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Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 I I SPACING MECHANISMS IN A HERMIT CRAB POPULATION Item Type text; Dissertation-Reproduction (electronic) Authors Snyder-Conn, Elaine Kay, 1948- 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 03/10/2021 22:24:01 Link to Item http://hdl.handle.net/10150/289616 INFORMATION TO USERS This material was produced from a microfilm copy of the original document. 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 original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document 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 thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 I i 77-26,961 SNYDER-CONN, Elaine Kay, 1948- SPACING MECHANISMS IN A HERMIT CRAB POPULATION. The University of Arizona, Ph.D., 1977 Zoology Xerox University Microfilms, Ann Arbor, Michigan 48106 SPACING MECHANISMS IN A HERMIT CRAB POPULATION by Elaine Kay Snyder-Conn 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 WITH A MAJOR IN ZOOLOGY In the Graduate College THE UNIVERSITY OF ARIZONA 19 7 7 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE I hereby recommend that this dissertation prepared under my direction by Elaine Kay Snyder-Corm entitled Spacing Mechanisms in a Hermit Crab Population be accepted as fulfilling the dissertation requirement of the degree of Doctor of Philosophy !Q 71 Dissertation Director Date After inspection of the final copy of the dissertation, the follovzing members of the Final E:camination Committee concur in its approval and recommend its acceptance:"" /6 77 Ma-,.-21 \1 77 3, /f 7~~7 ~ JL~*. 7 m? This approval and acceptance is contingent on the candidate's adequate performance and defense of this dissertation at the final oral examination. The inclusion of this sheet bound into the library copy of the dissertation is evidence of satisfactory performance at the final examination. 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 acknowledgment of source is made. Requests for permission for extended quotation from or reproduc­ tion 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 judgment the proposed use of the material is in the interests of scholar­ ship. In all other instances, however, permission must be obtained from the author. SIGNED: - Clcrvyo-^ ACKNOWLEDGMENTS Foremost, I thank Dr. Elisabeth Stull and Dr. Donald Thomson for encouraging me in this study, for conferring creative and useful advice, and for their thorough attention to manuscripts evolving from this study. Dr. Peter Pickens aided in the identification of hermit crabs species and also assisted in the experimental design of some of the rhythm studies. Dr. Ronald Pulliam also provided assistance in the experimental design and statistical treatment of the results, as well as providing able criticism of the manuscript. Mr. John Skeoch of the Villa School assisted in many of the experiments described in the final chapter. Dr. Robert Hoshaw gave valuable time in reading this manu­ script. Dr. Brian Hazlett and Dr. Ernst Reese graciously supplied unpublished data and ideas, in addition to stimulating my greater interest in the biology of hermit crabs. Numerous other researchers contributed unpublished observations to Chapter 3, including Drs. Richard Vance, William Grant, Jr., Elliot Norse, and Eldon Ball. In addition, Dr. Ball has contributed many research ideas to this study. Dr. Robert Kuehl suggested several of the statistical procedures followed in this study. Lastly, I would especially like to thank my husband, Mr. Jeffery Conn, for his aid in both research and writing phases of this study, and forbearance throughout this study. iii TABLE OF CONTENTS Page LIST OF TABLES vi LIST OF ILLUSTRATIONS viii ABSTRACT x 1. INTRODUCTION 1 Presentation of the Problem ..... 