Xerox University Microfilms 74-17,752
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Xerox University Microfilms 74-17,752 BOERaa, P. Dee, 1946- THE GALAPAGOS PENGUIN: A STUDY OF ADAPTATIONS FOR LIFE IN AN UNPREDICTABLE ENVIRONNENT. The Ohio State University, Ph.D., 1974 Ecology University Microfilms, A XEROX C o m p a n y , An n Arbor, Michigan © Copyright by P. Dse Boersnia 1974 THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. Tl-iE GALAPAGOS PENGUIN: A STUDY OF ADAPTATIONS FOR LIFE IN AN UNPREDICTABLE ENVIRONMENT DISSERTATION Presented in Partial Fulfillment of the Requirements for Tlie Degree Doctor of Pliilosophy in the Graduate School of The Chic State. University P. Dee Boersma, B.S. Hie Ohio State University 1974 Reading Coimnittee: Approved B y Paul A. Colinvaux Jerry F, Dov.nhower Rodger D. Mitchell Department of Zoology ACKNOWLEDGEMENTS I am deeply indebted to Dr. Paul Colinvaux for guidance and encouragement throughout this study. During my years as a graduate student, his knowledge, dedication and support were invaluable. I am also indebted to Drs. Jerry Downhower, Sheldon Lustick, Rodger Mitchell and Tony Peterle for their ideas, criticism and friendship. Madhukar Golhar provided statistical advice and the data were analyzed at The Ohio State University Computer and Information Science Center. This research was financed by the National Science Foundation under Grant GB-2906.SX, Principal Investigator, Dr. Paul Colinvaux. The author was supported by a University Fellowship during portions of the research. Personnel and scientists at the Charles Darwin Station, Lina A, Golden Cachalot and many residents of the Galapagos Islands provided assistance during this research. David Day, Dwayne Maxwell and Godfrey Merlin made general observations on penguin breeding and surface water temperatures while I was not in the islands. Nancy Jo, Chris Kjolhede and Sally Cloninger accompanied me as field assistants at different times. Without their deter mination and effort this study would not have been possible. Tlie duPont family took Sally Cloninger and myself on their yacht to Elizabeth Bay, Isabela and deserve special thanks. I am also grateful to the many people who either live or have passed through the Galapagos Islands and contributed to this research. VITA November 1, 1946... Born-Mt. Pleasant, Michigan 1968, 1969........ Spring field work, Research Assistant, Central Michigan University Museimi. Collecting and study of Florida birds. 1969.............. B.Sc., Cum Laude, Central Michigan University 1969.............. Board of Trustee Scholarship, Central Michigan University 1969.............. UTio's UTio in American Colleges and Universities 1969.............. National College Registary 1969.............. Member, Presidential Task Force on Women's Rights and Responsibjlities 1969-197 3......... University Fellow, T]ie Ohio State University, Columbus, Ohio 1970-197 6 ......... Trustee, Central Michigan University 1970-1974......... Teaching and Research Associate, The Ohio State University, Columbus, Ohio 1970, 1971, 1972... Summer field work. The Galapagos Islands, Ecuador 1971 .............. IVho's IVho in American Politics 197 2 .............. Winter field work. The Galapagos Islands, Ecuador 197 3.............. Representative to United States of American Department of State National Foreign Policy Conference 1973.............. United States Advisor to United Nation Commission of the Status of Women VITA CONTINUED CONTRIBUTIONS AND PUBLICATIONS A Matter of Simple Justice. U.S. Government Printing 1-33 (collective authorship). In press: Adaptations of Galapagos penguins for life in two different environments. ^ B. Stonehouse, The Biology of the Penguin. Macmillan, London. Adaptations of the Galapagos penguin to an unpredictable environment. Paper presented October 9, 1973 at the American Ornithologists' Union. The Ecologist and the Galapagos Penguin. 24 min., color 16 imm film, Department of Photography and Cinema, The Ohio State University, Production Associate. Breeding Strategies of Seabirds in the Galapagos Islands, A.A.A.S. Galapagos Symposium, February, 1974. TABLE OF CONTENTS Page ACKNOWLEDGEMENTS................................................ ü VITAE ........... iv LIST OF T A B L E S ..............................................viii LIST OF FIGURES............................................. x LIST OF P L A T E S ..............................................xiii INTRODUCTION ................................................ 1 PROCEDURE...................................................... 21 NATIJR.U HISTORY OF THE GALAPAGOS PENGUIN........................ 30 Distribution and Abundance ............................... 30 Beliavior.................................................. 43 The daily routine.................................... 43 Courtship behavior ................................ 44 Head movement........................................ 48 Pecking.............................................. 53 P r e e n i n g ............................................ 54 L o c o motion.......................................... 55 Vocal responses...................................... 61 Nesting.............................................. 65 Sexual and Age Dimorphism in Galapagos Penguins............. 66 Plumage......................................... 66 S i z e ................................................ 70 Movement and Mortality ................................... 82 Movement of males and females........................ 86 Movement of y o u n g .................................... 87 TABLE ÜL CONTENTS (CONTINUED) Breeding Activity ...................................... 90 Pair bonding...................................... 90 E g g s .............................................. 91 Incubation........................................ 9A C h icks............................................ 94 Synchrony in egg laying and hatching.............. 109 Frequency of breeding.............................. 110 Molting.................................................115 F o o d .................................................. 125 Predation and D e a t h .................................... 129 Breeding Seasons........................................ 132 Breeding Success ..................................... 133 ECOLOGY OF GALAPAGOS PENGUINS.................................135 Food as the Limiting Factor on the Population.......... 135 Further Test of the Mypotliesis that Food Limits the Population...........................................149 Advantages of Sexual and Age Dimorphism.................164 Maximizing Fitness and Minimizing the Cost of Reproduction ....................................... 172 M o l t i n g ......................................... 172 Pair b o n d s ....................................... 175 Synchrony......................................... 176 Egg shape......................................... 177 Incubation....................................... 179 Asynchrony in hatching ........................... 180 A clutch size of t w o ...............................181 Reproductive cost m o d e l .................. 186 CONCLUSION: THE STRATEGIES OF GALAPAGOS PENGUINS AND THEIR RELATIONSHIP TO OTHER ORGANISMS .................. 1G6 PLATES ...................................................... 203 LITER.ATURE C I T E D ............................................ 211 LIST OF TABLES Table Page 1 Three expected characteristics of organisms which live in unpredictable and predictable environments. ^ 2 English and official names of Galapagos Islands. • • • 10 3 Surface water temperatures at Pta. Espinosa, Fernandina, Cabo Douglas, Fernandina and Elizabeth Bay, Isabela between July