de la relación del potencial genético y las Se consideran principalmente las especies inducciones ambientales, con la capacidad que anidan en el Archipiélago Cubano (de limitativa ambiental (K; MacArthur 1972) de acuerdo con Garrido & García 1975, Garrido un área determinada. Esta última es: 1) poliva- & Kirkconnell 2000), así como las extinguidas lente cuando su concordancia es alta y se en aquél durante el cuaternario. Estas especies manifiesta en la frecuencia de los procesos (y subespecies) derivan de formas ancestrales cladogenéticos y en el mantenimiento de la que existieron en un espacio geográfico ame- homeostasis del desarrollo de las especies que 442 RELACIONES BIOGEOGRÀFICAS DE LA AVIFAUNA CUBANA la habitan; 2) mesovalente, si la concordancia determinación de los grupos corogénicos. El disminuye y el potencial genético es disconti- material bibliográfico examinado fue: Peter- nuo, lo que se refleja igualmente en las meno- son (1947), Blake (1953), Mayr & Short res tasas de cambio evolutivo sobre las (1970), Garrido & García (1975), Ridgeley combinaciones de caracteres y en la amplitud (1976) y Meyer de Schauensee & Phelps de los nichos ecológicos de sus especies para (1978). el logro del mantenimiento de la homeostasis Las relaciones con otras Antillas resultan del desarrollo de las mismas (expresado en su del compartimiento de especies orníticas gran potencial estasigenético; Vergara 1992); y entre ambos. Las últimas se dividan en cuatro 3) oligovalente, si ocurre una ruptura de la entidades, de acuerdo con sus propiedades cli- concordancia, lo que se manifiesta en una alta mático-fisiográficas y valencias evolutivas: tasa de extinciones, en la frecuencia de relic- Islas Bahamas, Islas Caimán, Antillas Mayores tos, en la disminución del potencial genético y y Antillas Menores. Se examinó Bond (1936, en un máximo del potencial estasigenético de 1985), Johnston (1975), Garrido & García sus especies. (1975), Lack (1976) y Voous (1983). Grupos corogénicos (sensu Vergara 1988b). Se dis- Método computacional. Se utilizó un programa en tingue dos grupos: el grupo corogénico norte- FORTRAN, mediante el empleo del coefi- americano, y el grupo corogénico suda- ciente de correlación Ø. Se desarrolló un pro- mericano. cedimiento de taxonomía numérica como índice de similitud (cluster analysis, mediante Fauna semiarmónica. Aquella que, aunque diver- el método de unión completa “complete sificada, no incluye todos los tipos básicos linkage”; Sneath & Sokal 1973), para conocer adaptativos, presentes en otra (aquí el área pri- las asociaciones de las áreas consideradas y el mordial centroamericana), que ecológica- grado de proximidad de estas con la cubana, mente los tiene en mayor medida (De acuerdo sobre la base del compartimiento de las espe- con la definición de fauna armónica de cies no endémicas que anidan en el Archipié- MacArthur & Wilson 1967). Aquí se consi- lago cubano (120), que también están dera fundamentalmente el número de gremios presentes en aquellas. trófico-conductuales y el atestamiento de especies contenidas en estos últimos. ÁREA ESTUDIADA Índice de diversidad de especies (de
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