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University Microfilms, Inc., Ann Arbor, Michigan the UNIVERSITY of OKLAHOMA This dissertation has been microfilmed exactly as received 68-14,206 QUANSTROM, Walter Roy, 1942- SOME ASPECTS OF THE ETHOECOLOGY OF RICHARDSON’S GROUND SQUIRREL IN EASTERN NORTH DAKOTA. The University of Oklahoma, Ph.D., 1968 Zoology University Microfilms, Inc., Ann Arbor, Michigan THE UNIVERSITY OF OKLAHOMA GRADUATE COLLEGE SOME ASPECTS OF THE ETHOECOLOGY OF RICHARDSON'S GROUND SQUIRREL IN EASTERN NORTH DAKOTA A DISSERTATION SUBMITTED TO THE GRADUATE FACULTY in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY BY WALTER ROY QUANSTROM Norman, Oklahoma 1968 SOME ASPECTS OF THE ETHOECOLOGY OF RICHARDSON'S GROUND SQQIRREL IN EASTERN NORTH DAKOTA APPROVED BY DISSERTATION COMMITTEE ACKNOWLEDGEMENTS Few tasks of any magnitude are accomplished without assistance. This one is no exception. Field studies by their nature require more than the usual amount of assistance. I am grateful to Mr. and Mrs. Teddy R. Erbele, Lehr, North Dakota; Mr. and Mrs. Chet Burley, Blanchard, North Dakota; Mr. and Mrs. Swede Johnson, Hunter, North Dakota; and Mr. and Mrs. Merton L. Sheldon, Mapleton, North Dakota, for use of their land while making this study. The Erbeles were especially kind and helpful during the two years we worked on their land. Mr. Merton L. Sheldon was willing to lend a hand in many ways, and provided a place to stay during the study. Mr. Sheldon's sensitivity to nature is re­ freshing. Mrs. Sheldon served meals many days to save more time for field work. I am grateful to the Department of Zoology, Uni­ versity of Oklahoma, and the National Aeronautics and Space Administration (N.A.S.A.) for financial help. \The research funds from N.A.S.A. made possible the filming many of the observations. I am indebted to Dr. Charles C. Carpenter, Professor of Zoology, for so many things that it would be most iii difficult, indeed, to list them all. Dr. Carpenter's love of all forms of animal life is deeply imbedded in my im­ pressions of him. One of the largest contributions to my graduate program that Dr. Carpenter has made is that I have always felt free to speak with him on any matter. Much of the basic concepts of this study were gained from Dr. Carpenter. I am also grateful to Dr. George Mitsch Sutton, Research Professor, not so much for actual help in this study but for teaching me the value of careful field observations. Dr. Sutton's constant attention to detail has helped me to strengthen a weak point. Professor Sutton's deep love for animals in their natural environment is something that I shall never forget. I am grateful to Drs. Charles C. Carpenter, Cluff E. Hopla, George J. Goodman, and Howard B. Haines for reading the manuscript and for making many helpful suggestions. I would also like to thank my wife, Fay, for her help in every aspect of this study. Without Fay's encouragement this study would not have been completed. IV TABLE OF CONTENTS Page ACKNOWLEDGEMENTS.................................... ill LIST OF TABLES...................................... vii LIST OF FIGURES................................. Ix FRONTISPIECE........................................ xi Chapter I. INTRODUCTION................................ 1 II. METHODS AND MATERIALS ...................... 6 III. THE ABIOTIC ENVIRONMENT.................... Geological History........................ 1^ Climate.................................. 16 IV. THE BIOTIC ENVIRONMENT...................... 21 V. BEHAVIOR .................................. 38 Growth and Pelage........................ 38 Yearly C y c l e s ............................ *+0 Emergence and Movement of Young Squirrels . ^8 Daily C y c l e .............................. 5l Posturing................................ 5^ L o c o m o t i o n .............................. 55 Burrows and Digging Behavior.............. 57 Tail Flicking............................ 64- Nest Building............................ 6M- Grooming.................................. 65 Elimination.............................. 66 Sexual Behavior .......................... 67 Parturition.............................. 72 Social Structure............. 7*+ Home Range and Territoriality............ 76 V Chapter Page Agonistic Behavior........................ 91 Food, Water, and Associated Behavior. 96 Vertebrate Associates .................... 107 VI. DISCUSSION.................................. Ill LITERATURE CITED.................................... 119 VI LIST OF TABLES Table Page 1. Maximum, minimum, and average air temperatures taken at Fargo (Hector Airport), North Dakota during 196$, 1966, 1967............. 17,18 2. Partial list of the plants found in associ­ ation with Richardson's ground squirrel in eastern North Dakota .................... 