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FCEN-UBA | Groba, Hernán F.. 2014 03 14 Tesis Doctoral Caracterización morfológica de las larvas y de los receptores olfativos del parasitoide Mallophora ruficauda (Diptera: Asilidae) y determinación del origen de las claves químicas en el hospedador Cyclocephala signaticollis (Coleoptera: Scarabaeidae) Groba, Hernán F. 2014-03-14 Este documento forma parte de la colección de tesis doctorales y de maestría de la Biblioteca Central Dr. Luis Federico Leloir, disponible en digital.bl.fcen.uba.ar. Su utilización debe ser acompañada por la cita bibliográfica con reconocimiento de la fuente. This document is part of the doctoral theses collection of the Central Library Dr. Luis Federico Leloir, available in digital.bl.fcen.uba.ar. It should be used accompanied by the corresponding citation acknowledging the source. Cita tipo APA: Groba, Hernán F.. (2014-03-14). Caracterización morfológica de las larvas y de los receptores olfativos del parasitoide Mallophora ruficauda (Diptera: Asilidae) y determinación del origen de las claves químicas en el hospedador Cyclocephala signaticollis (Coleoptera: Scarabaeidae). Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Cita tipo Chicago: Groba, Hernán F.. "Caracterización morfológica de las larvas y de los receptores olfativos del parasitoide Mallophora ruficauda (Diptera: Asilidae) y determinación del origen de las claves químicas en el hospedador Cyclocephala signaticollis (Coleoptera: Scarabaeidae)". Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. 2014-03-14. Dirección: Biblioteca Central Dr. Luis F. Leloir, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. Contacto: [email protected] Intendente Güiraldes 2160 - C1428EGA - Tel. (++54 +11) 4789-9293 UNIVERSIDAD DE BUENOS AIRES Facultad de Ciencias Exactas y Naturales Departamento de Ecología, Genética y Evolución Caracterización morfológica de las larvas y de los receptores olfativos del parasitoide Mallophora ruficauda (Diptera: Asilidae) y determinación del origen de las claves químicas en el hospedador Cyclocephala signaticollis (Coleoptera: Scarabaeidae) Tesis presentada para optar al título de Doctor de la Universidad de Buenos Aires en el área de CIENCIAS BIOLÓGICAS Hernán F. Groba Director de Tesis: Dra. Marcela K. Castelo Consejeros de estudios: Dra. Lilia S. Lauria de Cidre y Dr. Pablo E. Schilman Lugar de trabajo: Grupo de Investigación en Ecofisiología de Parasitoides Buenos Aires, Marzo 2014 RESUMEN Caracterización morfológica de las larvas y de los receptores olfativos del parasitoide Mallophora ruficauda (Diptera: Asilidae) y determinación del origen de las claves químicas en el hospedador Cyclocephala signaticollis (Coleoptera: Scarabaeidae) Autor: Lic. Hernán F. Groba Director: Dra. Marcela K. Castelo En un ambiente complejo y heterogéneo, el éxito de los individuos y de su descendencia depende de la capacidad de localizar los recursos y las condiciones necesarias para su supervivencia. El encuentro de los recursos y la evaluación de las condiciones se logran a través de la percepción de estímulos o claves, y particularmente en los insectos, los estímulos químicos juegan un rol central en estos procesos. En los parasitoides, la búsqueda y localización de los hospedadores, los cuales constituyen el recurso fundamental en su vida, puede involucrar diferentes tipos de estímulos, pero en particular son las claves químicas volátiles las que están mayormente implicadas en este comportamiento. El moscardón cazador de abejas Mallophora ruficauda es un parasitoide solitario de larvas de coleópteros de la familia Scarabaeidae (gusanos blancos del suelo) y una plaga de la apicultura de la Región Pampeana Argentina. En estado adulto es depredador de insectos, en particular de abejas melíferas, y en estado larval parasita preferentemente a larvas de tercer estadio de Cyclocephala signaticollis. La localización del hospedador por M. ruficauda está compartida entre la hembra y la larva. En primer lugar, la hembra localiza el hábitat del hospedador y ovipone en sustratos altos maximizando la dispersión de su progenie por el viento. En una segunda fase, las larvas que llegan al suelo se entierran, mudan y buscan activamente a su hospedador por medio de claves químicas que se originan en el tubo digestivo del gusano blanco. Hasta el momento, existen algunos aspectos de este mecanismo de localización que se desconocen. Por tanto, en la presente Tesis se estudiaron aspectos asociados al comportamiento de localización del hospedador por parte de M. ruficauda, tales como las características morfológicas de las estructuras sensoriales de los adultos y las larvas, los aspectos fisiológicos y morfológicos de las larvas asociados al período de búsqueda, y el rol ecológico y origen de las claves químicas I producidas