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Saissetia Oleae (Hemiptera: Coccidae) on Citrus and Olives: Population Structure and Biological Control

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Saissetia Oleae (Hemiptera: Coccidae) on Citrus and Olives: Population Structure and Biological Control UNIVERSIDAD POLITÉCNICA DE VALENCIA Escuela Técnica Superior de Ingenieros Agrónomos Departamento de Ecosistemas Agroforestales Black scale Saissetia oleae (Hemiptera: Coccidae) on citrus and olives: population structure and biological control DOCTORAL THESIS Presented by: Alejandro Tena Barreda Directed by: Dr. Ferran Garcia Marí Valencia 2007 DEPARTAMENTO DE ECOSISTEMAS AGROFORESTALES E.T.S. INGENIEROS AGRÓNOMOS Universidad Politécnica Valencia Camino de Vera, s/n Telf.96-3879260 -Fax96-3879269 46022 VALÈNCIA El Dr. Ferran Garcia Marí, Catedrático de la Universidad Politécnica de Valencia. CERTIFICA: Que la presente memoria titulada: “Black scale Saissetia oleae (Hemiptera: Coccidae) on citrus and olives: population structure and biological control”, realizado bajo mi dirección por D. Alejandro Tena Barreda, durante el periodo comprendido entre 2003 a 2007, constituye su Memoria de Tesis para optar al grado de Doctor, en el Departamento de Ecosistemas Agroforestales de la Universidad Politécnica de Valencia. Para que así conste a todos los efectos oportunos, firma el presente certificado Fdo: Dr. Ferran Garcia Marí Valencia, Mayo 2007 Agradecimientos Nombrar a cuantos han hecho posible la realización de esta memoria es cuanto menos una pretensión descabellada, así como injusto es pretender resumir en una simple frase toda su aportación. Así pués, y sabiendo de antemano que pecaré por omisión, quisiera expresar mi más sincero agradeciemiento al Dr. Ferran Garcia Marí, el meu mestre, per qui sent admiració i respecte, per haver- me donat la seua confiança i l’oportunitat de realitzar la tesis baix la seua supervisió, per haver-me guiat, aconsellat, dedicat el seu temps i sobretot per haver-me transmès la riquesa de la seua experiència respectant sempre les meues idees. to Dr. Robert F. Luck for giving me the opportunity of learning and working in his lab. For the long conversations about parasitoids and citrus pests. a la Dra. Antonia Soto porque sin su esfuerzo nada hubiera sido posible. Al Dr. Francisco Ferragut por su ayuda y por transmitirme su pasión por los “bichos”. A Carmen por su infinita paciencia y por sus consejos (y por las recetas), a Marta y Lupita por haberme introducido en la taxonomía y por haberme guiado cuando andaba muy perdido, a Miguel Angel por todo, a Tolis por descubrir el comportamiento de los parasitoides y hormigas juntos, a Cristina por transmitir mucha alegría (y organizar las comidas). A todos mis compañeros y amigos del Departamento, a los que siempre estuvieron ahí (Toni, Juanjo), a los que pasaron (Nabor, Juan, Laura, Pablo, María, Teresa,…) y a los recien llegados (Amparo, Alex, Amador, Raquel) por haberme ayudado siempre, por las risas, por los buenos momentos vividos y por haber aprendido juntos. to Dr. Apostolos Kapranas for transmitting all his knowledge about parasitoids, for teaching and guiding me in Riverside and for the drinks after work. To Porfirio Pacheco because “Metaphycus world” would not be possible without him, for teaching me English and for the “Mexican food”. To Bob’s lab people for helping me. To Dr. Gregory Walker for helping me with the montage and pictures of Metaphycus larvae. To Dr. Morse for his corrections in the second chapter and interesting conversations about thrips. To all my friends and colleagues in Riverside, Vicent and Aroa, Rory, Robbie, Rodrigo and Mariana, Diego, Jardel, Laura Simpson, Paul Flores, Laura,… for their friendship and support in the difficult moments. a mis compañeros y amigos de la ETSIA por todo lo vivido y por haberme ayudado en lo que hiciera falta. En especial a David y Juan por cuidarnos tanto. a les meues germanetes, a Vio y Carla, por haber crecido, aprendido y jugado juntos y sobretodo por todo el cariño y fuerza que me transmitis. a mis padres por habérmelo dado todo, por todos los esfuerzos que han tenido que realizar a lo largo de su vida y por haberme comprendido y apoyado en este camino. a Vero, por todo. A mi familia Table of contents Resumen .............................................................................................................. i Resum ................................................................................................................ iii Summary ............................................................................................................ v Chapter 1. Introduction .....................................................................................1 1.1- Systematic classification of Saissetia oleae 1.2. Importance of Saissetia oleae 1.2.1. Importance on citrus 1.2.2. Importance on olives 1.3. Biology of Saissetia oleae 1.3.1. Morphology 1.3.2. Life cycle, dispersal and migration 1.3.3. Seasonal history 1.3.4. Influence of abiotic factors 1.4. Damages 1.5. Biological control 1.5.1. Natural enemy complex 1.5.2. Augmentative releases 1.6. Metaphycus as parasitoids of soft scales 1.6.1. Host range and specificity 1.6.2. Immature development 1.6.3. Adult 1.7. Superparasitism 1.7.1. Host discrimination 1.7.2. Consequences of superparasitism 1.7.3. Ovicide Chapter 2. Justification and objectives ............................................................ 