24-27 3. List of the species of amphibians found in association with Richardson's ground squirrel in eastern North Dakota........... 28 4. List of the species of reptiles found in association with Richardson's ground squirrel in eastern North Dakota........... 29 5. List of the species of birds found in association with Richardson's ground squirrel in eastern North Dakota............. 30-35 6. List of the mammals found in association with Richardson's ground squirrel in eastern North Dakota......................... 36,37 7. Weights and measurements of selected groups of Richardson's ground squirrels............ 42 8. Appearance of adult females above ground, July, 1965.................................. ^7 9. Number and per cent of total squirrels above ground at half hour intervals on June 8 and June 10, 1965.................... 53 10. Size and weight of Richardson's ground squirrel shortly after birth................ 75 Vll Table Page 11. Number of young Richardson's ground squirrels observed above ground for each female in 1965........................................ 86 12. Home ranges of Richardson's ground squirrel . 92 Vlll LIST OF FIGURES Figure Page 1. Layout of the Mapleton study area............ 10 2. Average weekly temperature at Fargo, North Dakota (Hector Airport) during 19&5, 1966, 1967 ........................................... 4l 3. Underground soil temperatures taken at Fargo, North Dakota........................ 4k 4. Movement of young ground squirrels from their home burrow.......................... 50 5. Identification kiss.......................... 52 6. Postures given by Richardson's ground squirrel.................................. 56 7. Sequence of positions assumed while running. 58 8. Sketch of an excavated burrow............ 61 9. Sequence of positions assumed while digging. 63 10. Sequence of positions assumed in the pre- copulatory run of the m a l e ................ 68 11. Sequence of positions assumed before and during copulation.......................... 70 ,7^ 12. Home range of dominant female, 1965......... 77 13. Home range of dominant female, 1966......... 78 14. Home range of female 16 during 1965 and 1966 . 79 15- Home range of female 7 during 1965 and 1966. 80 16. Home range of female 13 during 1965 and 1966 . 8l IX Figure Page 17. Home range of female 10 during 1965 and 1966. 82 18. Home range of female 1^ during 1965 and 1966. 83 19. Home range of female h during 1965 and 1966 8^- 20. Home range and territory of female 11, 1965 87 21. Home range and territory of female 6, 1965. 89 22. Home ranges of all females, 1965.......... 90 23. Sequence of positions assumed while chasing 93 2h. Postures assumed after chase.............. 9^ 25. Face off; Submission-aggression posture . 95 26. Boxing; Three squirrel interaction........ 97 X ■RftWOHR Km t z ~ SOME ASPECTS OF THE ETHOECOLOGY OF RICHARDSON'S GROUND SQUIRREL IN EASTERN NORTH DAKOTA CHAPTER I INTRODUCTION From the time of Aristotle, who thought that every animal had its unique slot to fill and that the degree of perfection of a thing could be measured by how well it fit into this slot, man has recognized some basic zoological concepts of a particular animal's adaptation to its environ­ ment. These early natural historians recognized many obvious examples of these relationships such as gills for breathing in water, and lungs for breathing in air. Much present day research, still concerned with these adaptations, unearths some incredulously subtle animal-environmental relationships. It can be noted that in the past biologists have been primarily concerned with morphological adaptations. Such things as the rigid tail feathers of a woodpecker, or the stiff hair fringe on the hindfoot of a water shrew are pointed out as examples of structures which enable an animal to adapt to a particular environment. More recently, how­ ever, studies show that animals have evolved the ability to 2 use (i.e., behavior patterns) its morphology in ways that are advantageous. These behavior patterns must compliment the morphological structures in order for the organism to be successful. The associated behavior pattern is just as im­ portant as the morphological structure that it compliments, and the more specific the structure the more specific its be­ havioral pattern is liable to be. The American Avocet, Recurvirostra americana, with its unique bill structure, uses it to obtain food in a very specific manner; however, the Virginia opossum, Didelphis marsupialis. is a generalized animal and its feeding behavior is quite variable. A mature behavior pattern that compliments a spe­ cific anatomy can be accomplished by at least two means; an inherited behavior characteristic or behavior modified by experience. The type of behavior attempted, and also the amount of time spent performing a motor pattern
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