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Behavioral Aspects of Western Gray Squirrel Ecology

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Behavioral Aspects of Western Gray Squirrel Ecology Behavioral aspects of western gray squirrel ecology Item Type text; Dissertation-Reproduction (electronic) Authors Cross, Stephen P. 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 11/10/2021 06:42:38 Link to Item http://hdl.handle.net/10150/565181 BEHAVIORAL ASPECTS OF WESTERN GRAY SQUIRREL ECOLOGY by Stephen Paul Cross A Dissertation Submitted to the Faculty of. the DEPARTMENT OF BIOLOGICAL SCIENCES 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 6 9 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE I hereby recommend that this dissertation prepared under my direction by St e p h e n Paul Cross_______________________________ entitled B E H A V I O R A L A S P E C T S OF W E S T E R N G RAY__________ S Q U I R R E L E C O LOGY___________________________________ be accepted as fulfilling the dissertation requirement of the degree of D O C T O R OF P H I L O S O P H Y_____________________________ 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:* 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 allow- able without special permission, provided that accurate acknowledgment, of source is made. Requests for permission for extended quotation from or reproduction of this manu­ script 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 X wish to thank' Dr. E. LendeiX Cockrum, for his advice and assistance throughout my graduate studies, Drs, Russell P, Davis, C. Roger Hungerford and Stephen . < M. Russell contributed helpful advice and read the manu­ script critically, for which I am most grateful. Drs, Franklin Sturges and Joseph Hall were very helpful during the initial stages of the project, Messrs, Terry Rahe, Charles Phillips, Richard Walker, and Dr. William B . Purdom, assisted me in various phases of field work and captive animal studies. Two of my sons, Stephen and Michael, accompanied me on several field trips and assisted in mapping vegeta­ tion, My wife, Betty, in addition to her much appreciated encouragement and understanding, helped with some field work and in the preparation of this manuscript. Most of the field work for this study was conducted while I was a member of the faculty of Southern Oregon . College, Ashland. I am indebted to several individuals at that institution allowing me to carry on the work and for providing equipment, facilities, and monetary assist- .ance. iii iv . Additional financial .support was provided by The. American Museum of Natural History ~ Theodore Roosevelt Memorial Fund, The Society of the Sigma Xif and Tektronix Foundation. Field work during the summer 1968 was partially supported through a NASA Predoctoral Traineeship. TABLE OF CONTENTS - _ ' - . Page LIST OF ILLUSTRATIONS v. , „ , , , . , yxi LIST OF TABLES . „ » . , : . , , . • « „ . ix ABSTRACT 0 9 9 « .? « * * « « « » » ? « «. * % * • « » '■ 5C INTRODUCTION . , , . , * . , . , . .1 METHODS AND MATERIALS . , , . • , ... ' 6 Description of Study Areas . , . , . , . ... 6 . Trapping and Marking . ... .. ... « , . 15 Radio Tracking Techniques , ... ... 17 Home Range Plotting . .......... 21 Observation and Experimentation . .. 2 4 POPULATIONS AND INTERSPECIFIC RELATIONS . .. 29 Population Size and Characteristics ..... , . 29 Interspecific Relations . .. .. , , . , . 34 Parasites and Disease . , , ... 34 Predation . , . , , , . .. 36 Competitors . , . , « . 38 DAILY ACTIVITY PATTERNS . , , , . 42 Onset of Activity . .. , , .. .• . 42 Diurnal Activity . .. , . 47 Cessation of . Activity . ' . , . , ■ 53 EGOCENTRIC BEHAVIOR 56 ■ Maintenance Behavior . , . , . ... 56 Investigation, Alarm and Escape, and Play ..... 60 ■ Feeding.Habits 69 ■ Nesting Habits , , . , . « ... 79 Nest Description 79 Nest Utilization , . , , . » . , . 87 REPRODUCTIVE BEHAVIOR . , . , , .' . , V , . 99 Female Cycle , . .. , .' . .- 99 .. Male Cycle . , . ... ... .. 101 v ■ Vi . TABLE OF CONTENTS (CONTINUED) '' ' Page I Mating Behavior ., , . 102 MOVEMENTS . , , . , , .. , » „ . , , . , . , , . 108 Home Range , . , , , , , . , » v v , . 109 Population Aspects . , , » . ■ .......... Ill Individual Variations . , . 