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U·M·I University Mlcroflims International a Bell & Howell Information Company 300 North Zeeb Road Population biology of Octopus digueti and the morphology of American tropical octopods. Item Type text; Dissertation-Reproduction (electronic) Authors Voight, Janet Ruth. Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 04/10/2021 02:11:27 Link to Item http://hdl.handle.net/10150/185018 INFORMATION TO USERS The most advanced technology has been used to photograph and reproduce this manuscript from the microfilm master. UMI films the . text directly from the original or copy submitted. Thus, some thesis and dissertation copies are. in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. U·M·I University Mlcroflims International A Bell & Howell Information Company 300 North Zeeb Road. Ann Arbor. MI 48106·1346 USA 313 761·4700 800 521·0600 Order Number 9024636 Population biology of Octopus digueti and the morphology of American tropical octopods Voight, Janet Ruth, Ph.D. The University of Arizona, 1990 U·M·I 300 N. Zeeb Rd. Ann Arbor, MI 48106 POPULATION BIOLOGY OF OCTOPUS DIGUETI AND THE MORPHOLOGY OF AMERICAN TROPICAL OCTOPODS by Janet Ruth Voight A Dissertation Submitted to the Faculty of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 1990 ----~-.,--.~.------.-------~- ~------.- THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE 2 As members of the Final Examination Committee, we certify that we have read the dissertation prepared by __~T~auD~e~t~B~J!~t~h~~V~oui~g~h~t _______________________ entitled population B:jology of Octopus diglleti and the Morpholo~y of American Tropical Octopods and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor Of Phil osophy /3 IlJO':"; I ~~o Date Date Date Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. Dissertation Director Date 3 STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at the University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the library. Brief quotations from this dissertation are allowable without special permission, provided that accurate acknowledgement of source is made. Reuqests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author. 4 ACKNOWLEDGMENTS My research has benefitted from the guidance of three major professors; J.R Hendrickson; D.A Thomson and RE. Strauss. I thank each for his unique contributions to my work. I also thank O.G. Ward, K. W. FIessa, and M.J. Donoghue for their continued assistance and guidance as members of my committee and for their friendship. Space constraints prevent me from thanking each individual who has contributed to my work; my apologies to those not included. RT. Hanlon and colleagues at the University of Texas Marine Biomedical Institute provided vital assistance, especially early on. The staff of Centro D'estudios Desiertos y Oceanos (CEDO) made laboratory space and housing available without which, my field research would have been impossible. At the University of Arizona, the assistance of Rich Strauss has contributed greatly to the success of virtually every aspect of this research. Chapters 2, 3, and 4 exist only because of his help. Working with Rich as simply been the best, most productive phase of my scientific career. Throughout this research, Meriel Brooks has also provided invaluable assistance in diverse areas. In addition, she has helped me through times of utter despair and helped successes. Thanks Meriel. P.A Hastings, M.J. Porter, and G. Bharathan revealed to me the mysteries of reconstructing phylogenies. Michael Donoghue provided training and discussion which were essential to my work. In addition, cephalopod scientists have been extremely supportive, especially RB. Toll who taught me standard dissection and measurement techniques. Discussions with E.B. Hartwick, J.A Mather, R. Ambrose, K. Mangold, S.v. Boletzky, RK. O'Dor, M. Nixon RT. Hanlon and G.L. Voss, University of Miami, have offered what has been vital encouragement. W.M. Kier and A Smith, University of North Carolina, Chapel HilI, permitted me to cite unpublished work that is critical to understanding octopod evolution, and suggested major improvements to Chapter 4. I thank the curators who have allowed me to use collections under their care; W. Miller, University of Arizona; RT. Hanlon, University of Texas; c.F.E. Roper, United States National Museum; G.M. Davis, Academy of Natural Science, Philadelphia and K Boss, Museum of Comparative Zoology, Harvard. I thank the photographers who have documented specimens for me or provided me with otherwise unattainable photographs, especially, A Kerstitch; C. Schwalbe and J. Forsythe and R Aronson; F. Cipriano; R Hanlon; A Harvey; S. McLaughlin; R Lindner; A Rodaniche; S. Shuster; S. Sutter; and D.A Thomson. I thank Shawn McLaughlin not only for expert assistance in word processing, photography and figure preparation, but also for his emotional support, and most of all, his friendship. Financial assistance for research and travel was generously provided by the Hawaiian Malacological Society, Conchologists of America, and the Department of Ecology & Evolutionary Biology, University of Arizona. OC,topuses were collected under Mexican permits #0475 and 2999 issued to J.R Hendrickson in 1984-1985. This dissertation is dedicated to the memory of Gilbert L. Voss. Only because of the completeness of his biogeographic, taxonomic and systematic research have I been able to make the scientific advances reported here. 5 TABLE OF CONTENTS LIST OF ILLUSTRATIONS ................................... 9 LIST OF TABLES . .. 10 ABSTRACT .............................................. 11 GENERAL INTRODUCTION ................................ 13 CHAPTER ONE THE BIOLOGY OF A SEMI-ANNUAL ANIMAL: A YEAR-LONG STUDY OF Octopus digueti ................... 16 Summary. .. 16 Introduction ......................................... , 17 Materials and Methods ............................ 18 Study Area ................................ 18 Study Organism . .. 19 Field Techniques ........................... , 20 Laboratory Techniques ....................... 21 Data Analysis .............................. 23 Results ............................................. 25 Movement . .. 27 Predation ...................................... 29 Shelter Usage ................................... 31 Sex Ratio ...................................... 31 Definition of Age classes ............... .. 33 Growth ....................................... , 33 6 Table of Contents (Continued) Reproduction and Life span. .. 38 Discussion ........................... .. 41 Movement ...................................... 41 Predation ...................................... 43 Shell Usage . .. 45 Sex Ratios . .. 46 Age class definition ............................... 47 Growth ........................................ 53 Reproduction and Life span . .. 57 CHAPTER TWO VARIATION IN OcrOPOD SPECIMENS: REASSESSING THE MORPHOLOGY OF SOFf-BODIED ORGANISMS ............ 60 Summary ....................................... 60 Introduction .................................... 60 Materials and Methods ............................ 62 Results and Discussion ............................ 68 CHAPTER THREE AMERICAN TROPICAL SHALLOW-WATER OcrOPODINAE: A MORPHOLOGICAL STUDY ......................... .. 84 Summary ....................................... 84 Introduction .................................... 84 Materials and Methods ...........................
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