Pollination of Zamia Incognita A. Lindstr. & Idárraga on a Natural

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Pollination of Zamia Incognita A. Lindstr. & Idárraga on a Natural Pollination of Zamia incognita A. Lindstr. & Idárraga on a natural population in the Magdalena Medio, Colombia Trabajo de grado para el cumplimiento parcial de los requisitos para obtener el grado de Bióloga de la Universidad CES Estudiante: Wendy A. Valencia-Montoya Asesores: Dino Tuberquia y Juliana Cardona-Duque Programa de Biología Facultad de Ciencias y Biotecnología Universidad CES 2015 Al Magdalena Medio, para que la magia de sus bosques sobreviva a la indiferencia, y siga contando historias Agradecimientos Deseo agradecer a mis padres por todo el apoyo y aliento que desde muy temprano me dieron en este camino de estudiar formalmente los seres vivos. Especialmente a mi mamá Gloria Montoya, por su colaboración desde el procesamiento de datos hasta aportes económicos para el trabajo de campo; y principalmente porque es la síntesis de la capacidad de asombro, que para mí es el legado clave para estudiar biología. A mi papá Gedwin Valencia por su enorme apoyo y entusiasmo hablando de las zamias a todas las personas que conoce. A mis asesores Dino Tuberquia y Juliana Cardona porque además de todo el apoyo académico durante estos años han sido un apoyo emocional muy importante, porque ellos son maestros que forman personas éticas y apasionadas. A Dino gracias por enseñarme a cerrar los ojos para ver los seres vivos y a Juliana porque es la personificación de la nobleza. A Arturo Aristizábal, amante de las zamias, porque de alguna manera él fue quién nos introdujo en la apreciación de éstas plantas. También quisiera agradecer a todos mis profesores especialmente a Maria José Sanín, por su sensibilidad, brillantez y apoyo durante la formulación del proyecto. También de manera muy especial a Cristina López-Gallego, gran estudiosa de las cícadas, por los consejos, la literatura, los insectos y las invitaciones a sus salidas de campo de zamias. Agradezco a todas las personas que me acompañaron durante el trabajo de campo: Cornelio Bota, Yeyson Montoya, Miguel González, Laura Isabel Marín, Laura Gómez Mesa, Eliana, Héctor Manuel Arango, Edwin Hurtado, Camilo Flórez y Sebastián Cifuentes. Especialmente quiero agradecer a Cornelio Bota y Yeyson Montoya quienes estuvieron en la mayor parte del trabajo de campo en el zamial, por la alegría y la música; que enmarcados en ese hermoso bosque tropical constituyen de los momentos más preciados en este trabajo de grado. A Henry Arenas-Castro, Cristina López-Gallego, Maria José Sanín, Edwin Hurtado, Alejandra Duque y Nicolás González-Castro por sus sugerencias al manuscrito. Agradezco muy especialmente a Gustavo Londoño y a Juan Luis Parra por el préstamo desinteresado de los equipos para registrar termogénesis. A la Beca Colombia Biodiversa de la Fundación Manuel Ángel Escobar por la financiación parcial de este estudio. A Cornelio Bota y Camilo Flórez, mis dos naturalistas favoritos, porque para mí son el ejemplo vívido de la pasión por los insectos y el monte. A Sergio A. Muñoz-Gómez por su incondicional acompañamiento intelectual y emocional. A Henry Arenas-Castro, por todas las conversaciones y porque ojalá sigan siempre instaladas en lo que me es imprescindible. A Edwin Hurtado por todo su apoyo durante el tiempo de campo y escritura, y principalmente por sus matices de profundidad y ternura. A Yeyson Posada, la ceiba, por entender las zamias y regalármelas en cada ojo dibujado. A Michael Castaño, el Sol, por toda la alegría desprendida de su elocuente sensibilidad. A Duberney Chaverra por introducirme con paciencia en el maravilloso mundo de los insectos. A todos mis compañeros de la carrera: Vanessa Correa, Norman Echavarría, Elisa Lotero, Laura Gómez, Alejandra Duque, Natalia Duque, Valeria Zapata, Juan Diego Ospina, Lina Bolívar, Camila García, Tomás Vallejo, Pedro Gómez, Esteban Urrea, Hana Londoño y Paula Saravia por el acompañamiento tan importante durante el pregrado. A mis amigos Yenny Correa, Isabel Restrepo, Laura Muñoz, Camila Londoño, Lorena Quintero, Carolina Torres, Eliana Pineda, Juliana Herrera, Juliana Cerón, Sheela Turbek, Nancy Chen, Matt Dickinson, Héctor Arango, Yeyson Posada, David Taborda, Daniel Monsalve, Alejandra Clavijo, Blanca Arbeláez, Sebastián Cifuentes, Laura Sierra, Manuel Sánchez, Alejandro Ospina, Giovanny Pérez, Mauricio Zapata, Alejandro Ríos, Juan Guillermo Mesa, Juan David Sánchez, Alejandra Barrada, Liliana Palacio, Samuel Monsalve y David Mechas. También a Javier Cardona, David Cardona, Dolly Montoya, Alejandra Cardona, Rosario Giraldo, Celia Ramos, Carlos Montoya, Eliana Moyano, Paulina Valencia, Sandra Ortiz, Gloria Quintero, Juan Cardona y Margarita Cardona y todos los tíos y primos que se me escapan. A Juanita mi gata, lo más querido, porque este trabajo de