1 Theoretical Aspects of Spacing 2 Synopsis of the Research Approach 10 2. PATTERNS OF AGGREGATION AMONG MARINE CRUSTACEA 11 Introduction 11 Patches 11 Schools 13 Migratory Assemblages 15 Stratifications 22 Swarms and Clusters 24 Swarms 25 Clusters 33 Summary 58 3. SPACING IN HERMIT CRABS 61 Introduction 61 Clustering Species 61 Types of Clusters 69 Composition of Clusters 71 Size Composition 71 Species Composition 71 Environmental Variables 72 Depth 72 Substrate 72 Summary 73 iv V TABLE OF CONTENTS--Continued Page 4. AGGREGATION AND DISPERSION IN CLIBANARIUS DIGUETI: MECHANISMS 74 Introduction 74 Behavior in Nature 75 Sequential Diving Observations at Puerto Peflasco ... 75 Sequential Diving Observations at Bahfa San Carlos . 78 Transect Studies . • 79 Materials and Methods 79 Clustering Patterns within Transects 83 Clustering Patterns between Transects 84 Rhythmical Behavior under Constant Laboratory Conditions . 90 Activity Rhythms 90 Skototactic Rhythms 94 Phototactic Rhythm at Puerto Pefiasco 97 Rhythm of Gregariousness at Puerto Pefiasco 100 The Role of Rhythmical Behavior in Aggregation and Dispersal 101 Behavior in Response to the Environment 102 Components of the Rhythmically Changing Environment . 102 Components of the Static Physical Environment .... 126 Components of the Biotic Environment 129 Discussion and Conclusions 140 5. THE ADAPTIVE VALUE OF AGGREGATION IN CLIBANARIUS DIGUETI .... 146 Hypotheses 146 Shell Exchange 146 Mating 151 Shell Cleaning 152 Predator Avoidance 153 Resistance to Current 155 Resistance to Cold 155 Resistance to Desiccation 159 Discussion and Conclusions 164 APPENDIX: HERMIT CRAB VERSUS SHELL SIZE 169 REFERENCES 171 LIST OF TABLES Table Page 1. Types of Contagion Found in Marine Crustaceans 59 2. Spacing Patterns of Marine Hermit Crabs in Relation to Habitat Preference and Body Size 62 3. Measures of Aggregation 83 4. Basic Statistics and Indices of Clumping for Clibanarius digueti in Each Coquina Transect 85 5. Basic Statistics and Indices of Clumping for Clibanarius digueti in Each Boulder Transect 86 6. Temperatures during the Light Intensity Experiment 105 7. Response of Clibanarius digueti to Changes in Temperature . 106 8. Response of Clibanarius digueti to Changes of Current Direction 112 9. Basic Statistics and Indices of Clumping for All Hermit Crab Species Combined in Each Boulder Transect 132 10. Frequencies.of Shell Occupancy by Four Species of Hermit Crabs in Coquina and Boulder Transects 138 11. Results of One-Way Analyses of Variance in Size of Clibanarius digueti between and within Meters for Each Transect 148 12. Frequency of Shell Exchanges in Clustered versus Dispersed Clibanarius digueti 150 13. Taction and Mean Group Size of 16 Clibanarius digueti under Cold Conditions 156 14. Resistance to Cold by Clibanarius digueti in Large Clusters . 158 15. Resistance of Clibanarius digueti in Small Clusters to Desiccation 161 vi vii LIST OF TABLES--Continued Table Page 16. Resistance of Clibanarius digueti in Large Clusters to Desiccation 163 LIST OF ILLUSTRATIONS Figure Page 1. In Situ Pattern of Behavior of C_. digueti in the Puerto Peflasco Intertidal Zone 77 2. In Situ Pattern of Behavior of £. digueti in the Bahfa San Carlos Intertidal Zone 80 3. Clumping of Clibanarius digueti in Boulder and Coquina Transects by Month, as Indicated by the Variance to Mean Ratio 87 4. Clumping of Clibanarius digueti in Boulder and Coquina Transects by Month, as Indicated by Morisita's Index of Dispersion 89 5. Tidal Rhythm of Activity of Clibanarius digueti at Puerto Peflasco, Sonora, Mexico 92 6. Tidal Rhythm of Activity of Clibanarius digueti at Bahfa San Carlos, Sonora, Mexico 93 7. Tidal Rhythm of Skototaxis of CIibanarius digueti at Puerto Peflasco, Sonora, Mexico 96 8. Rhythm of Skototaxis of Clibanarius digueti at Bahfa San Carlos, Sonora, Mexico 98 9. Tidal Rhythm of Phototaxis of Clibanarius digueti at Puerto Peflasco, Sonora, Mexico 99 10. Locomotor Activity of Clibanarius digueti in Response to Different Light Intensities at Periods of High and Low Tide 104 11.
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