ECOLOGICAL FACTORS, PLEIOTROPY, AND THE EVOLUTION OF SEXUAL DIMORPHISM IN CHERNETID PSEUDOSCORPIONS (PHORESY, QUANTITATIVE GENETICS, SEXUAL SELECTION). Item Type text; Dissertation-Reproduction (electronic) Authors ZEH, DAVID WAYNE. 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 01/10/2021 14:50:45 Link to Item http://hdl.handle.net/10150/183995 INFORMATION TO USERS While the most advanced technology has been used to photograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. For example: • Manuscript pages may have indistinct print. In such cases, the best available copy has been filmed. • Manuscripts may not always be complete. In such cases, a note will indicate that it is not possible to obtain missing pages. • Copyrighted nlaterial may have been removed from the manuscript. In such cases, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, and charts) are photographed by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or as a 17"x 23" black and white photographic print. Most photographs reproduce acceptably on positive microfilm or microfiche but lack the clarity on xerographic copies made from the microfilm. For an additional charge, 35mm slides of 6"x 9" black and white photographic prints are available for any photographs or illustrations that cannot be reproduced satisfactorily by xerography. -_.. _-_ ... _-- 8708576 Zeh, David Wayne ECOLOGICAL FACTORS, PLEIOTROPY, AND THE EVOLUTION OF SEXUAL DIMORPHISM IN CHERNETID PSEUDOSCORPIONS The University of Arizona PH.D. 1986 University Microfilms International 300 N. Zeeb Road, Ann Arbor, MI48106 ---_.--_._-- PLEASE NOTE: In all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark_-I_. 1. Glossy photographs or pages ~ 2. Colored illustrations, paper or print ___ 3. Photographs with dark background ~ 4. Illustrations are poor copy ___ 5. Pages with black marks, not original copy __ 6. Print shows through as there is t~xt on both sides of page ___ 7. Indistinct, broken or small print on several pages ~ 8. Print exceeds margin requirements __ 9. Tightly bound copy with print lost in spine ___ 10. Computer printout pages with indistinct print ___ 11. Page(s) lacking when material received, and not available from school or author. 12. Page(s) seem to be missing in numbering only as text follows. 13. Two pages numbered . Text follows. 14. Curling and wrinkled pages __ 15. Dissertation contains pages with print at a slant, filmed as received / 16. Other--------------------------------- University Microfilms International ._--_._-_. __ .. _- - -----_._--.--_.- - • ECOLOGICAL FACTORS, PLEIOTROPY, AND THE EVOLUTION OF SEXUAL DIMORPHISM IN CHERNETID PSEUDOSCORPIONS by David Wayne Zeh 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 ARIZGNA 1 986 -------------------- THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by __D~av~i~d_W~a~yn~e~Z~eh~ _____________________________ entitled -------------------------------------------------------------------Ecological Factors, Pleiotropy, and the Evolution of Sexual Dimorphism in Cherne tid Pseudo scorpions and recommend that it be accepted as fulfilling the dissertation requirement Doctor of Philosophy for the Degree of ---------------------------------------------------------- /I ~, 17';r~ Date Date Date ~L!~,- Date 'O/t/7 a --.~ Date 7 7 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. ,-------.