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(Hymenoptera: Scelionidae), a Statistical Approach

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(Hymenoptera: Scelionidae), a Statistical Approach UNIVERSIDAD DE COSTA RICA SISTEMA DE ESTUDIOS DE POSGRADO PATRONES DE COLORACIÓN EN HYMENOPTERA: SU ORIGEN, EVOLUCIÓN Y POSIBLE FUNCIÓN EN MIRAS A LA APLICACIÓN BIOMIMÉTICA Tesis sometida a la consideración de la Comisión del Programa de Doctorado en Ciencias para optar al grado y título de Doctorado Académico en Ciencias REBECA ANDREA MORA CASTRO Ciudad Universitaria Rodrigo Facio, Costa Rica 2020 Ilustración1. Jennifer Bejarano. Título: forrageo de microavispa. ii DEDICATORIA ACKNOWLEDGEMENTS El presente documento, resultado de cuatro años y medio de investigación, no habría sido posible sin el apoyo de seres diversos, dentro y fuera de la academia, de diferentes especies, con diferentes formas de pensar. Les presento toda mi gratitud. Esta tesis está dedicada a Coco, gracias por entender que mamá quería hacer esta última “cosa rara” antes de poder estar en casa un poco más de tiempo, todos esos proyectos que hemos estado planeando tocan ahora. Fernando, es completamente posible que en ciertos momentos me hubiera ahogado en un mar de ansiedad si no hubieras estado allí para acompañarme y reír junto a mí, gracias por tantos momentos de calma y claridad y por ser una parte tan importante de mi vida y mi conciencia. Gracias a algunos hombres y mujeres de la Academia que en un inicio me desanimaron, diciendo que la ciencia no era una buena carrera para una mujer, madre sola con una hija, con solo medio tiempo y otro trabajo externo a la Universidad, me ayudaron e impulsaron, a perseverar aún más. A la banda Pulp y el regalo de la música en general por "consejos e inspiración continuos" durante mis momentos de mayor cansancio, a mis 4 perros por sus incondicionales buenos días y compañía. A mi jardín, huerta-tierra viva y visitas al bosque y a mi grupo de yoga de la Universidad Nacional, gracias por ser mi meditación diaria. Gracias a mis padres, Olga y Johnny y familia cercana, en especial a Adrián y Jessica, mi comunidad de queridos amigos, Carlos Cartín y familia, María Laura Montero, Adriana Pacheco, Ana de la Guardia, Anna Klochklov, Floria Mora, Al Uy, César Rodríguez, Ximena Miranda, familia Cordero Espinoza, por todo el apoyo, por no querer a ratos hablar conmigo de ciencia y por las increíbles oportunidades que me han brindado a lo largo de los años. Mi tutor, Paul Hanson, sin su apoyo y aliento, ¡estaría en apuros! Gran mentoría, siempre accesible y cálido. Nuestras discusiones académicas estuvieron llenas de calma y apertura, gracias por respetar y acuerpar cada una de mis propuestas y gracias por su propuesta de mi pasantía doctoral llena de insectos, osos, nieve y mucha naturaleza, junto a mi hija, en la maravillosa Canadá. Por tanto, me gustaría agradecer a todos los investigadores, y principalmente a mi asesor, Lubo Masner del Canadian National Collection of Insects, Arachnids and Nematodes (CNC) en el Ottawa Research and Development Centre en Ottawa, por su maravillosa colaboración para poder navegar en la vasta e increíble colección de Hymenoptera, insumo para uno de los artículos científicos de esta investigación. Inmensas gracias a mí comité y mis lectores, Paul Hanson, Marcela Hernández y Mauricio Fernández. Tuve el privilegio tener un comité muy activo con numerosas reuniones las cuales trajeron cada vez muchos comentarios y nuevas ideas que impulsaron la investigación. Deseo que todos los estudiantes de doctorado tengan un comité tan activo. Asimismo, agradezco a mis colaboradores del grupo del Centro de Investigación en Ciencia e Ingeniería de Materiales (CICIMA) en especial a Marcela Hernández, Esteban Avendaño y Marcela Alfaro. Esta tesis consistió de un trabajo arduo interdisciplinar con un grupo multidisciplinario, por tanto, agradezco a los tres por la habilidad de construir una estrategia de comunicación alineada con todos los investigadores. Cada disciplina de investigación tiene un "lenguaje" único, la jerga y los términos de uso común deben estar claramente definidos y los iii colaboradores que son relativamente nuevos en su campo pueden requerir explicaciones más detalladas. Gracias por la paciencia para explicar nuevos conceptos, por trabajar en sinergia, complementándonos para completar exitosamente el conjunto de herramientas esenciales y necesarias para lograr el objetivo final. Gracias Esteban Avendaño y Marcela Hernández por su visión de “ciencia abierta para todos” / “open floor science”, gracias por ofrecer el equipo del CICIMA y por garantizar un debido entrenamiento sin que los cobros excesivos opaquen el sentido de por qué hacemos ciencia. Me siento muy agradecida de poder incorporarme a un excelente grupo de investigación científica acuerpado por personas con calidad humana. Thanks to my external