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Pacific Northwest Forest Industry on Natural and Planted Regeneration

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Pacific Northwest Forest Industry on Natural and Planted Regeneration AN ABSTRACT OF THE THESIS OF ELVER HOWARD VOTH for the DOCTOR OF PHILOSOPHY (Name) (Degree) in ZOOLOGY; NATURAL HISTORY presented on October 10, 1967 (Major) (Date) Title: FOOD HABITS. OF THE PACIFIC MOUNTAIN BEAVER, APLODONTIA RUFA PACIFICA MERRIAM Abstract approved: Dr. Robert M. Storm The mountain beaver causes extensive loss to the Pacific Northwest forest industry on natural and planted regeneration sites, primarily to Douglas fir, by clipping or burying seedlings during its burrowing activity and also by damaging older trees by pruning or girdling. This project studied a heavy population and its annual harvest of vegetation on a 2- hectare site with alder succession problems on the north side of Marys Peak in the Oregon Coast Range from 1964 to 1966. Food habits methods were reviewed to include work by Adams, Aldous, Arata, Baumgartner and Martin, Dusi, Korschgen, Swift and Bishop, and Williams, and compared stomach contents analysis and fecal pellet analysis approaches. The methods used included histological preparation of reference material based on herbarium specimens and photographic plates of plant tissue slides to identify recognition items of 34 species of ingested plants from a site total of 141. Three series of fecal samples were analyzed from live trapped animals captured one or more at a time over a calendar year, the three series including adult males, lactating females (April, May, and June), and juveniles following weaning. Feeding experi- ments confirmed recognition items and sought to establish a leaf epidermis fragment equivalence index. A vegetation yield analysis made comparison of available vegetation with intake possible. Minimal statistical treatment justified number of samples, sample size, and counting procedures using 200X magnification of 10 1.0 mm diameter microscope fields for each sample taken from a fecal pellet collection. Results verified opinions of the mountain beaver as a primitive rodent eating a primitive plant diet. Deer - management studies, especially that by Crouch, have shown little dependence by deer on pteridophytes such as sword -fern or bracken. These two species composed 82 percent of the adult male mountain beaver diet. This rodent eats bracken, poisonous to other herbivores, with complete immunity. The animal occupies a food niche no other herbivore wants and which in respect to ferns is not in competition with deer. Lactating females take in a very large percentage of high - protein high-moisture-content young grasses. The juvenile diet after weaning corresponds to the adult male diet but includes more protein. Included were much behavioral and some food plant nutritional analysis information. Site excavations and biotelemetry served as accessory sources of data. Facilities were provided by the Oregon State University Forest Research Laboratory. Some support was provided by the National Science Foundation. Food Habits of the Pacific Mountain Beaver, Aplodontia rufa pacifica Merriam by Elver Howard Voth A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 1968 APPROVED: Professor of Zoology in charge of major Chairman of the Department of Zoology Dean of the Graduate School Date thesis is presented October 10, 1967 Typed by Gwendolyn Hansen for Elver Howard Voth ACKNOWLEDGEMENTS Numerous persons have given assistance in the furtherance of this project. Those to whom the author is particularly indebted include management personnel of the Forest Research Laboratory, especially Dr. Hugh Black for initial suggestions on the kind of research desired and for arranging many needed services and facilities, Edward Hooven for much helpful advice, and Richard Newman for planting the seedling areas. Dr. Kim Ching provided microscope facilities. Mr. John Thompson gave permission to use the site chosen on his property. Dr. William Lawrence of the Weyerhaeuser Company paid a visit to the Forest Research Laboratory on August 11, 1965, and included a stimulating private discussion on histological food habits techniques. Dr. Paul Weswig of the Forage Analysis Labora- tory kindly provided proximate analyses of several fern samples submitted. Dr. Robert Storm, major professor, first showed the author a suitable mountain beaver area,: subsequently listened to and answered many questions raised during the course of the work, gave much helpful guidance, and spent many hours in conference and reading manuscript in connection with the research. Gratitude is also expressed to colleagues, especially William Lovejoy and Sheldon Johnson, for their interest, and to James York, for his assistance during much of the summer of 1966. The National Science