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Estudio De Los Agregados De Dípteros Sarcosaprófagos Y Su Relación Con Los Ecosistemas Naturales De La Comunidad De Madrid

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Estudio De Los Agregados De Dípteros Sarcosaprófagos Y Su Relación Con Los Ecosistemas Naturales De La Comunidad De Madrid DEPARTAMENTO DE ZOOLOGÍA Y ANTROPOLOGÍA FÍSICA ESTUDIO DE LOS AGREGADOS DE DÍPTEROS SARCOSAPRÓFAGOS Y SU RELACIÓN CON LOS ECOSISTEMAS NATURALES DE LA COMUNIDAD DE MADRID Memoria presentada por Daniel Martín Vega, bajo la dirección del Dr. Arturo Baz Ramos, para optar al grado de Doctor Alcalá de Henares, 2011 DEPTO. DE ZOOLOGÍA Y ANTROPOLOGÍA FÍSICA Edificio de Ciencias. Campus Universitario 28871 Alcalá de Henares (Madrid) Tel.: 91 885 49 22 / Fax: 91 885 50 80 DON ARTURO BAZ RAMOS, PROFESOR TITULAR DEL DE- PARTAMENTO DE ZOOLOGÍA Y ANTROPOLOGÍA FÍSICA DE LA UNIVERSIDAD DE ALCALÁ, HACE CONSTAR: Que la presente memoria titulada “Estudio de los agregados de dípteros sarcosaprófagos y su relación con los ecosistemas natura- les de la Comunidad de Madrid”, presentada por Daniel Martín Vega para optar al grado de Doctor, ha sido realizada bajo mi dirección cumpliendo las condiciones exigidas para su presentación. Para que así conste a los efectos oportunos, firmo la presente en Alcalá de Henares, a 19 de mayo de 2011. Fdo.: Dr. Arturo Baz Ramos DEPTO. DE ZOOLOGÍA Y ANTROPOLOGÍA FÍSICA Edificio de Ciencias. Campus Universitario 28871 Alcalá de Henares (Madrid) Tel.: 91 885 49 22 / Fax: 91 885 50 80 El Dr. Jacinto Gamo García, Director del Departamento de Zoología y Antropología Física de la Universidad de Alcalá, en re- lación con la Tesis Doctoral “Estudio de los agregados de dípteros sarcosaprófagos y su relación con los ecosistemas naturales de la Co- munidad de Madrid”, de la que es autor Daniel Martín Vega, y que ha sido dirigida por el Dr. Arturo Baz Ramos, profesor titular del Departamento de Zoología y Antropología Física de la Universidad de Alcalá, CONFIRMA que dicha Tesis reúne los requisitos necesarios para su presentación y defensa. Para que así conste, firmo la presente en Alcalá de Henares, a 19 de mayo de 2011, con el VºBº del Tutor, Dr. Arturo Baz Ramos Fdo.: Dr. Jacinto Gamo García VºBº: Dr. Arturo Baz Ramos El presente trabajo ha sido financiado por: - Ministerio de Ciencia y Tecnología (Proyecto de Investigación BOS2003-00400) - Universidad de Alcalá (Proyecto de Investigación PI2003/016) Durante la elaboración de este trabajo, Daniel Martín Vega ha disfrutado de las siguientes becas: - Programa FPI (Formación de Personal Investigador) de la Univer- sidad de Alcalá - Programa FPU (Formación de Profesorado Universitario) del Min- isterio de Educación y Ciencia - Beca asociada al Proyecto de Investigación IUICP/PI2010/001 del IUICP (Instituto Universitario de Investigación en Ciencias Policia- les) de la Universidad de Alcalá ESTUDIO DE LOS AGREGADOS DE DÍPTEROS SARCOSAPRÓFAGOS Y SU RELACIÓN CON LOS ECOSISTEMAS NATURALES DE LA COMUNIDAD DE MADRID A mis padres. “Porque si el sol engendra gusanos en un perro muerto, besando la carroña, siendo un dios...” William Shakespeare, “Hamlet”. 