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University Microfilms International 300 N The Breeding Biology Of The Puffins: Tufted Puffin (Lunda Cirrhata), Horned Puffin (Fratercula Corniculata), Common Puffin (F. Arctica), And Rhinoceros Auklet (Cerorhinca Monocerata) Item Type Thesis Authors Wehle, Duff Henry Strong Download date 09/10/2021 09:57:35 Link to Item http://hdl.handle.net/11122/9312 INFORMATION TO USERS This was produced from a copy of a document sent to us for microfilming. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the material subm itted. The following explanation of techniques is provided to help you understand markings or notations which may appear on this reproduction. 1. Trie sign or “ target” fo r pages apparently lacking from the docum ent photographed is “Missing Page(s)”. If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure you of complete continuity. 2. When an image on the film is obliterated with a round black mark it is an indication that the film inspector noticed either blurred copy because of movement during exposure, or duplicate copy. Unless we meant to delete copyrighted materials that should not have been filmed, you will find a good image of the page in the adjacent frame. If copyrighted materials were deleted you will find a target note listing the pages in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photo­ graphed the photographer has followed a definite method in “sectioning” the material. It is customary to begin filming at the upper left hand corner of a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again -beginning below the first row and continuing on until complete. 4. For any illustrations that cannot be reproduced satisfactorily by xerography, photographic prints can be purchased at additional cost and tipped into your xerographic copy. Requests can be made to our Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In all cases we have filmed the best available copy. University Microfilms International 300 N. ZEEB RD.. ANN ARBOR. Ml 48106 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Wehle, D uff Henry Strong THE BREEDING BIOLOGY OF THE PUFFINS: TUFTED PUFFIN (LUNDA CIRRHATA), HORNED PUFFIN (FRATERCULA CORNICULATA), COMMON PUFFIN (F. ARCTICA), AND RHINOCEROS AUKLET (CERORHINCA MONOCERATA) University o f Alaska Ph.D. 1980 University Microfilms International 300 N. Zeeb Road. Ann Arbor, MI 48106 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE BREEDING BIOLOGY OF THE PUFFINS: TUFTED PUFFIN (LUNDA CIRRHATA) , HORNED PUFFIN ( FRATERCULA CORNICULATA), COMMON PUFFIN (F. ARCT1CA), AND RHINOCEROS AUKLET ( CERORHINCA MONOCERATA) A THESIS Presented to the Faculty of the University of Alaska in partial fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY By Duff Henry Strong Wehle, B .S ., M.S. Fairbanks, Alaska December 1980 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE BREEDING BIOLOGY OF THE PUFFINS: TUFTED PUFFIN (LUNDA CIRRHATA), HORNED PUFFIN (FRATERCULA CQRNICULATA) , COMMON PUFFIN (F_. ARCTICA) . AND RHINOCEROS AUKLET (CERORHINCA MONOCERATA) RECOMMENDED: . AoA)LX. \ ) Chairman, Advisory Committee Program I Vice Chancellor for ResBar’Ch arid Advanced Study iu Date Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT The natural histories of Tufted and Horned puffins ( Lunda cirrhata and Fratercula corniculata) were studied during the summer on Buldir Island, Alaska, in 1975 and on Ugaiushak Island, Alaska, in 1976 and 1977. Data from these and other recent studies have been presented and compared with that available for Common Puffins (F . arctica) and Rhinoceros Auklets (Cerorhinca monocerata) —actually a misnamed puffin—to provide synthesis of the natural history of the world's four species of puffins. Different aspects of puffin natural history show varying degrees of interspecific variability. Preferences of nesting habitats and patterns of colony settlement are similar for Tufted and Common puffins, while the length of incubation, brooding, and nestling periods are similar for Tufted Puffins and Rhinoceros Auklets. Sexual and social behaviors are similar for the congeneric Horned and Common puffins. All species have two to four vocaliza­ tions in common. Nest building, nest-site tenacity, nest-site cohab­ itation, territoriality, egg replacement, and the