Resource Partitioning and Reproductive Success of Three
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AN ABSTRACT OF THE THESIS OF Marcy Jean Cottrell for the degree of Master of Science in Wildlife Science presented on October 2, 1981 Title: Resource Partitioning and Reproductive Success of Three Species of Hawks (Butp"p) in an-Oreqç,Prairie Signature redacted for privacy. Abstract approved_ Dr. Charles E. Warren 2) In a native prairie of Oregon (363 red-tailed (Buteo jamaicensis), ferruginous (Buteo regalis), and Swainson's hawks (Buteo swainsoni) coexist in one of the highest nesting Buteo densities in North 2merica. In 1979 and 1980, the distribution, density, and productivity of the three species was studied in re- lation to availability of food and nesting habitat. The degree of ecological segregation among the three species was also investigated. Red-tailed and ferruginous hawks showed considerable overlap in the timing of their nesting cycles; the Swainson's hawk nested about one month later in the spring. The major food item of all three species was the Belding ground squirrel (Spennophilus beld- ingi). The total dietary overlap among the three hawk species was high: in 1979 and 1980, red-tailed and ferruginous hawks overlapped 90 percent and 80 percent, red-tailed and Swainson's hawks 88 percent and 83 percent, and ferruginous and Swainson's hawks 83 percent and 95 percent. Although timing and use of food resources exhibited considerable overlap, habitat was partitioned. Nine major habitat types occur- red within the study area. Red-tailed hawks had almost exclusive use of pine groves and cultivated areas with cottonwood trees. Fer- ruginous hawks had almost exclusive use of south exposure sites where large rocky outcrops occurred.Use of other habitat types overlapped to some degree. The total habitat overlap between the red-tailed and ferruginous hawk in 1979 and 1980 was 33 percent and 37 percent. The Swainson's hawk exhibited slightly greater overlaps in 1979 and 1980 of 54 percent and 52 percent with red-tailed hawks and 47 percent and 41 percent with ferruginous hawks. Habitat over- lap of the Swainson's hawk tended to be minimized by its selection of nest trees whose configuration differed from red-tailed and fer- ruginous hawks. Space was also partitioned by the maintenance of intraspecific and, to a lesser degree, interspecific nesting ter- ritories. Nest success of red-tailed and ferruginous hawks signifi- cantly declined when nesting pairs were visible to one another. This suggests that competition for space was a mechanism affecting distribution, density, and nest success. Based on the findings from this study, Buteo management could involve: (1) range management for prey species, which in this case is in accord with good management for cattle, (2) range management for nesting habitat, and (3) maintenance of properly spaced nest- ing sites. Resource Partitioning and Reproductive Success of Three Species of Hawks (Butco spp.) in an Oregon Prairie by Marcy Jean Cottrell A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science October 1981 Commencement June 1982 ACKNOWLEDGEMENTS This study would not have been possible without the interest, encouragement, and involvement of dozens of people. I want to thank Dr. Charles Warren, my major professor, for his wonderfulsupport and inspiring guidance throughout the course of this study. Dr. Charles Henny was instrumental in the planning and execution ofthe entire project, and his knowledge constantly instructive. I also wish to thank OSIJ's Department of Fisheries and Wildlife, andthe U.S. Fish and Wildlife Service, in particular Dr. Charles Stone,for sup- porting and funding this project. I will always be sincerely grateful for the kindnesses I re- ceived from the ranchers of Wallowa County. All the ranching f am- ilies of the Zumwalt Prairie granted me permission to walktheir land and study the wildlife and range. Many took an enthusiastic interest in the study and offered assistance anytime I was in need. Individuals from several state and federal agencies also extend- ed their time and effort.Biologists of the Oregon Department of Fish and Wildlife of Enterprise volunteered many hours to trackdown hawk nests and record raptor breeding activities. Employees of the U.S. Forest Service of Wallowa County were also helpful,and Ralph Anderson was particularly informative on the raptors of theZumwalt. I am very grateful to Chris Maser of the Bureau ofLand Management for providing invaluable assistance on the study of thehawks' food habits. Mike Kochert contributed important information on thetiming of raptor nesting activities at the Snake River Birds of PreyRefuge. Persons from the Soil Conservation Service and theAgricultural Stabilization and Conservation Service, both of Enterprise, also volunteered their assistance whenever possible.Here a most special thank you goes to my good friend Don Baldwin for his enthusiastic support, good humor, and