grado fue escrito en compañía de ella y de Pala. A todos los integrantes del Grupo Abarco, Tejuntas y al Colectivo Bidikay por soñar conjuntamente con un país que tenga una mirada más sensible de su rica diversidad biológica y cultural. Finalmente deseo agradecer a los habitantes de la vereda Santa Bárbara, especialmente a Doña Alba y sus hijas Darsy y Yireth, por recibirnos siempre con tanto cariño. A Don Arturo quién nos transportó hasta la vereda, y muy especialmente a Miguel González por ser un apasionado de la diversidad y la conservación. Medellín, 1-VI-2015 Manuscript for Submission to Arthropod-Plant Interactions (Original Version) Research Article Pollination of Zamia incognita A. Lindstr. & Idárraga on a natural population in the Magdalena Medio, Colombia WENDY A. VALENCIA-MONTOYA1, DINO TUBERQUIA1, 2 & JULIANA CARDONA- DUQUE1, 2 1Programa de Biología, Universidad CES-EIA, Medellín, Colombia Calle 10 A No. 22 - 04 2 Grupo Biología CES-EIA Correspondence to: Wendy A. Valencia-Montoya. E-mail: [email protected] Telephone number: (+57) (4) 444 0555 Ext. 1240 Fax number: (+57) (4) 3113505 Abstract: The genus Zamia (Zamiaceae: Cycadales) holds its greatest diversity in Colombia and most of its species are highly threatened by different factors. One of the most relevant and frequently ignored aspects for the establishment of effective conservation programs, is its highly specialized reproductive biology. Despite the importance of pollination for the viability of the Zamia populations, there are no studies about the pollination process for these cycads in Colombia. Herein we describe the pollination process of Zamia incognita A. Lindstr. & Idárraga, in a natural population from the Magdalena Medio valley in Colombia. We found Pharaxonotha beetles in the male cones where they complete all phases of their lifecycle. Cones produce heat, which generally follows a circadian pattern and its magnitude and extent, are positively correlated with the elongation of the cones and pollen shedding; therefore, the increment in cones temperature seems to play an important role in attracting the beetles to the cones. By following marked beetles with fluorescent dyes as well as by direct observations of the beetles on and into female cones, remaining at pollination droplets for a long period, we confirmed that Pharaxonotha sp. is the effective pollinator of Zamia incognita. We suggest that the pollination droplets may serve as reward to the pollinators. We also discuss the relationships of this Zamia species with other insects like Eumaeus butterflies, Atta ants, flies and Meliponinae bees. Key words: cycad pollination, Erotylidae, plant thermogenesis, plant-insect interactions. Introduction The pollination in Zamiaceae was considered to be anemopholus until the 80’s end. However, this was a misconception due to a generalization of most Gymnosperms condition, and it has been reassessed by extensive evidence of insect pollination (e.g. Donaldson 1997, Norstog et al. 1986, Suinyuy et al. 2009, Tang 1987a, Terry 2001, Terry et al. 2005, Vovides 1991, Vovides et al. 1997, Wilson 2002). During most of the XX century, entomophilous pollination was thought to be restricted to Angiosperms, and it has been considered as a key interaction that had played a pivotal role in promoting the overwhelming diversity of the interacting groups (Gorelick 2001, Norstog 1987). Pioneering studies in Zamia pollination challenged this paradigm (e.g. Norstog et al. 1986, Tang 1987a) and now we know that pollination in all the Zamiaceae genera relied on highly specialized relationships with insects, most of them beetles (Donaldson 1997, Norstog et al. 1986, Suinyuy et al. 2009, Tang 1987a, Terry 2001, Terry et al. 2005, Vovides 1991, Wilson 2002) and secondarily thrips (Terry 2001). The cycads exhibit an array of complex traits to attract insects: the female and male cones have differential attractants for feeding preferences; the pollinator life-cycles are coupled with plant phenology; and they volatilize their odors and have thermogenesis (Donaldson 1997, Norstog and Fawcett 1989). These features are shared with some beetle-pollinated angiosperm families (e.g. Cyclanthaceae: Eriksson 1994; Annonaceae: Gottsberger 1999; Arecaceae: Henderson 1986) and they are considered evolutionary convergences related to cantharophily (Norstog 1987, Terry et al. 2005). Due to the similarity of these cycad-insects interactions across different continents, and because most of the pollinator species belong to the not related beetle superfamilies Curculionoidea and Cucujoidea, it is now believed that the insect pollination has been independently evolved in different Zamiaceae genera (Suinyuy et al. 2009, Terry et al. 2004, 2005, Wilson 2002). These specialized associations indeed have important implications in cycad species conservation,
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