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Sensory Ecology of Spiders That Prey on Frogs

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Sensory ecology of spiders that prey on frogs Thesis submitted by Martha Patricia Torres Sánchez Grupo de Ecofisiología, Comportamiento y Herpetología Departamento de Ciencias Biológicas Universidad de los Andes Under the Supervision of Professor Adolfo Amézquita, PhD Dissertation presented in Partial Fulfilment of the Requirements for the Degree of Doctor of Sciences – Biology Universidad de los Andes Bogotá, Colombia August 2019 “Si nos aventuramos en el conocimiento y en la ciencia, lo hacemos tan sólo para regresar mejor equipados para la vida” Johann Wolfgang von Goethe, 1806 Dedication For my beloved son Martín, thank you for your laughter, for your games and for your caresses, thanks for being here and for making my life happy. For my life partner, Carlos for all the love and unwavering support you have given me. For my mother, who has supported me in everything and has helped me to become the person that I am today. For my family, who were always by my side and heard me talk about my research, even if they did not fully understand why I do what I do. They have all the gratitude and appreciation that I am able to muster. ii Agradecimientos La parte más difícil de la escuela de posgrado es lograr un equilibrio entre el trabajo y la vida personal. Durante mi tiempo como estudiante de posgrado en la Universidad de los Andes, tuve la suerte de conocer gente que hizo que los momentos de trabajo y vida fueran más productivos y agradables. Estoy inmensamente agradecida con Adolfo Amézquita por apoyar los diferentes caminos que recorrí durante mi doctorado. Me dio suficiente independencia para permitirme construir mi propia senda, y ayudarme a resolver cuando las cosas estaban a punto de salirse de las manos. Gracias por recibir a esta aracnóloga, que nunca se sintió como “mosco en leche” en el GECOH. Sobre todo, gracias por la paciencia y persistencia durante estos años. Adolfo es un editor absolutamente increíble y mi redacción y esta tesis se han beneficiado de ello. A todos los antiguos y actuales miembros del GECOH por su generosa ayuda durante este proceso. Gracias especiales a Pablo Palacios, Diana Motta, Mabel González, Sofía Rojas, Pedro Patricio, Sebastián Di Doménico, Sebastián Gómez, Catalina Orejuela, Santiago Mesa, Roberto Márquez, Iván Beltrán, Manuel Guayara, Leidy Barragán, Vicky Flechas y Javier Méndez por su ayuda durante la recolección de datos de campo y las primeras ideas de este trabajo. Mileidy Betancourt y Diana Galindo me han aconsejado, animado y apoyado – tanto en lo académico como en lo personal– a lo largo de estos años. Su amistad y ayuda han sido especialmente valiosas durante los últimos meses de redacción. Diana Pinto y Camilo Rodríguez hicieron extremadamente productivo y agradable el trabajo de campo. Les doy gracias por su paciencia para enseñarme a reconocer las ranitas y apoyar mis experimentos vibracionales y comportamentales. A mis colegas de la Universidad Central Javier Casas y Mauricio Vergara, quienes diseñaron y facilitaron generosamente equipos para realizar mis experimentos y respondieron el teléfono (¡incluso en navidad!) para solucionar iii problemas. Les agradezco por empujarme a pensar en nuevas formas de abordar mis preguntas y guiarme a través de los complicados conceptos de la física del sonido. Yeltsin Torres y Gustavo Lozano, estudiantes de ingeniería electrónica (UC) siempre fueron un oído dispuesto, me enseñaron habilidades útiles para la vida (¡como soldar!) y a no tener miedo de la física para medir la vibración con paciencia y buen sentido del humor. A Eileen Hebets de la Universidad de Nebraska en Lincoln (UNL), quien me recibió en su laboratorio durante la pasantía y puso a disposición mía su tiempo y los recursos necesarios para completar con éxito esta etapa del doctorado. Su puerta siempre estaba abierta, ya fuera para charlar sobre mi investigación, el desarrollo de los experimentos y análisis vibracionales, o solo sobre la vida. Allí tuve la oportunidad de conocer a un talentoso grupo de investigadores quienes se convirtieron en amigos de confianza. Agradezco a Rowan McGinley por ser tan increíblemente generoso con su tiempo, por apoyarme y guiarme durante mi estancia en UNL. Rowan, Laura Segura, Colton Watts, Tyler Corey, Bridget Bigner, Laura Gatch y Fiona Grey; me animaron a pensar en la logística y metodologías de muchas ideas de investigación presentes y futuras. Les agradezco su entusiasmo por mi investigación. Goran Mihaijlovik y Juan Carlos Tamayo directores de la Reserva Tanimboca, me brindaron su apoyo logístico para conducir esta investigación. Leonaldo, Sarita, Ramiro, José y Rodolfo proporcionaron un apoyo inestimable durante el trabajo de campo en Leticia. El Amazonas es uno de los lugares más especiales del planeta y lo que lo hace más especial, son las personas verdaderamente increíbles que allí conocí. He tenido experiencias inolvidables y aprendí mucho de su dedicación y grandes corazones. También agradezco a todos los colegas y amigos que han estado de una u otra forma. A Daniel Garavito por su acompañamiento, sus minuciosos y rigurosos comentarios y correcciones. iv Gracias a mi comité de evaluación por sus sugerencias constructivas para mejorar este trabajo. Eduardo Flórez, Jorge Molina y Rodrigo Villemart, ustedes han sido increíblemente generosos y comprensivos, y aprecio su paciencia y visión de este trabajo. Mi investigación no hubiera sido posible sin la asistencia técnica y administrativa y apoyo en el Departamento de Ciencias Biológicas. Mi más grande agradecimiento a mi familia. A mi madre, hermanas y sobrinos por animarme siempre a pensar y soñar en grande. Por su amor y apoyo infinitos a lo largo de estos años, asistiendo en el cuidado de Martín durante los meses lejos de mi hogar. Y, por último, agradezco a mi hijo Martín quien desde el vientre acompañó el trabajo de campo y se convirtió en mi compañero de travesía. A mi esposo Carlos, quien con gran entusiasmo recolectó arañas, observó ranas y transportó materiales y equipos. Me acompañó a lo largo del proceso de escritura, dándome un abrazo después de un día particularmente difícil o estupendo. No tengo palabras para agradecerles por simplemente estar allí y apoyarme incondicionalmente. Mi investigación fue financiada con el apoyo de la Universidad de los Andes a través de los Proyectos Semilla y la Asistencia Graduada, la Universidad Central a través del Programa de Apoyo a Estudios de Posgrado; la Unidad de Diseño, Innovación e Integración de Tecnología (DIT); y el Departamento de Ciencias Naturales y Colciencias a través del proyecto ID#52037. La investigación fue aprobada por el CICUAL de la Universidad de los Andes (Protocolo CFUA 16-014) y se llevó a cabo de conformidad con el Permiso Marco de Recolección de Especímenes de Especies Silvestres de la Diversidad Biológica con Fines de Investigación Científica No Comercial #1386. v Tabla de contenido Dedication ...........................................................................................................................ii Agradecimientos ................................................................................................................ iii General Abstract ................................................................................................................ 1 Resumen General .............................................................................................................. 3 General Introduction .......................................................................................................... 6 Thesis Aims and Objectives ........................................................................................... 9 Study System ............................................................................................................... 10 Literature cited ................................................................................................................. 13 Chapter I: Frog-eating Spiders: An Ecological Accident or a Specialized Guild? .............. 22 Abstract ........................................................................................................................... 23 Introduction ...................................................................................................................... 24 Methods ........................................................................................................................... 26 Data Collection and Search Criteria ............................................................................. 26 Sampling the Spatial Overlap Between Spiders and Putative Prey ............................... 28 Predation Experiments ................................................................................................. 29 Results ............................................................................................................................ 30 Geographic distribution and spider pervasiveness in anuran predation ........................ 30 Co-occurrence and behavioural interactions between spiders and frogs: How Important are Anuran Predation? ....................................................................................................... 33 To what extent do spiders feed on anurans? ................................................................ 34 Discussion ....................................................................................................................... 34 Acknowledgments ...........................................................................................................
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  • Reflections on the Tapetum Lucidum and Eyeshine in Lycosoid Spiders

