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Anthropic Habitats As Novel Environments

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Anthropic Habitats As Novel Environments Anthropic habitats as novel environments: consequences for evolution and conservation Maider Iglesias-Carrasco PhD Thesis 2017 (c)2017 MAIDER IGLESIAS CARRASCO Title: Anthropic habitats as novel environments: consequences for evo- lution and conservation. Author: Maider Iglesias Carrasco. University: Faculty of Science and Technology – University of the Basque Country. Universidad del País Vasco/Euskal Herriko Unibertsita- tea (UPV/EHU). Year: 2017 PhD supervisors: Dr. José Martín, Dr. Carlos Cabido Cite as: Iglesias-Carrasco, M. 2017. Anthropic habitats as novel environ- ments: consequences for evolution and conservation. University of the Basque Country, Faculty of Science and Technology, Spain. 196 pp. Cover: Carlos Cabido Finance: The present Thesis has received financial support from the Minis- try of Education (Spanish Goverment): Ayudas para la formación de profesorado universitario (FPU), convocatoria 2012. A mis padres y abuela Index Agradecimientos________________________________________________________1 Summary______________________________________________________________3 Introduction ___________________________________________________________5 1. Anthropic environments as new habitats _______________________________7 1.1 Urbanization _______________________________________________8 1.2 Exotic tree plantations _______________________________________9 2. Adaptive and maladaptive responses of animals to anthropic environments __11 2.1 Effects on animal diversity __________________________________11 2.2 Anthropic habitats as ecological and evolutionary traps ____________14 2.3 Sensory disturbance ________________________________________17 2.4 Disease and pathogens ______________________________________18 2.5 Behavioural responses to changing conditions ___________________19 3. Amphibians and reptiles as study species _____________________________22 4. Conservation interest of anthropic habitat ____________________________23 4.1 Urban areas ______________________________________________23 4.2 Exotic Plantations __________________________________________24 5. References _____________________________________________________24 Objectives ___________________________________________________________33 Chapter 1. Exotic plantations lead to lower species richness, density and condition of amphibians: linking field observations and physiological responses in meso- cosms. ________________________________________________________37 Chapter 2. Condition-dependent trade-offs between sexual traits, body condition and immunity: the effect of novel habitats.________________________________63 Chapter 3. Secondary compounds from exotic tree plantations change female mating preferences in the palmate newt (Lissotriton helveticus). _________________89 Chapter 4. Leaf extracts from an exotic tree affect responses to chemical cues in the palmate newt (Lissotriton helveticus). _______________________________109 Chapter 5. Urban habitats can affect body size and body condition but not immune response in amphibians. __________________________________________135 Chapter 6. Habitat dependent effects of experimental immune challenge on lizard antipredator responses. ___________________________________________155 General conclusions __________________________________________________177 Annex 1. Resumen general _____________________________________________183 Annex 2. Contributions ________________________________________________191 Agradecimientos Han sido muchas las personas que me han animado y ayudado en este largo y arduo proyecto que ahora culmina, y a todas ellas quiero transmitir mi más sincero agradecimiento. Me gustaría comenzar diciendo a mis padres lo importante que ha sido para mí su compañía y su apoyo en este trayecto, ayudándome tanto emocionalmente en los momentos difíciles (que no han sido pocos), como participando activamente en muchos de los experimentos realizados: recogiendo hojas de los bosques, acompañándome en excursiones diurnas y nocturnas, con frío, lluvia y viento, en busca de tritones, sapaburus y lagartijas... ¡Menos mal que nadie perdió un dedo por congelación! Mi familia siempre ha estado a mi lado, pero quiero mencionar, en especial, los cuidados y el cariño de mi abuela, Eintxels, que me ha protegido siempre, ¡aunque sin entender muy bien por qué aún continúo “estudiando”! Carlos ha compartido conmigo este periplo desde el principio, cuando sólo era una idea. Con él empecé a vislumbrar y valorar algunos comportamientos animales en los que nunca antes había reparado, manteniendo largas conversaciones sobre relación y evolución de diferentes especies, que generaron ilusión y proyectos. Su entusiasmo por la ciencia ha sido fundamental