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Of American Crocodiles Observed (Spotlight Surveys) and Captured (Mark-Recapture) by Size Class and Age Group, Respectively, Across 2013 on Coiba Island, Panama

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Of American Crocodiles Observed (Spotlight Surveys) and Captured (Mark-Recapture) by Size Class and Age Group, Respectively, Across 2013 on Coiba Island, Panama Natural History of the American crocodile in a Tropical Pacific Island in Panama, Central America by Sergio A. Balaguera Reina B.S. A Dissertation In Biological Sciences Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Approved Llewellyn D. Densmore III Chair of Committee Nancy McIntyre Richard Strauss Richard Stevens David Ray Mark Sheridan Dean of the Graduate School May, 2018 Copyright 2018, Sergio A. Balaguera-Reina Texas Tech University, Sergio A. Balaguera-Reina, May 2018 ACKNOWLEDGMENTS I want to express my gratitude to my academic advisor Dr. Llewellyn D. Densmore III for his continuous support and guidance during my studies in the Biological Sciences program at Texas Tech University. I would like to extend this gratitude to the members of my academic advisory committee Drs. Nancy McIntyre, Richard Strauss, David Ray, and Richard Stevens for their support and encouragement to pursue my academic goals. Their guidance and critical minds help me to shape and structure my scientific path and grow-up both personally and intellectually. The last five years at Texas Tech University have been a great experience at both personal and professional levels. It is why I want to express my gratitude to the chair of the department Dr. Ron Chesser as well as all professors I have worked under as a teacher assistant Drs. Robert Bradley, Rhonda Boros, Michael Dini, Richard Strauss, Lou Densmore, and Nicte Ordoñez-Garza. I also want to thank to Judy Edwards, Lanita Ladd, Lisa Dion, Pat Moore, Raquel Miranda, Kunju Li, Carol Espinosa, Vina Khan, and Christopher Long for all your support and help through this time. My gratitude to all my friends and colleagues who I met and shared time with at Texas Tech University, but especially to Stephen Roussos, Mike Sager, Ashish Bashyal, Elizabeth Mack, Brandon Gross, Austin Osmanski, Mark Lee, Andrés Rodríguez-Cordero, Betzaida Rivera, Ryan Vazquez, Abbie Ince, Juan Pablo Carrera, Nicte Ordoñez-Garza, Cibele Sotero, Diana Vargas, Pablo Tovar, Paulina Vélez, and Diana Palacios. I want to acknowledge the support of the Colombian government specially the Colciencias Institute who awarded me a scholarship to study abroad as well as the Colfuturo Institute staff for their help and administrative support. I would like to extend this acknowledge to the Panamanian Environmental Agency (ANAM), the Panamanian National Secretariat of Science, Technology, and Innovation, the Smithsonian Tropical Research Institute (STRI), the AERONAVAL, and the IUCN/SSC/Crocodile Specialist Group for their economical and logistical support to carry out the field part of this dissertation. I want to thank to Dr. Miryam Venegas- ii Texas Tech University, Sergio A. Balaguera-Reina, May 2018 Anaya for her support during this project as well as all the undergraduates and local people who helped me collect data in the field, especially to Betzaida Rivera, Jhon Gaitan, Tatiana Rincon, Tatiana Niño, Valeria Beltran, Alejandra Cristancho, Alejandro Morales, Italo Arbeláez, and Andrés Sánchez. Finally, I want to recognize the unconditional help and support of my beloved girlfriend Lorena, my father José Vicente, mother María Pureza, brother Diego Armando, and sister Magda Ximena, as well as cousins, uncles, aunts, grandparents and close friends that walked this long path with me. This work is dedicated to all my friends that could not see this dream come through, your names will be never forgotten. “Carpe diem, quam minimum credula postero” Horace 23 (BC) “We have the duty of formulating, of summarizing, and of communicating our conclusions, in intelligible form, in recognition of the right of other free minds to utilize them in making their own decisions” Ronald Fisher iii Texas Tech University, Sergio A. Balaguera-Reina, May 2018 ABSTRACT Conservation of large predators has long been a challenge for biologists due to the limited amount of information we have about their ecology and generally low numbers in the wild. The last decade has made it clear that to implement sound conservation practices, we must increase our understanding of large predators’ ecology, covering all possible aspects. I investigated the natural history of the American crocodile (Crocodylus acutus) on Coiba Island, Panama, addressing a comprehensive ecological framework including four specific topics: crocodile reproductive, population, spatial, and trophic ecology. As part of this work, I also developed two novel approaches focused to identify American crocodiles at the individual level based on the dorsal scute pattern (individual identification pattern recognition IIPR) and to estimate population sizes (using spotlight data) based on the sampling distribution (via bootstrapping) with adjusted confidence intervals. American crocodile reproductive ecology works as a primary driver of the spatial patterns found on Coiba Island, which are also highly influenced by environmental conditions such as precipitation. Hierarchization based on age group and sex through space and time was identified in the study area. Dietary niche overlap analysis showed clear ontogenetic dietary partitioning among juveniles, subadults, and adults. However, the general pattern postulated for crocodylians (small individuals eating invertebrates and adults feeding on larger prey) was not supported by the data collected from Coiba Island, indicating that even adult American crocodiles dwelling in coastal areas rely on more on easy-to-catch/abundant prey such as crustaceans. These results have important implications for the way we see the species as a functional entity of the community, including its interactions and more complex roles in the system than thought before. Overall, American crocodile life history varies considerably from insular to mainland populations and from coastal to in-land populations, demonstrating how “flexible” C. acutus is in terms of habitat requirements. Data suggest that realized niches in the insular populations “shrink” compared to mainland populations due to a reduced availability of resources and an inherent increase of intra and interspecific competition. Results from the present study allow future management and iv Texas Tech University, Sergio A. Balaguera-Reina, May 2018 conservation planning to be based on the comprehensive integration of information on the ecology of a Neotropical crocodylians. Key words: Crocodylians; nesting ecology; population ecology; spatial ecology; trophic ecology. v Texas Tech University, Sergio A. Balaguera-Reina, May 2018 TABLE OF CONTENTS ACKNOWLEDGMENTS ........................................................................ ii ABSTRACT .............................................................................................. iv TABLE OF CONTENTS ......................................................................... vi LIST OF TABLES ................................................................................... xi LIST OF FIGURES ............................................................................... xiv CHAPTER I ............................................................................................... 1 INTRODUCTION ..................................................................................... 1 CROCODYLUS ACUTUS IN PANAMA: A STATUS REPORT ...... 1 OBJECTIVES ....................................................................................... 6 ORGANIZATION OF CHAPTERS ..................................................... 7 CHAPTER II ............................................................................................. 8 REPRODUCTIVE ECOLOGY ............................................................... 8 INTRODUCTION ................................................................................ 8 MATERIALS AND METHODS ........................................................ 10 RESULTS ........................................................................................... 13 DISCUSSION ..................................................................................... 24 CHAPTER III ......................................................................................... 32 POPULATION ECOLOGY ................................................................... 32 INTRODUCTION .............................................................................. 32 MATERIALS AND METHODS ........................................................ 35 Study area ...................................................................................... 35 Data collection .............................................................................. 35 Pre-sampling ................................................................................. 36 Sampling ....................................................................................... 38 vi Texas Tech University, Sergio A. Balaguera-Reina, May 2018 Spotlight surveys data analyses ..................................................... 39 Mark-recapture data analyses ........................................................ 40 RESULTS ........................................................................................... 41 Spotlight survey ............................................................................ 42 Mark-recapture .............................................................................. 43 DISCUSSION ..................................................................................... 47 Source of bias in population
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