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Effects of Environmental Variation on the Reproduction of Two Widespread Cervid Species

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Effects of Environmental Variation on the Reproduction of Two Widespread Cervid Species UNIVERSIDAD POLITÉCNICA DE MADRID ESCUELA TÉCNICA SUPERIOR DE INGENIERÍA DE MONTES, FORESTAL Y DEL MEDIO NATURAL EFFECTS OF ENVIRONMENTAL VARIATION ON THE REPRODUCTION OF TWO WIDESPREAD CERVID SPECIES DOCTORAL DISSERTATION MARTA PELÁEZ BEATO Ingeniera Técnica Forestal Máster en Investigación Forestal Avanzada 2020 PROGRAMA DE DOCTORADO EN INVESTIGACIÓN FORESTAL AVANZADA ESCUELA TÉCNICA SUPERIOR DE INGENIERÍA DE MONTES, FORESTAL Y DEL MEDIO NATURAL EFFECTS OF ENVIRONMENTAL VARIATION ON THE REPRODUCTION OF TWO WIDESPREAD CERVID SPECIES DOCTORAL DISSERTATION MARTA PELÁEZ BEATO Ingeniera Técnica Forestal Máster en Investigación Forestal Avanzada 2020 THESIS ADVISORS: ALFONSO RAMÓN SAN MIGUEL AYANZ PEREA GARCÍA-CALVO Doctor Ingeniero de Montes Doctor Ingeniero de Montes LECTURA DE TESIS Tribunal nombrado por el Sr. Rector Magnífico de la Universidad Politécnica de Madrid, el día _____________de ________________de 2020. Presidente/a: _____________________________________ Secretario/a: _____________________________________ Vocal 1º: ________________________________________ Vocal 2º: ________________________________________ Vocal 3º: ________________________________________ Realizado el acto de defensa y lectura de la Tesis el día ____ de _______de 2020, en la Escuela Técnica Superior de Ingeniería Forestal y del Medio Natural, habiendo obtenido calificación de _______________________. Presidente/a Secretario/a Fdo.:_______________________ Fdo.:_______________________ Vocal 1º Vocal 2º Vocal 3º Fdo.:_______________ Fdo.:_______________ Fdo.:_______________ iii MENCIÓN DE DOCTORADO INTERNACIONAL INTERNATIONAL DOCTORATE MENTION Esta Tesis ha sido informada positivamente para su defensa en exposición pública por los siguientes investigadores: This Ph.D. Thesis has been positively evaluated for its public defense by the following external reviewers: Dr. Florianne Plard Department of Statistics University of Lyon 2 (France). Dr. Francisco Ceacero Department of Animal Science & Food Processing Czech University of Life Sciences Prague (Czech Republic) v Este trabajo ha sido posible gracias a la financiación del Ministerio de Educación, Cultura y Deporte a través de un contrato predoctoral para la Formación de Profesorado Universitario (FPU13/00567). Las estancias en el extranjero han podido llevarse a cabo gracias a dos ayudas de movilidad, para estancias breves en otros centros españoles y extranjeros y para traslados temporales a centros extranjeros, dirigidas a beneficiarios del subprograma de Formación del Profesorado Universitario (EST14/00304 y EST16/00095) y a la convocatoria de ayudas del Consejo Social de la Universidad Politécnica de Madrid para el Fomento de la Formación y la Internacionalización de Doctorandos para el curso 2015-2016. También se ha contado con la ayuda del Ministerio de Educación, Cultura y Deporte para cubrir el pago de los precios públicos de las matrículas por tutela de tesis en los programas de doctorado. vi A mis padres y a Rafa vii Agradecimientos – Acknowledgements Es realmente difícil hacer mención en pocas líneas de tantas personas que han recorrido este camino conmigo, haciendo posible el desarrollo de esta tesis. Sirvan estas palabras como muestra de mi gratitud hacia ellas. Me gustaría comenzar agradeciendo de forma muy especial la ayuda de mis directores de tesis, Profesor Dr. Alfonso San Miguel y Dr. Ramón Perea. Les agradezco enormemente la libertad y confianza que me han dado, apoyándome en todo momento en mis decisiones y animándome a elegir temas de investigación acordes con mis intereses científicos. Los admiro por su gran conocimiento académico y científico, pero más aún por su humildad y trato personal. Con ellos me he sentido siempre muy valorada. Ellos han sido mis maestros en el ámbito académico y personal, y no puedo imaginar mejores directores. A mi compañera de despacho, Aida, le agradezco el tiempo que me ha dedicado desde el principio de la tesis, siempre dispuesta a escuchar mis dudas y a orientarme. Admiro su actitud positiva ante la vida y me encanta tenerla de compañera de despacho. I would also like to give my sincere thanks for their warm hospitality to my supervisors during my research stays at the University of Stanford, Laval and Lyon. Thanks to Prof. Rodolfo Dirzo, Prof. Côté and Prof. Gaillard. It has been an honour to be able to join their labs and to learn directly from them. I really admire their work, which has inspired much of my research. I also thank the support of all lab members and friends I made during my research stays. We had really fun moments during the Friday’s happy hour at Stanford, the Thursday beers in Quebec and l’heure de l’apèro at Lyon and Chizé. Finally, thanks to all the students that helped me with laboratory, field and research work. Quiero dar las gracias a los directores de Los Quintos de Mora, Carlos Rodríguez y Ángel Moreno por apoyar en todo momento la continuidad de los estudios en la finca y por facilitar al máximo la recogida de muestras; a todo el