para llegar hasta aquí. Gracias a Jose por su ayuda con el papeleo. Al final tengo que darle la razón y constatar que el doctorado, más que para hacer ciencia, ¡te prepara para lidiar con la burocracia del país! Gracias también por su ayuda y comentarios en los manuscritos e introducción de la tesis, y por las mañanas en el Ventorrillo organizando los experimentos. Iván Gómez-Mestre y Pablo Burraco me recibieron en Doñana, donde tuve que enfrentarme a un verdadero trabajo de laboratorio con el que no contaba cuando empecé la tesis. Supuso toda una experiencia, con altibajos y dificultades, pero también con buenos momentos. Gracias, Iván, por tu apoyo y tus consejos, ¡nunca los olvidaré! Aitor Larrañaga ha hecho posible el primer capítulo de esta tesis. Gracias por aguantar todas las idas y venidas del manuscrito. Mención aparte merecen Michael Jennions y Megan Head. Ellos me acogieron, orientaron y guiaron en Canberra cuando más frustrada me encontraba. Desde el comienzo me hicieron sentir como en casa. Su apoyo constante me situó de nuevo en la dirección adecuada y con ellos ¡publiqué mi primer paper! ¡Qué gran momento! Y, aunque después ha habido varios rechazos a la hora de publicar, sus alentadoras palabras consiguieron que no me rindiera en ningún momento. Han sido unos mentores geniales e increíbles compañeros. Thank you very much, without you I would not get it. Martin Whiting ha sido otro pilar en Australia. Gracias a su amable aceptación en su lizard lab pude pasar varios meses en Sydney conociendo gente nueva y 1 volviendo a tener contacto con herpetos en un país lejano. Un recuerdo muy especial a Aranzadi y a su gente, sobre todo a Ion Garin. Gracias a él y a su ayuda en el campo (¡cientos y cientos de horas!) he podido realizar muchos más experimentos de los planeados, y muchas gracias, también, a toda la gente que me ha ayudado a conseguir animales: Iñaki, Egoitz, Aitor Laza, Janire, Julen..., y buenos amigos interesados en temas de naturaleza como Floren y familia. Mi sincero agradecimiento, así mismo, a Alberto Gosá, por ayudarme a mejorar la escritura, y a Josema y Mertxe, por su ayuda en los proyectos. Agradecer, naturalmente, a todos aquellos que han sufrido a mi lado la temida, tediosa y agotadora burocracia, sobre todo Juan Carlos Iturrondobeitia y Raimundo. ¡Gracias por vuestra paciencia! Todos mis amigos se han mantenido a mi lado en los momentos más bajos. Gracias por quitar hierro a situaciones difíciles. Entre ellos, Oier y Alejandra han sido únicos. Gracias por las innumerables horas en el campo y las pizzas nocturnas en Aranzadi entre grabación y grabación. Quisiera mencionar también algunos amigos dentro del ámbito científico como Xim Cerdá, Elena Angulo, Sergio Naretto, Fran Encinas, Vlatka Horvat o John Endler, que me han ayudado con inestimables consejos acerca de cómo abordar la ciencia, la estadística y la publicación de manuscritos. Por último, David ha sido durante este último año una compañía imprescindible, apoyándome y tendiéndome la mano con paciencia y optimismo, incluso en situaciones ciertamente complicadas. Quiero expresar mi agradecimiento por su chévere, férreo e incondicional apoyo. Mila esker danoi. 2 Summary This Thesis explores how the rapid transformation of the natural habitat to an- thropic environments alters the behavioural responses and life-history traits of amphibi- ans and reptiles. This project focuses on two of the most widely extended anthropic hab- itats of the world: urban habitats and exotic tree plantations. Anthropic environments are actual new habitats, with selective pressures different from natural ones. The popu- lations inhabiting these disturbed habitats need to quickly evolve or to show phenotypic plasticity to adapt and survive in the new environmental conditions. In that sense, an- thropic habitats provide a unique opportunity to study the adaptation process (or the lack of it) and the ability of the species to plastically respond to the new environmental pressures, so in the last years the interest of the scientist on this research area has in- creased drastically. This Thesis delves into the effect of the presence of these two novel environ- ments on the functional ecology of individuals, using amphibians and reptiles as model study species. The Thesis includes both field work and laboratory experiments, some- times combining both techniques in a single chapter to refine the results. Exotic plantations First, field work showed that the richness and abundance of amphibians was lower in pine and eucalypt plantations compared to the natural oak forest. We combined this data with an experimental study in mesocosms were we found that the presence of leachates released by the exotic trees may explain, at least in part, the results found in the field (Chapter 1). The next three chapters (Chapters 2-4) are based
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