collaborators, especially the Biomimicry Institute and the Copenhagen Institute of Interactive Design, although we are from different countries we are connected through lasting intellectual ties, thanks for self-organizing through common interests and not through top-up/below partnerships and for transcending political, religious and linguistic barriers. I am really grateful of joining forces with you and for your new approaches and different types of expertise needed to renew science, thanks for educating me on inter- and trans-disciplinarily research and for the creation of more spaces for science to engage with different publics and vice-versa. A todos los estudiantes que colaboraron en este proyecto Alejandra Sánchez, Steven Zamora, José Vargas, Soren Pessoa, Edgar Pérez, Jennifer Bejarano, Andrés Pintor, Anthony Ulate, Geovanna Rojas, Paulina Morales y Natalia Jiménez gracias y mantengan ese maravilloso entusiasmo siempre y a mis estudiantes en docencia, gracias por permitirme hacer algo que amo, enseñar, aprender y compartir. Gracias al personal de Protolab y Proinnova UCR por apoyar siempre mis ideas, sin censura, dando como resultado la patente de la jaula automatizada programable que facilitó este trabajo doctoral. Por último, gracias al Programa de Doctorado en Ciencias y a la Universidad de Costa Rica, a esta tesis y mis modelos de estudio, artrópodos e insectos, por darme conocimiento, felicidad y sorpresas en el campo y por reforzar lo que para mí significa hacer ciencia. Aunque a ratos no lo parezca, afortunadamente, la profesión se adapta a muchas personas diferentes, las cuales pueden encontrar su propio nicho, en el que el "éxito" no debe definirse por la longitud de un CV, factores de impacto de la revista o el "reconocimiento de nombre" de la institución. Para mí eso no necesariamente equivale a “felicidad en la ciencia”, yo creo en otra métrica de éxito, una que puede lograr cualquier persona, con buen corazón, mucha curiosidad, motivación e interés social. “El mundo necesita de la ciencia y la ciencia necesita de mujeres” Bokova, directora Unesco. iv Índice de contenidos Páginas Preliminares Portada i Dedicatoria………………………………………………………………………………. .iii Hoja de aprobación……………………………………………………………………… …v Tabla de contenido……………………………………………………………………… …vi Resumen…………………………………………………………………………………. .vii Lista de Tablas…………………………………………………………………………… ..xi Lista de Ilustraciones……………………………………………………………………… .xii Cuerpo de Trabajo Introducción 1 Justificación 12 Capítulo 1 15 Capítulo 2 17 Capítulo 3 19 Capítulo 4 21 Capítulo 5 23 Discusión y conclusiones 26 Anexos 39 Referencias 40 vi RESUMEN El presente estudio proporciona la primera caracterización física y funcional del patrón negro- naranja-negro (BOB) que se encuentra en avispas parasitoides de muy pequeño tamaño (3-10 mm). Aunque se había reportado previamente que la coloración BOB ocurre en himenópteros, especialmente en Scelioninae, los resultados de esta investigación demuestran que este patrón está mucho más extendido de lo que se creía. Este patrón abarca tanto ecto- como endoparasitoides, idiobiontes y koinobiontes, de una diversidad de hospederos, así como también sínfitos fitófagos. Con respecto al tamaño, la coloración BOB parece ser especialmente común (no exclusiva) en himenópteros con una longitud corporal entre 3 y 10 mm. Con respecto a la distribución altitudinal de la coloración BOB, la gran mayoría de especímenes que muestran este patrón de color fueron recolectados debajo de los 2,000 msnm. Finalmente, este patrón de color se encuentra en todas las regiones biogeográficas, aunque con menor frecuencia que en el neotrópico. Para ahondar más en la caracterización de este patrón más allá de la taxonomía y hábitos biológicos, se utilizó la técnica óptica de la microspectrofotometría para generar espectros de reflectancia para ocho géneros de sceliónidos. Estos datos de reflectancia se analizaron mediante análisis univariados y análisis de datos funcionales los cuales demostraron ser superiores al ofrecer una mejor representación de la información física, dado que tienen en cuenta la curva espectrofotométrica completa. Adicionalmente, se implementó el método de componentes espectrales RGB para estudiar el origen del color en los casos en que no es posible dilucidar una composición química directa. En el caso de los géneros estudiados, se descubrió que los componentes espectrales azules del color naranja y negro eran casi idénticos, lo que sugiere que existe un compuesto común para los pigmentos. Se utilizó una correlación entre los valores medios de las características de los componentes de color en un intento de agrupar géneros que muestran valores similares y algunos, pero no todos, son similares a sus posiciones en las filogenias publicadas en la literatura. Profundizando más en la investigación, se propuso realizar
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