118 Non-routine Movements . , . , .. , . .. 125 Ma1 1 ng » « % * « ^ ^ * $ « « » * * * * « 12 6 Feeding ... 126 Avoidance . « « . « . « . 12J. Dispersion and Exploration , , , , » ... , 129 SOCIAL BEHAVIOR « , . , 131 Communication and Agonistic Behavior 131 Social Hierarchy . , . , ... » , . , » . , » , 13 5 DISCUSSION AND CONCLUSIONS . .......... 145 LITERATURE C I T E D ......... .......................... .. 161 XjIST OF ILLUSTRATIONS. Figure . ,Page 1, Map of the vicinity of Ashland, Oregon showing areas mentioned in text 7 2, Temperature-moisture climogra'ph for Ashland, Oregon ........... .. , . 8 3, Map of the Baldy study area 10 4, Map of the Emigrant Lake study area , . » » . , 13 5, The Emigrant Lake study area (looking from east to west) » „ , 14 6 , Diagram of a typical radio unit used for tracking « , . 19 7, A feeder attached to a ponderosa pine 25 8, Western gray squirrel population trends for Jackson and Josephine counties, Oregon . , . , 32 9, Emergence times in relation to sunrise . « . , 43 10, Variation in activity after sunrise 48 11, Interactions at a feeder, as an index of activity ........... , . , 51 1 2 , Proportion.of squirrels in nests at time intervals before civil sunset , , . , . 54 13, Stretching posture of a squirrel descending a tree . , ...................................... 59 14, Typical posture assumed during loafing relaxation 61 15, Two variations in posture associated with alert relaxation . 62 16,' Attentive posture . 6 6 17, A tree-hole nest site 81 vii viii LIgT OF ILLUSTRATIONS (CONTINUED.) . Figure ; page 18, A stick nest .(top view) in a ponderosa pine , 8 6 19, Seasonal changfe in relative size of the ..birlbo-urethral gland in living squirrels . 103 2 0 , ■ :Home range determinations for eight Western gray squirrels at the Emigrant Lake study area during late winter- , early spring, 1967 . , . , , . , , , . , , 115 21, Home range plots for squirrels 8 and 23 during different time periods . .. , .. , . , 120 22, Home range plots for squirrel 14 during different time periods . , , , , . , , 1 2 1 23, Home range plots for squirrel 13 during different time periods ......... 1 2 2 24, Aggressive postures associated with agonistic behavior 133 25, Interactions between gray squirrels at the Emigrant Lake feeder 137 26, Interactions between gray squirrels art the Baldy feeder 13.8 LIST OF- TABLES ! Table Page 1* The number of gray squirrels inhabiting the permanent trap area of the Emigrant Lake study area at various trap periods during the study „ „ . , „ » . „ „ „ „ , , , , 31 2, Selected data concerning onset of activity of individual S'ciurus griseus at the Emi­ grant Lake study area , .. , . , . - 45 3„ Percent occurrence of foods utilized by j3. griseus in southern Oregon during the y ear « « « « « ■ « « * • • « « » « ? < « n « » '*7 0 4. Selected data concerning Western gray squirrel nest characteristics' . , , « , » , , 82 5. Nest utilization by some squirrels of the Emigrant Lake population , , . , , „ , , 90 6 . Nest utilization by three squirrels during nine days of a 15 day period ......... 95 7. Total recorded range for several individual squirrels in the Emigrant Lake population . 112 8 . Late: winter-spring home ranges of Western gray squirrels in the Emigrant Lake study area ........... .................... 114 9. Late summer, 1967, home ranges of Western gray squirrels from two study areas ........... 117 10. Late summer, 1966 and 1968, home ranges of Western gray squirrels in. the Emigrant Lake study area , . , , , , . , . , 119 11, Correlation of social rank and home range size of Western gray squirrels - 142 ix ABSTRACT The role of behavior of the Western gray squirrel, Sciurus griseus Ord, in relation to the basic ecology of the species was investigated during a 36 month period? near Ashland, Oregon. Data were collected regarding popular tion characteristics, daily activity patternsr egocentric behavior, reproductive behavior, movements and social be­ havior. Two separate populations of squirrels were studied, one of which was estimated to be at least twice as dense as the other. The higher population density probably resulted from the presence of a thicker and more continuous
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