participation by both sexes in incubation and feeding of young are characteristic of all puffins. For all colony-years reported, the average range of breeding success rates for puffins are 50-60% for laying success, 75-90% for hatching success, and 53-82% for fledging success. Puffins exhibit considerable seasonal and geographic inter-and intraspecific variation in their foraging habitats. Fish is the most important prey for all adult puffins, although squid, polychaetes, and crustaceans are con- 3 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 4 sumed to varying degrees by each species. Sand lance ( Ammodytes s p p .) is the most common prey fed to all puffin nestlings. Nestling growth rates are highest in Tufted and Horned puffins when sand lance are supplemented with Capelin ( Mallotus villosus) and in Common Puffins when sand lance are supplemented with sprats ( Sprattus sprattus) . Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS The field research for this dissertation was supported, in part, by the U .S. Fish and Wildlife Service through the Aleutian Islands National Wildlife Refuge and the Office of Biological Services--Coastal Ecosystems. I am grateful to the following people and organizations for their assistance in various phases of this study: Dr. O. H. Hewitt and Dr. A. Borror, Shoals Marine Laboratory, Portsmouth, New Hampshire, for instilling in me the interest in and appreciation for seabirds which ultimately led to this study. The members of my doctoral committee, Dr. R. D. Guthrie, Dr. S. F. MacLean, J r., Dr. C. P. McRoy, and Dr. G. C. West for their advice and constructive criticism and especially Dr. Brina Kessel, my advisor, for her continued guidance and encouragement throughout this study. R. H. Day, B. E. Lawhead, K. D. Powers, E. P. Hoberg, G. V. Byrd, J. L. Trapp, D. L. Woolington, E. B. Rhode, and others who assisted me in the field. The Aquatir Collections, University of Alaska Museum, and par­ ticularly K. O. Coyle, G. E. Keiser, G. Mueller, and K. Vogt for identification of puffin prey items. The Terrestrial Vertebrate Collections, University of Alaska Museum, Fairbanks, for providing specimens and information used in connection with this study. 5 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 6 M. J. Amaral; R. E. Ashcroft; P. A. Baird; G. Burrell; A. R. DeGange; G. j. Divoky; Z. Eppley; D. A. Frazer; D. D. Gibson; R. Gill, J r.; P. J. Gould; M. P. Harris; S. A. Hatch; G. L. Hunt, Jr.; L. L. Leschner; D. A. Manuwal; R. A. Moe; D. A. Nysewander; G. A. Sanger; A. L. Sowls; K. Taylor; J. L. Trapp; and K. Vermeer, who kindly provided me with unpublished information used in this dis­ sertation. B. Hunt, R. Hunt, S. Watson, and N. Yamamoto, for assistance in the illustrations, graphics, and typing of this dissertation. Fellow graduate students and friends, especially R. H. Day, B. E. Lawhead, D. L. Musgrave, S. Paulsen, T. R. Seastedt, F. C. Seyfert,. and P. Tryon, for their help and encouragement in various aspects of this study. The Institute of Terrestrial Ecology, Banchory, for logistic sup­ port in Scotland. Dr. M. P. Harris, Institute of Terrestrial Ecology, Banchory, for guidance, encouragement, and constructive criticism in the preparation of this dissertation and Dr. M. P. Harris and K. Taylor, St. Andrews University, St. Andrews, for their generosity and hospitality extended during my stay in Scotland. Finally, I dedicate this dissertation to my wife Sara, who typed its initial drafts and who has provided me with constant encouragement, understanding, and assistance through the long months of writing. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS Page L IS T OF FIG U RES ....................................................................................... 11 LIST OF TABLES ....................................................................................... 12 INTRODUCTION .................................................................................... 16 CHAPTER I: COMPARATIVE BREEDING BIOLOGY AND FEEDING ECOLOGY OF TUFTED AND HORNED PUFFINS, LUNDA CIRRHATA AND FRATERCULA CORNICULATA .... 20 NESTING HABITAT ............................................................... 20 THE PRE-EGG STAGE ..........................................................
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