Pontificia Universidad Católica De Chile Facultad De Ciencias Biológicas Programa De Doctorado En Ciencias Biológicas Mención Ecología
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Pontificia Universidad Católica de Chile Facultad de Ciencias Biológicas Programa de Doctorado en Ciencias Biológicas Mención Ecología TESIS DOCTORAL: LIVING IN THE FAST LANE: FORAGING ECOLOGY OF THE ANTARCTIC FUR SEAL (Arctocephalus gazella) AT THE EDGE OF THEIR BREEDING DISTRIBUTION Por RENATO FRANCISCO BORRÁS CHÁVEZ Diciembre 2020 Pontificia Universidad Católica de Chile Facultad de Ciencias Biológicas Programa de Doctorado en Ciencias Biológicas Mención Ecología TESIS DOCTORAL: LIVING IN THE FAST LANE: FORAGING ECOLOGY OF THE ANTARCTIC FUR SEAL (Arctocephalus gazella) AT THE EDGE OF THEIR BREEDING DISTRIBUTION Tesis presentada a la Pontificia Universidad Católica de Chile como parte de los requisitos para optar al grado de Doctor en Ciencias Biológicas mención Ecología Por RENATO FRANCISCO BORRÁS CHÁVEZ Director de Tesis: Dr. José Miguel Fariña R. Co-Director de Tesis: Dr. Francisco Bozinovic K. Comisión de Tesis: Dr. Luis Ebensperger P. Dr. Mauricio Lima A. Dr. Pablo Sabat K. Diciembre 2020 LA DEFENSA FINAL DE LA TESIS DOCTORAL TITULADA “LIVING IN THE FAST LANE: FORAGING ECOLOGY OF THE ANTARCTIC FUR SEAL (Arctocephalus gazella) AT THE EDGE OF THEIR BREEDING DISTRIBUTION” Presentada por el Candidato a Doctor en Ciencias Biológicas Mención Ecología de la Pontificia Universidad Católica de Chile SR. RENATO FRANCISCO BORRÁS CHÁVEZ Ha sido aprobada por el Tribunal Examinador, constituido por los profesores abajo firmantes, calificándose el trabajo realizado, el manuscrito sometido y la defensa oral con nota ___7.0_____ (Siete) DR. MAURICIO LIMA A. DR. JUAN A. CORREA M. Coordinador Comité de Tesis Decano Facultad de Ciencias Biológicas-UC Facultad de Ciencias Biológicas-UC DR. FRANCISCO BOZINOVIC K. DR. JOSÉ MIGUEL FARIÑA R. Co-Director de Tesis Director de Tesis Facultad de Ciencias Biológicas-UC Facultad de Ciencias Biológicas-UC ` DR. LUIS EBENSPERGER P. DR. PABLO SABAT K. Miembro del Comité de Tesis Miembro Externo del Comité de Tesis Facultad de Ciencias Biológicas-UC Universidad de Chile Santiago, 26 de noviembre de 2020.- LIVING IN THE FAST LANE: FORAGING ECOLOGY OF THE ANTARCTIC FUR SEAL (ARCTOCEPHALUS GAZELLA) AT THE EDGE OF THEIR BREEDING DISTRIBUTION 6 DEDICATORIA No hay nadie en el mundo que haya celebrado más mis logros académicos que mi familia, y nadie tan presente en el arduo camino que implica la obtención de este grado que mi madre y padre. La presente tesis está dedicada a: Miguel Borrás Geisse y Sandra Chávez Ordenes. Quien soy hoy, es fiel reflejo de su incondicional amor y apoyo. También dedico esta tesis a mi sobrino, Miguel Borrás Cerda. Por el sacrificio que implicó para mí alejarme tantos meses de tu lado en tus primeros años, tiempo congelado en los parajes Antárticos que nunca recuperaré… 7 ACKNOWNLEDGMENTS I first want to thank to my Ad-honorem advisor dr. Michael Goebel for the huge opportunity given when he included me as a PhD student on the pinniped program. Dr Goebel played a key role in the construction development and execution of the pinniped monitoring program of the Antarctic Marine Living Resources division (AMLR) of the National Oceanic and Atmospheric Administration (NOAA) and was the leader of this program at Cape Shirreff for more than 20 years including the years that this thesis was conducted. “Mike” was a great mentor in this process. He was the most qualified person in the understanding of both the species and the ecosystem studied in this thesis, especially at this location. He also managed to get the funding and support necessary to conduct this research through the AMLR program. We have become friends along the way and for all of this I’m grateful with him. I would like to thank the AMLR program of NOAA for their funding and logistic support. I’m especially grateful with its director, Dr. George Watters, for allowing me to conduct my research within the program. I also want to thank all the field assistants hired by NOAA during these years: Kevin Pietrzak, Mckenzie Mudge, Tony Kroeger, Wiley Archibald, Whitney Taylor, Sam Woodman, Jesse Senzer, and especially, Dr. Douglas Krause, the new leader of the pinniped program. Furthermore, I would like to thank those who assisted me at the AMLR-NOAA laboratory facilities in San Diego, USA: Jen Walsh, Becky Nitschelm and Korrine Ettl. I want to thank the Chilean Antarctic Institute (INACH) for the funding INACH- DT_02_15 that allowed me to do my field work at the Guillermo Mann refuge, the purchase of some instruments and the hiring of two assistants to whom I’m deeply grateful for their dedication; Camila Vargas and Eduardo Fuentes. Also, I appreciate the laboratory assistance provided at the Catholic University by Daniela Oryan. I also want to