Behavioral Ecology and Spawning Periodicity of the Gulf of California Grunion, Leuresthes Sardina
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Behavioral ecology and spawning periodicity of the Gulf of California grunion, Leuresthes sardina Item Type text; Dissertation-Reproduction (electronic) Authors Muench, Kevin Allen, 1941- 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 23/09/2021 20:26:50 Link to Item http://hdl.handle.net/10150/565393 BEHAVIORAL ECOLOGY AND SPAWNING PERIODICITY OF THE GULF OF CALIFORNIA GRUNION, LEURESTHES SARDINA by Kevin Allen Muench 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 1 9 7 7 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE I hereby recommend that this dissertation prepared under my direction by Kevin A11 en Muench_______________________________. entitled Behavioral Ecology and Spawning Periodicity of the Gulf of California Grunion, Leuresthes sardina_______________ be accepted as fulfilling the dissertation requirement of the degree of Doctor of Philosophy____________________ _____________ Dissertation Director Date After inspection of the final copy of the dissertation, the following members of the Final Examination Committee concur in its approval and recommend its acceptance:*’* '7~? 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 scholarship. In all other instances, however, permission must be obtained from the author. SIGNED: ACKNOWLEDGMENTS My introduction to the richness and beauty of the Gulf of California came from Don Thomson. We shared the excitement and mystery of our first grunion run in 1968, which stimulated this study. Don has provided aid, counsel, guidance, and interest throughout the study. His stimulus has contributed immeasurably to the completion of this work. I thank the multitude of persons who assisted me in various ways during grunion runs, especially Chuck Constant, Ernie Kehl, Bill Phebus, Bill Yarwood, Frank Tavernier, and those whose names may not be listed but are appreciated. Boyd Walker provided valuable comments during two Gulf grunion runs on the differences between the two species. Logistics for the beginning phase of this study were supported by ONR grant NR 104-897. Several close Mexican friends at El Golfo assisted my efforts, especially Juan Rodriguez and Emmanuel Carlos, who gave "Thunderpig" (my special research beach buggy) and I a place to stay, work, and socialize. The assistance and prodding from Judy and Allyson during the years of data collecting and analyzing kept me going, albeit intermit tently. Draft preparation could never have been done without the help and handwriting interpretation of my friend and typist, Paula Christy. My thanks to Chris van Dyck who did the illustrations, layout, coordi nated manuscript preparation, and assisted me in my absence from the campus. Tor Hansen provided the line drawing of the Gulf grunion. iii Valuable comments and special assistance were given me by my committee to enable me to finish the manuscript by the deadline. My special thanks to Don Thomson, Elisabeth Stull, Charles Ziebell, Robert Hoshaw, and Robert Mellor. I wish to dedicate this research to my mentor and friend, Don Thomson, who stimulated this first research on the Gulf grunion and all succeeding work which now makes him the world's authority on the Gulf grunion, Leuresthes sardina. TABLE OF CONTENTS Page LIST OF ILLUSTRATIONS ............................. vii LIST OF T A B L E S ...................................................... ix ABSTRACT ................................... x INTRODUCTION........................................................ 1 Literature Review K> .H rfr Scope of Study . Taxonomy of L_. sardina and L. tenuis Range ............................. Study Site ....................... METHODS AND MATERIALS .............................................. 12 R E S U L T S ................. ........... ............................. 15 Behavioral Ecology ............. 15 Movement to and from Spawning B e a c h e s ................... 15 Movement onto the Spawning Beach ......................... 18 The Spawning A c t .......................................... 22 Waves and Spawning B e h a v i o r ............................. 38 Periodicity of Spawning R u n s ................................. 40 Spawning Season .......................................... 40 Tidal Influence on Diel Spawning......................... 41 Duration of Runs .......................................... 44 Run S e r i e s ................................................ 53 Timing of Run S e r i e s ...................................... 54 Prediction of Gulf Grunion R u n s ......................... 59 Conservation of the Gulf G r u n i o n ............................. 60 DISCUSSION AND CONCLUSIONS.................... 65 Spawning Periodicity and Biological Clocks ................... 65 Spawning Behavior of the Gulf Grunion ......................... 68 Timing and Prediction.......................................... 70 APPENDIX A: TIDE HEIGHTS AND TIMES OF NIGHT GRUNION RUNS AFTER NEW MOON P H A S E ................ 72 v vi TABLE OF CONTENTS--Continued Page APPENDIX B: TIDE HEIGHTS AND TIMES OF NIGHT GRUNION RUNS AFTER FULL MOON P H A S E ....................................... 74 APPENDIX C: TIDE HEIGHTS AND TIMES OF DAY GRUNION RUNS AFTER NEW MOON P H A S E ...................................................... 76 APPENDIX D: TIDE HEIGHTS AND TIMES OF DAY GRUNION RUNS AFTER FULL MOON P H A S E .................................................. 78 APPENDIX E: SUMMARY OF ALL DATA ON GULF GRUNION RUNS AT EL GOLFO DE SANTA CLARA, SONORA, MEXICO, FROM 1968 TO 1973 USED IN COMPUTING THE MEAN VALUES IN TABLES 4 AND 5 ............... 80 LIST OF REFERENCES.................................................. 86 LIST OF ILLUSTRATIONS Figure Page 1. The Gulf grunion, Leuresthes sardina (Jenkins and Evermann, 1 8 8 8 ) .................................................... 5 2. Known distribution and range of the Gulf grunion, Leuresthes sardina .................................................. 6 3. Known spawning beaches of the Gulf grunion along the Sonoran coast (outlined in dark b a n d s ) ......................... 8 4. El Golfo beach showing megaripples......................... 10 5. Study beach showing grunion spawning area ................. 11 6. Daytime grunion run showing beach where fishhave spawned . 16 7. Scouts in the surf and on the b e a c h ....................... 19 8. A heavy daytime r u n .......... 21 9. People chasing grunion during a daytime r u n ............... 23 10. Night run showing male bumping female ..................... 24 11. Night run showing three erect females ..................... 25 12. Spawning fish under minimal surf c o n d i t i o n s ............... 27 13. Female grunion beginning to dig into the s a n d ............. 29 14. Grunion run under normal surf conditions................... 30 15. Females extruding eggs into s a n d ........................... 32 16. Female beginning to leave after egg laying ................. 33 17. Egg pod buried about 55 cm in the s a n d ..................... 34 18. Males encircling finger mimicking vibrating fe m a l e .......... 36 19. Sequence of behaviors in grunion spawning ................. 39 vii viii LIST OF ILLUSTRATIONS— Continued Figure Page 20. Times of the daily higher high tides (HHW) for the years 1967-1973 .......................................... 43 21. Observations made of Gulf grunion runs at El Golfo showing night, day, and transition between night and day runs . 45 22. Gulf grunion runs at El Golfo showing transition of higher high tides to daytime and beginning of daytime runs . 46 23. Scheme representing the time relations that are basic to the hypothesis that the spawning times of the California grunion depend upon the initiation of egg (or sperm) development at the time of the moon phases and on a constant period of development ......................... 55 24. Scheme representing the time relationships for predicting the spawning time of the Gulf g r u n i o n ................... 58 25. Fisherman capturing grunion during a day run ........ 61 26. Tire tracks on a grunion spawning b e a c h ................... 62 27. Stranded grunion attracted by lights ....................... 63 LIST OF TABLES Table Page 1. Number