ix personal y guardería de Los Quintos de Mora, y, en especial, a Carmen López (q.e.p.d.), Amanda, José Polo y Justi, gracias por su apoyo y ayuda en todo momento. It has also been a pleasure to have the opportunity to work with Prof. Kurt Bollmann and Dr. Maik Rehnus. I’m grateful for the opportunity to participate in the “Rehkitzmarkierung Schweiz” project and for thier help and support as co-authors. I also thank the help and support of Dr. Marco Heurich for his collaboration on this project. A Juan Carlos Peral, director del Centro Cinegético de Valsemana, le agradezco su generosidad al facilitarnos los valiosísimos datos que, con tanto esfuerzo y sacrificio, han recogido durante años, y a José Luis Álvarez su inestimable trabajo en la elaboración y estructuración de la base de datos. También quiero dar las gracias a Emmanuel Serrano y Santiago Lavín por su trabajo como co-autores de este estudio y por su hospitalidad (y cocina riquísima) durante nuestras salidas de campo. También deseo dar las gracias al equipo del Proyecto Aequilibrium que, aunque no está directamente relacionado con ningún capítulo de esta tesis, me ha ayudado mucho en otros aspectos de mi formación predoctoral. Gracias a Enrique Navarro, María Moreno, Pablo Ortega, Jonatan García, Javier Gamonal y Mario Bregaña. Gracias también a la Asociación del Corzo Español (ACE) por facilitar la toma de muestras durante las jornadas de descastes de corzas que, sin duda, ayudarán a conocer mejor el estado reproductivo de las hembras en la península. También deseo dar las gracias a todos mis compañeros y al personal de la Escuela de Montes; a mis compañeros de comedor, por contribuir a despejarme con risas y paseos a mediodía y a mis compañeros de doctorado, por los cafés y charlas sobre nuestras preocupaciones por la tesis; a Sonia Roig, por su apoyo y ayuda como coordinadora del programa de doctorado, y a Almudena Izquierdo, por su ayuda en los complejos trámites de la tesis. x Y, por supuesto, a mis padres a los que, junto con Rafa, dedico esta tesis, porque un trabajo de investigación es fruto también del apoyo que nos ofrecen las personas que nos quieren. A mi padre le agradezco haberme enseñado que, con constancia y trabajo, se puede conseguir casi todo lo que uno se proponga. A mi madre le doy las gracias por su compañía, cariño y cuidados durante las vacaciones en las que ella asumió las tareas domésticas que me correspondían para que yo pudiese centrarme en la tesis. Por último, a Rafa, sin duda, el que más me ha acompañado, sufriendo el día a día de esta tesis. A él le doy las gracias por su ayuda incondicional, así como por su paciencia y apoyo en los momentos de desánimo. Le doy las gracias también por haberlo dejado todo para venirse conmigo a cualquier lugar del mundo y, por supuesto, mil gracias por haberme iniciado en el mundo de la programación y por todo lo que me ha enseñado. Por toda su ayuda, este trabajo puede considerarse también suyo. Muchas gracias a todos xi Abstract Populations living at the edge of a species' range of distribution often experience sub-optimal conditions and different selective pressures than those located at more central locations. Thus, in the context of climate change, edge populations may provide key information to understand the evolution of species under harsher climatic conditions, helping to predict future demographic changes and distribution shifts. Our main goal was to assess the effect of environmental variation on the reproduction of red deer (Cervus elaphus) and roe deer (Capreolus capreolus) populations living at the edge of their range (i.e. Mediterranean and Alpine ecosystems). This thesis is structured in three chapters that contain 6 research studies. Chapter I analyzed how environmental conditions affected female reproductive timing of both species. Study 1 revealed that red deer displayed high phenotypic plasticity in conception dates in Mediterranean environments, being population traits and the effect of climatic conditions on plant productivity the most influential factors. Thus, dry springs with low plant productivity and high densities produced delayed conception dates. Studies 2 and 3 assessed the ability of roe deer parturition timing to track environmental changes along two different axes: space and time. While roe deer parturitions were delayed with increasing latitudes and elevations, they displayed little plasticity to rapidly adjust them to the increasingly earlier plant phenology of Alpine environments. Chapter II studied the variation of demographic parameters along geographical gradients. Thus, study 4 showed an increase in roe deer litter size with latitude but a decrease with elevation. This opposite relationship, known as the fecundity gradient paradox, has never been reported for a large mammal. Finally, Chapter III analyzed how environmental conditions and individual factors influenced the development of male secondary sexual traits in Mediterranean environments. Study 5 indicated xiii that population traits and food availability were the main factors influencing antler size in red deer, in line with Study 1. Finally, roe deer early-life investment on antlers was a reliable indicator of future adult performance (i.e.
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