thank those that silently worked very hard for us to be safe and sound while on Antarctica: The INACH crew involved in the 8 administrative process, especially Wendy Rubio, leader or the expedition department, and the Chilean air forces and Army. I thank the Vicerectoría de Investigación of the Pontificia Universidad Católica de Chile and “The State of the Antarctic Ecosystem (AntEco)” program for funding a fellowship and a conference participation respectively. I want to thank the “Capital Humano Avanzado” program and the PhD scholarship ANID N°: 21130059 that allowed me to get my degree. I am grateful to the Center of Applied Ecology and Sustainability (CAPES) for the opportunity to carry out this research and to ANID PIA/BASAL FB0002 for funding. I’m also grateful with every single co-author of the upcoming publications for their priceless contribution to this study: M. E. Goebel, S. Villegas-Amtmann, L.A. Hückstädt, C. Rivera-Rebella, D. P. Costa, S. M. Budge, A. Guerrero, C.J. van Calcar, F. Santa Cruz, L. Krüger, C. A. Cárdenas and especially my two advisors for trusting me and for their constant support: Dr. Francisco Bozinovic and el Dr. José Miguel Fariña. Also, I want to thank Catalina Fuentealba for creating the fur seal cover illustration, and the artist that created the illustrations of each chapter, Alejandra “Momo” Ramírez, who I have admired all my life and has participated in this thesis from digging poop (scats analysis) to revise my presentations. I appreciate the help of reviewers that provided insights on early draft of the manuscripts, research proposal and helped with the methodology: Paulina Möller, Leah Segui, Theresa Keates, and Sarah Kienle. To my dear 2013 PhD generation for many battles fought together: Bryan Bularz, Brian Latorre, Magdalena Fuentealba, Belen Gallardo, and Francisco Díaz. A special mention to my beautiful friends and family (mis negros, Pete,Magda and my great siblings, especially my sister, Alejandra Borrás, who also worked in this document), they have walked with me and support every decision along my academic journey. Finally, I would like to thank Caroline. Ik wil Caroline bedanken. Voor alle ondersteuning, ondanks onze afstand, mijn stemmingswisselingen en op het laatst ook mijn scriptie stress. Alleen jij en ik weten hoe leuk en mooi onze reis samen is. This manuscript would not have been possible if any of these pieces of the puzzle would have been missing, and for this, I am deeply grateful to all of them. 9 10 LIST OF CONTENTS DEDICATORIA _____________________________________________________________________ 6 ACKNOWNLEDGMENTS _____________________________________________________________ 7 LIST OF CONTENTS ________________________________________________________________ 10 LIST OF FIGURES __________________________________________________________________ 14 LIST OF TABLES ___________________________________________________________________ 18 LIST OF ABBREVIATIONS ____________________________________________________________ 21 RESUMEN _______________________________________________________________________ 22 ABSTRACT _______________________________________________________________________ 25 INTRODUCTION __________________________________________________________________ 28 FACTORS SHAPING FORAGING BEHAVIOR ____________________________________________________ 29 MEASURING FORAGING BEHAVIOR IN MARINE MAMMALS_________________________________________ 31 OTARIIDS FORAGING BEHAVIOR __________________________________________________________ 33 THE ANTARCTIC FUR SEAL. AN EXTENSIVELY STUDIED OTARIID ______________________________________ 34 PROJECT OUTLINE, RESEARCH OBJECTIVE, AND HYPOTHESES _______________________________________ 36 GENERAL METHODOLOGY __________________________________________________________ 39 STUDY SITE AND LOGISTICS _____________________________________________________________ 39 ANIMAL CHARACTERISTICS ______________________________________________________________ 41 CAPTURE PROTOCOL _________________________________________________________________ 42 INSTRUMENT CONFIGURATION AND DATA PROCESSING __________________________________________ 44 DAY AND NIGHT DIVING DIFFERENCES ______________________________________________________ 44 DIVING EFFORT _____________________________________________________________________ 45 11 CHAPTER 1. TIME AND BEHAVIORAL CONSTRAINTS OF LACTATION IN HIGHLY HETEROGENOUS ENVIRONMENTS ______________________________ 47 INTRODUCTION __________________________________________________________________ 52 MATERIALS AND METHODS _________________________________________________________ 55 FORAGING EFFORT ___________________________________________________________________ 56 STATISTICAL ANALYSIS _________________________________________________________________ 58 RESULTS ________________________________________________________________________