FISHERY BIOLOGY AND FEEDING ECOLOGY OF RAYS IN BAHÍA ALMEJAS, MEXICO A thesis submitted to the faculty of San Francisco State University In partial fulfillment of The requirements for The degree Master of Science In Marine Science by Joseph John Bizzarro San Francisco, California December, 2005 Copyright by Joseph John Bizzarro 2005 ACKNOWLEDGMENTS I consider my thesis committee to be the greatest ever assembled for my subject matter. Dr. Enric Cortés’ advice was especially valuable in the design and analysis of my third chapter (feeding ecology). Dr. Lara Ferry-Graham has an incredible knack for organization, presentation, and pragmatism. She was very helpful throughout the analysis and editing process and provided me with myriad ideas and critical comments. Dr. Ralph Larson is an esteemed ecologist and a great editor. His comments greatly improved this document. Dr. Gregor M. Cailliet, my advisor and friend, is someone for whom I have a great deal of respect. He is an example of all that is good in science and provided me with countless helpful suggestions and opportunities during my protracted tenure in his ichthyology laboratory. This study would not have been possible without the considerable assistance of several friends and colleagues. Wade D. Smith provided unwavering moral and logistic support throughout the duration of my enrollment at Moss Landing Marine Laboratories (MLML). He is a great friend, a passionate and dedicated scientist, and an incomparable field biologist. The support and companionship of Annie Beesley and Paul Collins during 1999 fieldwork were greatly appreciated. Carolina Downton Hoffmann and students of Dr. Carlos Villavicencio-Garayzar’s Laboratorio de los Elasmobranquios at the Universidad Autónoma de Baja California Sur and Dr. Felipe Galván-Magaña and his students at CICIMAR were essential to the coordination of survey trips and the attainment and storage of supplies and samples. Several MLML faculty and staff members were also integral to the completion of my M.Sc. degree. Joan Parker and the library staff provided me with a substantial amount of literature, much of which was obscure and/or foreign, in a timely manner and never restricted my numerous requests. Special thanks to Kathleen Baker and Donna Kline for helping me navigate the SFSU system and graduate without any major dilemmas. They were magicians. Dr. Gary Greene has employed me for almost a decade and helped me to develop a second-career in habitat mapping and associated fish/invertebrate studies. He has become a good friend and also has provided me with many opportunities that piqued my scientific interests and put money in my pockets. Dr. Dave Ebert allowed me the time and flexibility to complete my thesis while working for the Pacific Shark Research Center, and for that I am very grateful. I decided on a career in marine ecology during a term at the Williams College- Mystic Seaport Program. Dr. Jim Carlton was the program director and his passion, dedication, and superlative tutelage in the classroom and laboratory have been an inspiration to me throughout my life. Dr. Janie Wulff and the other students on that program also contributed to create an incredible working and social environment that I envy to this day. I would also like to thank Dr. John J. Gilbert, who served as my undergraduate thesis advisor at Dartmouth College and who hosted the marine biology portion of a tropical ecology program that I participated in and treasured during my senior year. Finally, I would like to thank my family and friends, who mean more to me than anything else. My parents taught me the importance of education at an early age and always encouraged me to pursue what I enjoyed most in my studies and my career. I cannot fully express how much their support and love has meant to me over the years. My father (Joseph B. Bizzarro) is an avid outdoorsman and my appreciation of the natural world was fostered through the time I spent with him “in the woods”, often at our vacation home in the rural Catskill Mountains of upstate New York. My mom (Bernadette M. Bizzarro) is an extremely caring and sensitive woman who is always there with a hug and a kind word. When I think of her, it makes me smile. My brother, (Dr. Matthew J. Bizzarro) is one of the most disciplined people that I have ever met and combines that trait with intelligence and pragmatism. His “way” is truly inspirational to me and I thought of him long and often during the writing of this document. Stori Oates has made everything in my life better and I love her dearly. She is also a great editor who helped me with my thesis much more than should be expected of any girlfriend. I’ve enjoyed the company of many good friends during my time in graduate school, especially: Erik Cordes, Jim McCarthy, Erica Burton, Jason Cope, Lisa Kerr, Matt Levey, Jeff Field, Eric Sandoval, and brother Wade. I greatly appreciated and benefited from Mason Dean’s inquisitive nature and especially our correspondence re: Narcine. He is destined to become a monster in the field and I am looking forward to watching it happen. I was lucky enough to have several old friends living in the area during my graduate work, including: Chris Schin, Andrew Harris, Andy Peay, and Ben Wang. Correspondence with Chris Maloney kept me sane, made me laugh, and provided much needed non-scientific