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Phd Thesis Phd BEHAVIOURAL AND PHYSIOLOGICAL STRESS RESPONSES TO ENVIRONMENTAL AND HUMAN FACTORS IN DIFFERENT SMALL MAMMAL SPECIES: IMPLICATIONS FOR THEIR CONSERVATION Álvaro Navarro Castilla Madrid 2015 PhD Thesis PhD UNIVERSIDAD AUTÓNOMA DE MADRID Facultad de Ciencias Departamento de Biología Fotografía de portada: ratones de campo Apodemus sylvaticus, Cantabria, España. Autor: Ángel Ruiz Elizalde UNIVERSIDAD AUTÓNOMA DE MADRID Facultad de Ciencias Departamento de Biología BEHAVIOURAL AND PHYSIOLOGICAL STRESS RESPONSES TO ENVIRONMENTAL AND HUMAN FACTORS IN DIFFERENT SMALL MAMMAL SPECIES: IMPLICATIONS FOR THEIR CONSERVATION RESPUESTA COMPORTAMENTAL Y DE ESTRÉS FISIOLÓGICO A FACTORES AMBIENTALES Y HUMANOS EN DIFERENTES ESPECIES DE MICROMAMÍFEROS: IMPLICACIONES PARA SU CONSERVACIÓN Álvaro Navarro Castilla TESIS DOCTORAL Madrid 2015 UNIVERSIDAD AUTÓNOMA DE MADRID Facultad de Ciencias Departamento de Biología BEHAVIOURAL AND PHYSIOLOGICAL STRESS RESPONSES TO ENVIRONMENTAL AND HUMAN FACTORS IN DIFFERENT SMALL MAMMAL SPECIES: IMPLICATIONS FOR THEIR CONSERVATION RESPUESTA COMPORTAMENTAL Y DE ESTRÉS FISIOLÓGICO A FACTORES AMBIENTALES Y HUMANOS EN DIFERENTES ESPECIES DE MICROMAMÍFEROS: IMPLICACIONES PARA SU CONSERVACIÓN Memoria presentada por Álvaro Navarro Castilla para optar al grado de Doctor Europeo en Ciencias Biológicas por la Universidad Autónoma de Madrid Bajo la dirección de la Dra. Isabel Barja Núñez, Profesora del Departamento de Biología de la Universidad Autónoma de Madrid El doctorando Vº Bº de la Directora Álvaro Navarro Castilla Dra. Isabel Barja Núñez Madrid, Noviembre 2015 A mi familia A ti AGRADECIMIENTOS Por fin ha llegado el gran momento que tanto esperaba… La elaboración de una tesis doctoral conlleva mucho esfuerzo y si he llegado hasta aquí no ha sido sólo por el mérito propio sino también gracias a todos vosotros. Por eso me gustaría, a través de estas líneas, expresar mi más sincero agradecimiento a todas las personas que de una manera u otra han colaborado en la realización de esta tesis doctoral. En primer lugar me gustaría agradecer a mi directora de tesis, la Dra. Isabel Barja Núñez, por haberme ofrecido esta gran oportunidad y por haber confiado en mí desde el principio. Muchas gracias por poner a mi disposición todos los medios que estaban a tu alcance, por la acertada orientación y formación durante todos estos años y por animarme y hacerme partícipe de tantas ideas, colaboraciones, congresos y proyectos. Gracias porque si he llegado tan lejos ha sido porque tú me has transmitido tus conocimientos y experiencia. No tengo palabras suficientes de agradecimiento porque sin tu esfuerzo, dedicación infinita e inestimable ayuda, sin ti, simplemente esto no hubiera sido posible. Por eso, el mayor reconocimiento te lo debo a ti, por ser una gran científica, persona y amiga. Gracias Isa, de corazón. También me gustaría expresar mi agradecimiento a la Universidad Autónoma de Madrid y en particular a la Unidad de Zoología donde he podido formarme como docente e investigador. Gracias a todos vosotros por ayudarme y aconsejarme y por los buenos momentos que he compartido con cada uno de vosotros. Especialmente me gustaría agradecer al Dr. Javier de Miguel por haber estado desde el comienzo de esta aventura y aceptar con agrado también participar en el final, gracias por ser mi tutor durante los cursos de doctorado y por los buenos momentos a lo largo de estos años. No me puedo olvidar del Dr. Juan Moreira por compartir conmigo sus conocimientos en investigación “punteira” y de Beatríz Sánchez por ser una compañera estupenda y por ayudarme con los últimos retoques de este trabajo. No me quiero olvidar de todas aquellas personas con las que pude participar en distintos proyectos y trabajos de campo, muchas gracias a la Dra. Ana Piñeiro, Dr. Pedro Olea, Dra. Patricia Mateo-Tomás, Dr. Juan Malo, Dra. Cristina Mata y a Pablo Ruíz-Capillas, gracias por las buenas experiencias y por vuestra ayuda porque parte de AGRADECIMIENTOS esta tesis es gracias a vosotros. Y especialmente me gustaría agradecerle al Dr. Mario Díaz por el apoyo logístico, por tus revisiones y comentarios, y por tu paciencia infinita para explicarme y aclararme cualquier duda que me surgía, gracias Mario. También debo de agradecer al centro de naturaleza Cañada Real (Peralejo, Madrid), especialmente a José España, por proporcionarme el material necesario para poder realizar parte de los experimentos que comprenden esta tesis. I am also especially grateful to Dr. Rupert Palme, Dr. Carlos Fernandes and Dr. Dustin J. Penn for accepting me in their corresponding laboratories, for sharing your precious knowledge and experience and for the revision of several manuscripts of this thesis making useful comments to improve them. Thank you very much for your generous help along these years. También