dedication to the project. The range manage- ment portion of the study could not have been done without his most generous help. Several other friends volunteered their time to assist with the field work. I will always be especially indebted to Christopher Carey, Tracy Fleming, Carol Barla Gregory, and John Houle, all of whom spent countless hours walking over the prairie to locate and moniter hawk nests. Their aid and constant friendship is very deep- ly valued. I also want to thank the students and staff at Oak Creek Laboratory of Biology for their support, good cheer, and lively con- versation. Lastly, my deepest gratitude goes to my parents, Dr. and Mrs. George Cottrell, for their encouragement and love, and fortheir humor which always saw me through. TABLE OF CONTENTS Introduction 1 Methods Results and Interpretation 8 Habitat 8 Buteo Density and Distribution inRelation to Nesting Habitat 16 Use of Nest Sites 25 Food Utilization 30 Food Availability 35 Timing of Nesting Activities 38 Territoriality 39 Reproductive Performance 42 Discussion Management Implications 53 Bibliography 58 Appendices 62 Appendix I ... 63 Appendix II 65 Appendix III 68 IST O' 'IGURES Figure Page 1. Photographs of the seven different ecological range 9 sites and of the cultivated land of the Zumwalt Prairie a. Rolling Hills Range Site 10 b. Biscuit Scabland Range Site 10 South Exposure Range Site 11 North Exposure Range Site 11 Dry Mountain Meadow Range Site 12 Pine Grove Range Site 12 Aspen Grove Range Site 13 h Cultivated Land -Southern Unit 13 i. Cultivated Land -- Prairie 14 2. Range sites and cultivated land used by the three 24 species of Buteo during 1979 and 1980 3. Nest substrates used by the three Buteo species in 27 the Zumwalt Prairie during 1979 and 1980 4. The heights of trees selected by red-tailed, 29 ferruginous, and Swainson's hawks for nesting, the heights of the nests in the trees, and the nest-tree diameter at breast height 5. Intraspecific and interspecific distances between 41 nests, and the.success of the nests, of the three species of Buteo during 1979 and 1980 LIST OF TABLES Table Page Evaluation of red-tailed hawk's preference or 18 avoidance of the nine different habitat types that occur in the Zuinwalt Prairie Evaluation of ferruginous hawk's preference or 20 avoidance of the nine different habitat types that occur in the Zumwalt Prairie Evaluation of Swainson's hawk's preference or 21 avoidance of the nine different habitat types that occur in the Zumwalt Prairie Prey of red-tailed hawks in the Zumwalt Prairie 31 during 1979 and 1980 Prey of ferruginous hawks in the Zuinwalt Prairie 32 during 1979 and 1980 Prey of Swainson's hawks in the Zumwalt Prairie 33 during 1979 and 1980 Nuntber of Belding ground squirrel burrows and 37 northern pocket gopher mounds per 200 m x 3 m transects conducted throughout the Zuinwalt Prairie Number of nestlings reared to fledging by red- 45 tailed, ferruginous, and Swainson's hawks in the Zumwalt Prairie during 1979 and 1980 RESOURCE PARTITIONING AND REPRODUCTIVE SUCCESS OF THREE SPECIES OF HAWKS (BUTEO spp.) IN AN OREGONPRAIRIE INTRODUCTION In a native grassland prairie of northeastern Oregon,red-tailed (Buteo jamaicensis), ferruginous (Buteo regalis), andSwainson's hawks (Buteo swainsoni) coexist in one of the densest concentrationsof nesting Buteo hawks in the United States. In 1979 and 1980, an average of 50 pairs of red-tailed, 24 pairs offerruginous, and 8 pairs of Swainson's hawks nested annually in the 363km2area, commonly re- ferredto as the Zumwalt Prairie. The Zumwalt Prairie is a relatively isolated system. Ranging in elevation from 1219 m to 1585 m, it is bordered to thesouth and west by mountains of 3050 m, to the east by steep canyonsdropping 475-914 m, and to the north by dense forest.Few shrubs occur in the prairie except along intermittent streams, and interfingeringsof the forest community reach out into the grassland in some areas.The Zuinwalt Prairie is the southernmost extension of the Palouse Prairie, anim- portant wheat producing region of the U.S..It is located 194 km south of the section of the Palouse where Fitzner(1978) conducted his intensive study of the Swainson's hawk. While most of the Palouse, including Fitzner's area, has had a large part of itsnative vegeta- tion destroyed by cultivation and water impoundment (Fitzner1978), the Zuiiwalt Prairie still is primarily composed ofnative bunchgrass species. It is one of the best examples in thenorthwest of native prairie in good to excellent condition (Baldwin, pers. comm.1980). Species such as red-tailed, ferruginous, and Swainson'shawks, 2 exhibitin9 very simix piches, must segregate themseLves ecologically through thepartitioning of resources, if they are tocontinueto coexist.- (Elton 1946, Lack 1971, Price 1975). On the basis of most empirical studies, it has been conjectured that interspecific compe- tition determines the patterns of resource partitioning among closely related species.