distractions; Mr. Moneschaegn, I salute you. Paul Marashlian is my A.C. Cowlings in times of crisis and my Dean Martin when times are good. We are long-term survivors of the west coast experiment. Funding for this project was provided by the: David and Lucile Packard Foundation, Homeland Foundation, JiJi Foundation, California Sea Grant College System, PADI Project Aware, General Services Foundation, World Wildlife Fund, Christensen Fund, Moss Landing Marine Laboratories, Mote Marine Laboratory, Instituto Nacional de la Pesca, Dr. Earl H. and Ethyl M. Myers Oceanographic and Marine Biology Trust, and National Oceanic and Atmospheric Administration/National Marine Fisheries Service (to the National Shark Research Consortium, Pacific Shark Research Center). My parents provided fortuitous supplemental funding throughout the duration of this study. This project was conducted in accordance with and under the approval of the SFSU Committee for the Protection of Human and Animal Subjects Protocol #02-030. TABLE OF CONTENTS List of Tables. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi List of Figures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xx List of Appendices. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxxvi VOLUME 1 ⎯ CHAPTER 1 AND CHAPTER 2 CHAPTER 1 ⎯ Physical and biological characteristics of the Bahía Magdalena Lagoon Complex (Baja California Sur, Mexico), with comments on elasmobranch faunal composition and nursery utilization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 Results and Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Physical Characteristics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Biological Characteristics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 Fisheries. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Elasmobranchs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 CHAPTER 2 ⎯ The artisanal ray fishery of Bahía Almejas (Baja California Sur, Mexico). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .115 Field Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .115 Data Analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .117 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 Artisanal Fishing Camps and Related Activities. . . . . . . . . . . . . . . . . . . . . 120 Catch Composition – Puerto Viejo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .122 Catch Composition – Additional Camps. . . . . . . . . . . . . . . . . . . . . . . . . . . 126 Sex Ratios. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .127 Size Composition. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 Morphometric Relationships. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 Biomass and Numerical Landings Estimates. . . . . . . . . . . . . . . . . . . . . . . .137 Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165 Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166 VOLUME 2 ⎯ CHAPTER 3 CHAPTER 3 ⎯ Diet, foraging ecology, and trophic relationships of a ray assemblage from Bahía Almejas (Baja California Sur, Mexico). . . . . . . . . . . . . . . . . . . . . . . . . . . . 226 Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228 Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .236 Data Collection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .236 Data Analyses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .241 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 Shrinkage Experiment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 Rhinobatos productus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .252 Narcine entemedor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .259 Dasyatis dipterura. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 Gymnura marmorata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .272 Rhinoptera steindachneri. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .278 Interspecific Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278 Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Diet Composition and Comparison with Congeners. . . . . . . . . . . . . . . . . .282 Dietary Overlap – Intraspecific Comparisons. . . . . . . . . . . . . . . . . . . . . . .293 Dietary Overlap – Interspecific Comparisons. . . . . . . . . . . . . . . . . . . . . . .299 Trophic Ecology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303 Conclusions and Recommendations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .308 Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .311 Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .313 LIST OF TABLES Table Page CHAPTER 1 1. Grain size (mm), classification, sorting, and dominant clast of sediment samples collected in Bahía Almejas during August 1999. Sample collection locations are displayed in Fig. 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 2. Systematic list of elasmobranchs noted within the Bahía Magdalena lagoon complex, Baja California Sur, Mexico. Species occurrences summarized from: Castro-Aguirre and Torres-Orozco, 1993; Cruz-Agüero et al., 1994; Villavicencio-Garayzar and Abitita-Cárdenas, 1994; Galván-Magaña et al., 2000; and personal observations. Taxonomy follows Compagno (1999). General eastern North Pacific distribution categorized as follows: Tropical (TR), Subtropical (STR), Temperate (TM), and Widespread (WD). . . . . . . . . . . . . . . 94
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