por los hospedadores. Se encontró que las antenas de los adultos de M. ruficauda están especializadas en la captura e intercepción de estímulos químicos volátiles, posiblemente involucrados en los comportamientos de apareamiento y oviposición. Las larvas de primer y segundo estadio del parasitoide poseen una clara diferencia morfológica en sus estructuras cefálicas, lo que podría estar vinculado al comportamiento diferencial observado entre estos estadios durante el proceso de búsqueda del hospedador. Además el estudio del tamaño corporal de las larvas de primer estadio que nacen a lo largo de la temporada reproductiva sugiere que habría una asignación diferencial de recursos energéticos por parte de las hembras, pero no se ha encontrado un patrón claro al respecto. Por último, se determinó que tanto las larvas de estadio II de M. ruficauda como las de estadio III de C. signaticollis se orientan positivamente hacia extractos de la cámara de fermentación de las larvas de estadio III de C. signaticollis. Estos resultados sugieren que las larvas de M. ruficauda podrían estar explotando el sistema de comunicación química de su hospedador durante la búsqueda y localización. Se determinó indirectamente el origen de estas claves, observándose la presencia de microorganismos asociados al hospedador en esta región del tubo digestivo. Mediante experimentos comportamentales se demostró que los productos metabólicos de estos microorganismos serían las claves que orientan a las larvas del parasitoide durante la localización del hospedador. Sin embargo, no se determinó la función de estas sustancias químicas en las interacciones intraespecíficas de C. signaticollis. En conclusión, M. ruficauda presenta estructuras sensoriales especializadas vinculadas a la percepción de estímulos químicos volátiles, aspectos fisiológicos que permiten la sobrevida de las larvas hasta el encuentro con el hospedador, y el uso de claves altamente confiables y específicas durante la búsqueda, las cuales contribuyen a la eficiencia en la localización de recursos por larvas y hembras de M. ruficauda. Esta Tesis es una contribución al conocimiento de la historia de vida de M. ruficauda y un aporte al entendimiento de otras especies de parasitoides con características y estrategias de vida similares. PALABRAS CLAVE: Mallophora ruficauda, Scarabaeidae, búsqueda del hospedador, sensilias, claves químicas. II ABSTRACT Morphological characterization of larvae and olfactive receptors in the parasitoid Mallophora ruficauda (Diptera: Asilidae) and determination of chemical cues origin in the host Cyclocephala signaticollis (Coleoptera: Scarabaeidae) Author: Lic. Hernán F. Groba Supervisor: Dra. Marcela K. Castelo Individuals and their offspring success depend on their ability to locate the resources and optimal conditions required for survival in a complex and heterogeneous environment. The finding of resources and evaluation of these conditions is given through stimuli or cues. In most animals, especially insects, chemical cues play a central role in the location of resources. Parasitoid’s hosts are a fundamental resource on parasitoid life history, as without those parasitoids are not able to develop its life cycle. Host-seeking behaviour may include different types of cues or stimuli but particularly volatile chemicals are mainly involved. The robber fly Mallophora ruficauda is a solitary ectoparasitoid of scarab beetle larvae (white grubs) and a major pest of beekeeping in the Pampas Region of Argentina. Adults are predators of flying insects, especially honeybees, and larvae parasitized mainly third instar larvae of Cyclocephala signaticollis. Mallophora ruficauda have a split host location strategy between female and larva. First, female locates the host habitat and lay eggs in clusters on elevated available sites increasing larval dispersal and hence encounter success with larval hosts. Second, once in the ground, larva starts to dig and actively seeks for its host detecting chemical cues arisen from white grub’s digestive tube. So far, some characteristics of host finding mechanism are still unknown. Therefore, in this Thesis some features associated to M. ruficauda host searching behaviour, such as morphological characteristics of sensory structures of adults and larvae, larvae morphological and physiological aspects associated to searching process, and ecological role and origin of chemical cues produced by hosts were studied. Antennae of M. ruficauda adults are specialized for interception and capture of volatile chemical stimuli, possibly involved in mating and oviposition behaviour. First and second parasitoid larval instars showed morphological differences in cephalic structures which could be associated with the differential behaviour
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