Foundation provided a fellowship during the summer of 1964. TABLE OF CONTENTS Page INTRODUCTION 1 Usual Limits of Food Habits Research 1 Review of Literature on Food Habits Research Methods 8 Review of Literature on the Mountain Beaver and its Distribution 20 METHODS AND MATERIALS 26 Limits of the Study 26 Site Selection 27 Animal Project 34 Trapping Program 34 Population Analysis of the Study Site 37 Use of Biotelemetry 42 Pellet Collection and Preservation 43 Captive Animal Colony 44 Plant Project 46 The Recognition Item 46 The Nature of Leaf Epidermis and Some Other Leaf Tissue 72 The Reference Item 7 5 Statistical Project 89 Counting Procedures 89 Tests of Significance 99 RESULTS AND DATA 106 Population of the Study Site 106 Vegetation of the Study Site 114 Analysis of Diet Composition by FPA Epidermal Fragment Count 127 Adults (Predominantly Males) 135 Lactating Females 136 Juveniles 136 Comparison of Available Vegetation and its Place in the Diet 136 Annual Production, Phenology, and Aspection 136 Annual Harvest 139 Uses or Destruction of Vegetation for Non -diet Connected Purposes 153 Page DISCUSSION 158 Characteristics and Behavior Related to Feeding 158 The Activity Period 158 Fossorial Behavior 167 Harvesting Behavior 179 Feeding Behavior 186 Ingestion 194 Digestion 196 Defecation 200 Refection 201 Nutritional Requirements of the Animal 202 CONCLUSIONS 207 Significance of the Diet 207 Problems Associated with the Method 208 Management Implications 211 BIBLIOGRAPHY 213 APPENDICES I. Plant List 247 II. Plates 256 LIST OF FIGURES Figure Page 1. Successive treatments of the distribution of Aplodontia rufa subspecies since Taylor, 1918. 24 2. Investigated known locations of pacifica populations. 29 3. Map of the Blodgett -Marys Peak area. 30 4. Map of the study site showing locations of animal captures, burrow systems, burrow excavations, vegetation study quadrats, and Douglas fir planted seedling areas. 32 5. Sweeping pattern used in slide examination. 93 6. Distribution of study site and vicinity trapping efforts and animal captures through the trapping calendar, for 32 animals numbered 12 -44 (except for Animal 23, an accession from Valsetz). 109 7. Mountain beaver population density fluctuation curve inferred for normal study site conditions, 1964 -1965. 113 8. Proportions of population belonging to different age classes at different times, and animal biomass com- parisons, for one hectare. 115 9. Percentage composition of diet, males, based on number of epidermal fragments appearing in fecal pellets in 50 microscope fields per pellet collection per individual. 132 10. Percentage composition of diet, lactating females, based on number of epidermal fragments appearing in fecal pellets in 50 microscope fields per pellet collection per individual. 133 11. Percentage composition of diet, juveniles, based on number of epidermal fragments appearing in fecal pellets in 50 microscope fields per pellet collection per individual. 134 Figure Page 12. Number of plant species in diet, captured males, based on identification of epidermal fragments appearing in fecal pellets in 50 microscope fields per pellet collection per individual. 13 5 13. Aspection of layer dominants and other plants of the study site. Ferns. 140 14. Aspection of layer dominants and other plants of the study site. Grasses. 140 15. Aspection of layer dominants and other plants of the study site. Trees and low perennial dicot ground cover. 140 16. Aspection of layer dominants and other plants of the study site. Monocots other than grasses, and annuals. 141 17. Aspection of layer dominants and other plants of the study site. Perennial herbs. 141 18. Aspection of layer dominants and other plants of the study site. Rosetting evergreen herbs. 141 19. Dependence on seasonal and layer availability. 153 20. Plan of burrow system. Excavation 1. 170 LIST OF TABLES Table Page I. Comparative Treatment of Ingested Materials and Behavior Related to Digestion, and their Usefulness in Determining Methods of Terrestrial Mammal Food Habits Study 16 II. Trapping Efforts and Success on the Study Site 35 III. Sex and Age Class Criteria for Pacifica Animals Taken from the Study Site 41 IV. Summary of Feeding Experiment Data: Species Fed 50 V. Summary of Feeding Experiment Data: Relative Quantities Ingested 51 VI. Summary of Feeding Experiment Data: Pellet Weights Obtained 52 VII. Summary of Feeding Experiment Data: Fragments Counted From Last Day Pellet Samples, 1/8 Pellet (Approx. 0.06 g) Spread Under 22x40 mm Coverslip 53 VIII. Summary of Feeding Experiment Data: Fragments Counted from Stomach Samples, 0.06 g Spread Under 22x40 mm Coverslip 53 IX. Summary of Feeding Experiment Data: Fragments Counted Not Belonging to Single Species Fed by Day of Last Pellet Collection 54 X. Summary of Feeding Experiment Data: Sample Frag-
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