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Tesis De Doctorado TESIS DE DOCTORADO: DIVERSIDAD, DISTRIBUCIÓN TEMPORAL Y TRÓFICA, Y PATRONES DE NIDIFICACIÓN DE UN AGREGADO DE ESPECIES DE ESCARABEIDOS COPRÓFAGOS (COLEOPTERA) EN UN CAMPO NATURAL PASTOREADO (CERRO COLORADO, DPTO. DE FLORIDA, Uruguay) Tesista: Enrique Morelli Área: Biología Subárea: Zoología Orientador: Dr. Fernando Pérez-Miles. Sección Entomología, Facultad de Ciencias. Coorientador: Dr. Arturo Baz-Ramos. Dpto. de Biología Animal, Facultad de Alcalá de Henares, España. Laboratorio de Ejecución: Sección Entomología, Facultad de Ciencias. Montevideo 2005 DIVERSIDAD, DISTRIBUCIÓN TEMPORAL Y TRÓFICA, Y PATRONES DE NIDIFICACIÓN DE UN AGREGADO DE ESPECIES DE ESCARABEIDOS COPRÓFAGOS (COLEOPTERA) EN UN CAMPO NATURAL PASTOREADO (CERRO COLORADO, DPTO. DE FLORIDA, URUGUAY). Sección I: Capítulo 1- Introducción, objetivos y estructura de la Tesis Capítulo 2- Superfamilia Scarabaeoidea Latreille, 1802 Capítulo 3- Ecología de los Scarabaeoidea coprófagos Capítulo 4- Preferencias tróficas de los Scarabaeoidea coprófagos Capítulo 5- El estiércol como recurso Capítulo 6- Los Scarabaeoidea coprófagos y los ecosistemas de pastoreo Capítulo 7- Antecedentes Johannes Macarius, Abraxas y Apistopistus, Amberes, 1657 Es el escarabajo unigénito autor de sí mismo, compuesto de mortal especie hecha de sórdida masa, sin ayuntamiento venéreo, y formador de la figura del mundo. Este que se engendra y vive entre la inmundicia hedionda, muere con el olor suave de la rosa; y así pintado por sí solo, es símbolo y jeroglífico del hombre valeroso y valiente que huye de los deleites y regaladas blanduras, ocupado en continuo trabajo y por esto los traían los soldados esculpidos en los anillos y por divisa los capitanes romanos....... Jerónimo de Huerta, Hist. Nat. X, XXVIII-anotación ÍNDICE Resumen.............................................................................................................. 1 SECCIÓN I: CAPÍTULO 1 Introducción.......................................................................................................... 3 CAPÍTULO 2 Superfamilia Scarabaeoidea Latreille, 1802......................................................... 9 CAPÍTULO 3 Ecología de los Scarabaeoidea coprófagos.......................................................... 21 CAPÍTULO 4 Preferencias tróficas de los Scarabaeoidea coprófagos........................................ 65 CAPÍTULO 5 El estiércol como recurso................................................................................... 71 CAPÍTULO 6 Los Scarabaeoidea coprófagos y los ecosistemas de pastoreo............................ 88 CAPÍTULO 7 Antecedentes........................................................................................................ 99 SECCIÓN II: CAPÍTULO 8 Material y métodos.............................................................................................. 104 CAPÍTULO 9 Taxocenosis......................................................................................................... 125 CAPÍTULO 10 Fenología .......................................................................................................... 148 CAPÍTULO 11 Selección del recurso.......................................................................................... 166 CAPÍTULO 12 Ciclos de vida Patrones de nidificación..................................................................................... 213 CAPÍTULO 13 Conclusiones finales......................................................................................... 238 CAPÍTULO 14 Bibliografía....................................................................................................... 266 Apéndice: TABLAS........................................................................................ 24 TESIS DE DOCTORADO: DIVERSIDAD, DISTRIBUCIÓN TEMPORAL Y TRÓFICA, Y PATRONES DE NIDIFICACIÓN DE UN AGREGADO DE ESPECIES DE ESCARABEIDOS COPRÓFAGOS (COLEOPTERA) EN UN CAMPO NATURAL PASTOREADO (CERRO COLORADO, DPTO. DE FLORIDA) Tesista: Enrique Morelli Área: Biología Subárea: Zoología Orientador: Dr. Fernando Pérez-Miles. Sección Entomología, Facultad de Ciencias. Coorientador: Dr. Arturo Baz-Ramos. Dpto. de Biología Animal, Facultad de Alcalá de Henares, España. Laboratorio de Ejecución: Sección Entomología, Facultad de Ciencias. Fecha de aprobación: 27/IV/2000 RESUMEN Un fenómeno importante dentro del ecosistema edáfico es la coprofagia. Muchos organismos detritívoros son coprófagos, ingiriendo regularmente estiércol después de que éste ha sido enriquecido y modificado por la actividad microbiana del medio. Los estudios sobre comunidades coprófagas tienen gran interés tanto desde el punto de vista teórico como aplicado. Cada excremento de bóvido constituye un microhábitat efímero, delimitado y heterogéneo