    Reflections on the Tapetum Lucidum and Eyeshine in Lycosoid Spiders

    2013. The Journal of Arachnology 41:43–52 Reflections on the tapetum lucidum and eyeshine in lycosoid spiders Kari Benson: School of Sciences, Lynchburg College, 1501 Lakeside Dr., Lynchburg, Virginia 24501, USA. E-mail: [email protected] Robert B. Suter: Department of Biology, Vassar College, 124 Raymond Avenue, Poughkeepsie, New York 12603, USA Abstract. In the lycosoid spiders, the secondary eyes possess a grate-shaped tapetum lucidum that reflects light, causing eyeshine when these spiders are viewed with approximately coaxial illumination. This guanine-based reflective surface is thought to increase visual capabilities in low light. We explored the eyeshine of the posterior medial eye in eight taxa of pisaurid and lycosid spiders. The taxa included four pisaurids: Dolomedes tenebrosus Hentz 1844, D. triton (Walckenaer 1837), D. scriptus Hentz 1845 and D. vittatus Walckenaer 1837; and four lycosids: Gladicosa pulchra (Keyserling 1877), Hogna sp. (cf. Lycosa lenta (Hentz 1844) sensu Wallace 1942), Rabidosa punctulata (Hentz 1844) and Varacosa avara (Keyserling 1877). We found that there were significant family- and species-level differences in both the reflected spectra and the intensity of reflection. Although the peaks of the reflected spectra were in the green range for all spiders, the mean peak was further toward the blue end of the spectrum for the lycosids than for the pisaurids. Variation among species (about 54% of the total variation) was dominated by G. pulchra (Lycosidae) and D. vittatus (Pisauridae), both of whose spectra peaked near yellow, vs. V. avara (Lycosidae), whose spectra peaked to the blue side of green. The lycosid spiders showed overall brighter eyeshine.