tengo mucho que agradecerle a mi gran amiga la Dra. Sandra Revilla- Fernéndez, Sandri (que suena mejor entre nosotros), muchas gracias de verdad por tu amistad a lo largo de estos años, por darme la oportunidad de aprender tanto en ese magnífico laboratorio y por enseñarme tantas técnicas y protocolos, pero sobre todo por hacerme sentir especial al confiar en mí para llevar todo a cabo. ¡Gracias! Son mucho los amigos que han estado a mi lado a lo largo de esta estapa, algunos implicados directamente en este trabajo ayudándome en el campo, resolviendo dudas o simplemente siendo un gran apoyo aunque no llegasen a entender lo que hacía… Pues aquí lo tenéis, espero que esto os ayude a comprender lo importante que era para mí y lo mucho que habéis significado. A mis amigos de hoy y siempre, Laura, Lu, Adrián, Wander y especialmente a Will que aunque ya no estés aquí, esta tesis también va dedicada a ti. A mi gente de Viena (Susana, Paloma y Karolina) por vuestra magnífica compañía y amistad durante mis estancias allí, y a mis chicas de veterinaria Anita, Didi, Encarna, Gaby, Mari, y especialmente a Patri porque tú más que ninguna sabes lo que esto ha costado, gracias por tu apoyo, tus ánimos, tu compañía durante tantos días interminables y por todos los momentos inolvidables. Mil gracias o mejor dicho millones de gracias a Alejandro Bajo (tranquilo hombre que no me había olvidado de ti) por las magníficas ilustraciones que has realizado para cada uno de los capítulos de esta tesis porque sin ellas este trabajo no sería lo mismo. Pero también me gustaría agradecerte todo tu apoyo incondicional y los AGRADECIMIENTOS ánimos que me has dado durante todo este tiempo ya que han sido muy importantes para mí y me han impulsado cada día a seguir avanzando hasta el final. ¡Gracias por haber estado y seguir estando ahí! Finalmente, pero con un cariño muy especial por ser los más importantes, quiero agradecerles a mis padres, Santi y Justi, todo lo que me habéis enseñado, que habéis confiado en mí apoyándome y facilitándome el camino para que pudiera estudiar, formarme y poder dedicarme a lo que más me gusta. Gracias porque aunque no os lo diga muy a menudo sabéis que os quiero mucho y que esto es tan vuestro como mío. También a mis hermanos, Alejandro y Alberto, porque vosotros también habéis aportado ayudándome con las mil cosas que os he pedido y siempre que lo he necesitado. Y a mi tía favorita (sí Marivi, eres tú) porque siempre me has animado y aconsejado para poder alcanzar esta meta. Gracias también a mis abuelos, María que aún está y a los que ya se fueron, a vosotros os dedico esta tesis. Esta tesis ha sido posible gracias a una beca de Formación de Profesorado Universitario (FPU) del Ministerio de Educación y Ciencia de España. Y a la financiación parcial y colaboración con los siguientes proyectos de investigación: - SEK02B06 - CGL2006-05047/BOS - CCG10-UAM/AMB-5325 - CENIT-OASIS - REMEDINAL-2 (S-2009/AMB/1783) - ACORNSIZE (CGL2006-06647/BOS) - MONTES (CSD2008–00040) - RISKDISP (CGL2009- 08430/BOS) - VULGLO (CGL2010-22180-C03-03) CONTENTS ABSTRACT ................................................................................................................... 1 RESUMEN ..................................................................................................................... 3 INTRODUCTION ........................................................................................................... 5 Risk of predation and chemical communication ................................................... 5 Interspecific competition ..................................................................................... 7 Human disturbances .. .......................................................................................... 9 Physiological stress responses ....................................................... ......................12 Study species – Small mamals ..................... ……………………………..…….15 RESEARCH AIMS ....................................................................................................... 19 CHAPTER 1. RISK OF PREDATION .......................................................................... 21 PAPER I. Antipredatory response and food intake in wood mice (Apodemus sylvaticus) under simulated predation risk by resident and novel carnivorous predators ............................................................................................................... 23 Abstract ............................................................................................................. 23 Introduction ....................................................................................................... 24 Objetives ..........................................................................................................
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