---,---- STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment 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. Request 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. SIGNED: ACKNOWLEDGMENTS I wish to thank S.J. Arnold, J.H. Brown, V. Delesalle, L. Delph-Lively, D.J. Futuyma, JoR. Hendrickson, A. Kodric-Brown, C.M. Lively, M. McGee, W.B. Muchmore, S. Nelson Jr., W.L. Nutting, S.K. Sakaluk, R.L. Smith, R. Hensley Thomas, P. Weygoldt, and especially Jeanne Anne Zeh for critically commenting on the manuscript. C.C. Hoff, W.B. Muchmore, S. Nelson Jr., and P. Weygoldt provided considerable advice, reprints and/or unpublished data. W.B. Muchmore confirmed the identification of ~. arizonensis. David Robinson kindly recorded data on the last few replications of the phoresy experiment. I thank the staff at Saguaro National Monument for permission to collect in the Tucson Mountain Unit. I am most grateful to J.R. Hendrickson for encouraging me to embark on a doctoral thesis. His support continued despite my shift in research from marine ecology to sexual selection in pseudoscorpions, considered by the uninformed to constitute a most inconsequential group of organisms. I was privileged to write this dissertation in the company of Jeanne Zeh, who provided considerable companionship and intellectual guidance. Her iii ----._.. _---_._-------_... _----- iv comments on all aspects of my research greatly improved the clarity and impact of this dissertation. Finally, research grants from the National Academy of Sciences (Henry Joseph Henry Fund), Sigma Xi, and the University of Arizona Graduate College (Graduate Student Development Fund; Summer Support Fund) contributed much appreciated financial aid. --- -_._--_._._------------- TABLE OF CONTENTS Page LIST 'OF TABLES vi i LIST OF ILLUSTRATIONS • ix ABSTRACT xi 1. SEXUAL SELECTION: AN OVERVIEW 1 2. AGGRESSION, DENSITY AND SEXUAL DIMORPHISM IN CHERNETID PSEUDOSCORPIONS ••• 5 Materials and Methods 11 Literature Analysis of North American Chernetidae • • • • • ••••• 13 Experiments with Dinocheirus arizonensis 15 Results ••••..•••••••.••.•. 19 Literature Analysis of North American Chernetidae • • • • • • • • • • • • • 19 Experiments with Dinocheirusarizonensis 25 Discussion • • • • • • • • • • • • • • • • 31 3. AN EXPERIMENTAL STUDY OF FEMALE SEXUAL RECEPTIVITY 41 Materials and Methods ••• ••• •• 45 Mating Experience and Female Receptivity 45 Spermatophore Deposition Rate and Female Receptivity ••••••••.•• 49 Results 49 Mating Experience and Female Receptivity 49 Spermatophore Deposition Rate and Female Receptivity ••••••• 51 Discussion • • • • • • • • • • • 54 v vi TABLE OF CONTENTS--Continued Page 4. THE PSEUDOSCORPION, THE SAGUARO AND THE FLY: PHORETIC DISPERSAL IN A DESERT ARACHNID 69 Materials and Methods •••••••••• 74 Experiment 1: Do Pseudoscorpions Preferentially Prey on or Attach to Newly Eclosed Neriids? •••••••• 75 Experiment 2: Do Gender and Social Context Influence the Incidence of Phoresy and Predation? • • • • • • • • • • • • 76 Colonization Study: Do Colonization and Dispersal Patterns Reflect Gender Differences in Phoretic Behavior? 77 Results • • • • • • • •••••••••• 80 Experiment 1: Do Pseudoscorpions Preferentially Prey on or Attach to Newly Eclosed Neriids? •••••••• 80 Experiment 2: Do Gender and Social Context Influence the Incidence of Phoresy and Predation? ••••••••••• 80 Colonization Study: Do Colonization and Dispersal Patterns Reflect Gender Differences in Phoretic Behavior? 83 Discussion • • • • • • 87 5. LIFE HISTORY CONSEQUENCES OF SEXUAL DIMORPHISM IN A CHERNETID PSEUDOSCORPION 96 Materials and Methods 100 Experimental Populations ••• 100 Experimental Design • • • • • • • • •• 101 Statistical Analysis • • • • • • • • • 103 Results • • • • • • • • • •• 104 Somatic Allocation Comparison. 104 Developmental Period/Heritability Study 106 Discussion • • • • • 112 APPENDIX 1: LITERATURE DATA ON SEXUAL DIMORPHISM IN CHERNETIDAE • • • • • • • • • • • • • 119 APPENDIX 2: LITERATURE DATA ON SEXUAL DIMORPHISM IN CHEIRIDIOIDEA • • • • • • • • • • • • • • • 122 LIST OF REFERENCES 123 ----------- LIST OF TABLES Page Table 1. Comparison between Males and Females for Carapace Length and Chela Silhouette Area among North American Species of Chernetidae • • • • •. • • • • 12 2. Comparison of Generic Means for Sexual Dimorphism • • •• • • • • 23 3. Summary of Stepwise Linear Regression