con respecto al medio en el que se produce un relevo faunístico característico, a medida que éste se homogeiniza consustrato. En esta microsucesión, los Coleoptera Scarabaeoidea coprófagos, representan el mayor número de taxa de insectos. Los escarabajos coprófagos participan activamente en el proceso de descomposición sobre las heces de los herbívoros, contribuyendo a mantener el flujo de nutrientes en el sistema suelo-planta-animal, disminuyendo la pérdida de superficie para pastoreo, favoreciendo la fertilidad , textura, aereación y permeabilidad al agua de los suelos, así como la reducción de poblaciones de parásitos que se desarrollan en el estiércol. La composición y estructura de las comunidades de Scarabaeoidea coprófagos estarían determinadas por el reparto de recursos, permitiendo la coexistencia de especies potencialmente competidoras y los patrones de colonización y distribución entre parches de recurso. En el presente estudio se plantean los siguientes objetivos: • Determinar la riqueza y abundancia de coleópteros Scarabeoidea coprófagos asociados a excrementos bovino y ovino en un predio de campo natural sometido a pastoreo. • Establecer las fenologías y preferencias tróficas de las distintas especies. • Describir para el mayor número de especies posible, los ciclos de vida y los patrones de nidificación. CAPÍTULO 1 INTRODUCCIÓN Los excrementos al igual que otros tipos de recursos como los cadáveres, frutos y madera muerta, presentan una serie de características especiales que afectan a la dinámica y estructura del conjunto de especies que los utilizan. Por un lado, constituyen unidades o parches de alta calidad a lo que se ha atribuido el hecho que alberguen una elevada diversidad de especies (Atkinson & Shorrocks, 1981). Pero por otro lado, su duración en el tiempo es muy reducida, de forma que las especies solo pueden completar una generación dentro de ellos (Hanski, 1991b). Los mamíferos, y en menor grado, las aves, los reptiles y algunos moluscos terrestres, eliminan al final de su ciclo digestivo, masas más o menos importantes de excrementos, en general de consistencia pastosa. El volumen de excremento producido por un individuo a lo largo de su vida, representa una masa infinitamente superior al peso corporal del animal. Estas deyecciones contienen, conjuntamente con aquellos elementos orgánicos que han escapado a la digestión, una parte de los elementos intermediarios de esta digestión que no han sido absorbidos durante el tránsito intestinal, y una rica flora bacteriana y fúngica que se ha desarrollado en el curso de las diversas fases de la digestión y de su tránsito intestinal. Las deyecciones ofrecen así, una reserva potencial de moléculas orgánicas más o menos complejas, y una reserva energética, que hacen pensar en el excremento como un micro-ecosistema particular. El excremento como recurso, presenta algunas características que le son propias, y otras que comparte con otros micro-sistemas como cadáveres y frutos (Paulian, 1993). Su característica más sobresaliente es la tendencia a una distribución espacial y temporal fuertemente discontinua. La distribución espacial no es totalmente aleatoria. Los excrementos son depuestos, aunque se trate de especies que estén en continuo movimiento, en lugares con cierta preferencia próximo a los abrevaderos, bajo árboles, refugios, etc. Otras especies, de hábitos sedentarios, defecan en un lugar establecidos dentro de sus madrigueras. Inclusive, cuando se constituyan depósitos relativamente importantes, éstos quedan alejados unos de otros, y responden a una distribución siempre ampliamente discontinua. Por otra parte, aunque las concentraciones de excremento sean importantes, y haciendo la salvedad de aquéllos que son depuestos bajo tierra, el sol deseca rápidamente estas masas o son dispersadas por la lluvia. De esta forma, este recurso presenta un periodo limitado en el tiempo para ser utilizado. En la región mediterránea, por ejemplo, una placa de excremento vacuno expuesto al sol en una pradera, pierde todo poder atractivo en 36 horas. En regiones intertropicales, el periodo es aún más breve. Se ha observado que los agregados de especies que utilizan recursos parcheados muestran una elevada variación en el número de individuos que alberga cada parche (excrementos: Hanski, 1987a; Holter, 1982; cadáveres: Kneidel, 1985; Hanski, 1987a; frutos: Atkinson & Shorrocks, 1984). Esta distribución agregada ha sido interpretada 3 como un mecanismo que permite la coexistencia de especies potencialmente competidoras (Atkinson & Shorrocks, 1981; Hanski, 1981, 1987a; Ives, 1988). Las heces son consumidas primordialmente por toda una comunidad de insectos coprófagos, entre los que se destacan los Scarabaeoidea coprófagos. Su actividad acelera la desaparición de las
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