15 Agradecimientos ebo comenzar expresando mi agradecimiento al Dr. Artu- ro Baz, director de esta tesis doctoral (con todo lo que ello implica) y responsable de todos los resultados positivos que pueda contener. Asimismo, debo dar también espe- Dcialmente las gracias a las Drs. Luisa Díaz Aranda y Blanca Cifrián, que me han ayudado en distintos momentos y aspectos de todo este tiempo como doctorando. El agradecimiento se extiende a todo el Departamento de Zoología y Antropología Física de la Universidad de Alcalá, con mención especial para los compañeros de tercer ciclo que han pasado por allí y con los que he compartido muchas, muchas horas: Aída, Alfonso, Carolina, Chusa, Cristina, Irene, Laura, Noemí, Silvia y Txomin. Quisiera expresar también mi agradecimiento al Dr. Verner Michelsen, de la Universidad de Copenhague, por su valiosa colabo- ración y su generosidad en el trabajo sobre el hallazgo de la especie Thyreophora cynophila. Igualmente me encuentro en deuda con to- dos aquellos entomólogos que identificaron parte del material inven- tariado en este estudio y cuyos nombres se especifican en el capítulo 2 de esta memoria, así como con todos aquellos investigadores que han atendido algunas de mis dudas y consultas. Por supuesto, debo dar también las gracias al personal del servicio de préstamo interbi- bliotecario de la Universidad de Alcalá, por satisfacer todas mis peti- ciones bibliográficas, algunas de ellas realmente complicadas. Con la ayuda de todos ellos este trabajo ha podido llegar a mejor término. Querría aprovechar además para dar también las gracias a los Drs. 16 Los dípteros sarcosaprófagos en la Comunidad de Madrid Francisco Ponce Gordo y Rafael Martínez Díaz, de la Universidad Complutense de Madrid y la Universidad Autónoma de Madrid, res- pectivamente, por permitirme colaborar en sus proyectos y aprender con ellos cuando era un recién licenciado con muchas dudas sobre hacia dónde dirigirme. Por último, no puedo dejar de dar las gracias a todos los fa- miliares y amigos que, aunque muchas veces no entiendan muy bien qué hago estudiando “los bichos que se comen a los muertos”, me han dado durante todo este tiempo el ánimo necesario para evitar que me quedase a medio camino. Como mis padres y mi hermano. O mis abuelas y abuelos, tías y tíos, primas y primos. O Pilar Menéndez. O Mariluz Mateos. O Álvaro Barranco, o los dos Albertos (Pérez y Carrasco), o Pablo Malavé, o Raúl Diego, o Javier Cardenete, o Carlos Domínguez, o Javier Cacho, o José Juan Gallardo, o Paloma Pérez, o Pablo G. Cuadrado, o Susana Sánchez. O muchas más per- sonas que harían esto interminable. A todos ellos, gracias. 17 Índice Resumen / Abstract. 21 CAPÍTULO 1. INTRODUCCIÓN. 25 1.1 La carroña como recurso. 26 1.2 Importancia y aplicaciones de los dípteros sarcosaprófagos 29 1.3 La importancia de los estudios faunísticos regionales. Co- nocimiento sobre los dípteros sarcosaprófagos en la Península Ibérica. 32 1.4 Justificación y objetivo del estudio. 35 CAPÍTULO 2. ÁREA DE ESTUDIO. METODOLOGÍA. 37 2.1 Descripción del área de estudio. 37 2.1.1 Variables topográficas y climáticas. Pisos biocli- máticos. 38 2.1.2 Variables edáficas. 41 2.1.3 Variables de uso del territorio. 43 2.2 Metodología. 44 2.2.1 Diseño de muestreo. Selección de localidades. 44 2.2.2 Validez de la selección de localidades. 50 2.2.3 Trampas. 60 2.2.4 Procesado e identificación de las muestras. 66 2.2.5 Análisis y tratamiento de los datos. 67 18 Los dípteros sarcosaprófagos en la Comunidad de Madrid CAPÍTULO 3. COMPOSICIÓN FAUNÍSTICA. 