37 Chapter 3. Density and structure of Saissetia oleae populations on citrus and olives: relative importance of the two annual generations................................. 41 Chapter 4. Parasitoid complex of black scale Saissetia oleae on Citrus and Olives: seasonal trend and impact on host population. ..................................... 57 Chapter 5. Host discrimination, superparasitism and ovicide by Metaphycus flavus an endoparasitoid of Coccus hesperidum. ............................................... 79 Chapter 6. Conclusions .................................................................................. 109 Resumen La caparreta negra es la principal plaga de cítricos y olivos del género Coccidae en todo el Mundo. Su incidencia es especialmente importante en la cuenca del Mediterráneo donde ambos cultivos se encuentran ampliamente distribuidos y coexisten a nivel local. Diversos enemigos naturales, principalmente parasitoides, han sido introducidos con el fin de controlar las poblaciones del cóccido. Sin embargo, su incidencia parece ser limitada y la caparreta negra continua siendo una plaga ocasional en muchos países Europeos, incluido España. Para mejorar el control de esta especie bajo los principios del Manejo Integrado de Plagas en ambos cultivos hemos estudiado el número y la importancia de cada una de sus generaciones, sus parasitoides a lo largo de los año 2003-05, así como algunos aspectos de la biología del parasitoide Metaphycus flavus. Las poblaciones de caparreta negra mostraron una tendencia similar en ambos cultivos, presentando un máximo en julio, cuando las larvas móviles emergieron tras el periodo de puesta de las hembras adultas. A continuación se produjo una disminución poblacional debida a la alta mortalidad de las larvas de primera edad durante los meses de verano. Se observó una segunda salida de larvas móviles parcial, heterogénea y variable en otoño-invierno. Sin embargo, las poblaciones no aumentaron durante este periodo debido al efecto de las bajas temperaturas en la supervivencia de las larvas de primera edad y a la menor fertilidad de las hembras adultas, las cuales fueron la mitad de grandes y fecundas que las de primavera. No se encontraron diferencias entre el tamaño de las hembras adultas desarrolladas sobre cítricos y olivos. Si se realizan tratamientos para controlar posibles explosiones de la plaga recomendamos su aplicación a finales de julio, cuando las poblaciones son homogéneas, todas las larvas han emergido y las larvas de primera edad predominan en las poblaciones. Los parasitoides más abundante y ampliamente distribuidos de la caparreta negra en cítricos y olivos del este de España fueron Scutellista caerulea, Metaphycus flavus y Metaphycus lounsburyi. Scutellista caerulea fue encontrada bajo el 35.4 ± 7.5% y el 22.4 ± 3.5% del cuerpo de las hembras fijadas en cítricos y olivos respectivamente. Sin embargo, su eficacia parece limitada porque las poblaciones de los dos parasitoides aumentan demasiado tarde para poder prevenir explosiones poblacionales del cóccido. i Metaphycus flavus fue el parasitoide de estadios inmaduros más abundante. Metaphycus helvolus, introducido hace 30 años, no ha desplazado a M. flavus como en otras regiones del Mediterráneo. Metaphycus flavus es un endoparasitoide gregario facultativo que tiende a superparasitar cuando se cría sobre Coccus hesperidum, por lo que se llevó a cabo un estudio del comportamiento del parasitoide cuando este se encuentra con huéspedes previamente parasitados, así como las consecuencias de su comportamiento. Las hembras de M. flavus con y sin experiencia previa fueron capaces de distinguir entre los cóccidos sanos y parasitados, pero no pudieron distinguir entre aquellos parasitados por ellas mismas o por otras hembras de su misma especie. Las hembras no disminuyeron el tamaño de la puesta al superparasitar. Metaphycus flavus destruye los huevos previamente depositados en el cóccido antes de depositar los suyos, es decir, practica el ovicidio. Las hembras detectaron los huevos por la presencia del pedúnculo de estos, utilizando posteriormente el ovipositor para destruirlos. El número final de parasitoides que emergió de los huéspedes superparasitados se vió afectado por tres factores, encapsulación, competición entre larvas y ovicidio. El porcentaje de huevos encapsulados por el cóccido disminuyó desde alrededor
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