71 3.1 El inventario de especies. 71 3.1.1 Familia Calliphoridae. 77 3.1.2 Familia Dryomyzidae. 85 3.1.3 Familia Heleomyzidae. 86 3.1.4 Familia Muscidae. 91 3.1.5 Familia Piophilidae. 100 3.1.6 Familia Platystomatidae. 106 3.1.7 Familia Sarcophagidae. 106 3.1.8 Familia Scathophagidae. 117 3.1.9 Familia Sepsidae. 118 3.1.10 Familia Tachinidae. 119 3.1.11 Familia Ulidiidae. 124 3.2 Las familias no identificadas. 127 3.3 La importancia de inventariar la biodiversidad: El caso ex- cepcional de la familia Piophilidae. 129 3.4 La presencia de especies no necrófagas en el inventario. 138 3.5 Sex-ratio. 141 CAPÍTULO 4. RIQUEZA Y DIVERSIDAD DE ESPECIES. 149 4.1 Estimas de la riqueza de especies. 149 4.1.1 Curvas de acumulación de especies. 151 4.1.2 Estimadores no paramétricos. 158 4.1.3 Discusión sobre las diferentes estimas de riqueza de especies obtenidas. 164 4.2 Patrones de diversidad. 168 4.2.1 Índices de diversidad, dominancia y equitatividad 168 4.2.2 Influencia del hábitat en la diversidad de especies 186 CAPÍTULO 5. ESTACIONALIDAD Y FENOLOGÍA. 197 5.1 Variación estacional en el número de especies y ejemplares 198 5.1.1 Variaciones estacionales. 198 5.1.2 Variaciones estacionales en los diferentes pisos bioclimáticos y hábitats. 202 5.2 Rango y amplitud de nicho estacional. 217 5.2.1 Rango estacional. 217 5.2.2 Amplitud de nicho estacional de las especies más significativas. 223 Índice 19 5.3 Fenología de las especies más representativas. 226 5.3.1 Familia Calliphoridae. 231 5.3.2 Familia Muscidae. 242 5.3.3 Familia Heleomyzidae. 251 5.3.4 Familia Sarcophagidae. 259 5.3.5 Familia Piophilidae. 264 5.3.6 Familia Ulidiidae. 271 CAPÍTULO 6. LOS AGREGADOS DE DÍPTEROS SARCO- SAPRÓFAGOS. 275 6.1 Variación espacial de los agregados de especies de dípteros sarcosaprófagos y preferencias de hábitat de las principales es- pecies. 276 6.2 Variación espacio-temporal de los agregados de especies de dípteros sarcosaprófagos. 306 6.2.1 Verano. 306 6.2.2 Otoño. 315 6.2.3 Invierno. 323 6.2.4 Primavera. 328 Conclusiones. 337 Referencias. 343 ANEXO. ARTÍCULOS PUBLICADOS. 389 Artículo publicado en Entomologia Generalis. 391 Artículo publicado en Boletín de la Asociación española de Entomología. 401 Artículo publicado en Systematic Entomology. 405 Artículo publicado en European Journal of Entomology. 415 Artículo publicado en Medical and Veterinary Entomology. 419 20 Los dípteros sarcosaprófagos en la Comunidad de Madrid 21 Resumen l orden Diptera constituye un grupo de insectos especial- mente diverso y de gran importancia económica y sani- taria, aspectos que se ven acentuados en el caso de aque- llas especies que se desarrollan sobre materia orgánica en Edescomposición. Sin embargo, a pesar del interés y las aplicaciones de estos dípteros sarcosaprófagos, el conocimiento sobre su ecología en la Península Ibérica es aún muy limitado. Este trabajo presenta un estudio sobre los agregados de dípteros sarcosaprófagos en los siete ecosistemas naturales más característicos de la Comunidad de Madrid como región representativa del centro peninsular. Un total de 22202 ejemplares pertenecientes a 11 familias y a 87 especies de dípteros sarcosaprófagos fueron capturados mediante el uso de trampas cebadas con carroña en 21 localidades entre junio de 2006 y mayo de 2007.
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