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Growth and Reproductive Ecology of Larix Laricina in Interior Alaska

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Growth and Reproductive Ecology of Larix Laricina in Interior Alaska AN ABSTRACT OF THE THESIS OF Kevin Brown for the degree of Master of Science in Botany and Plant Pathologypresented on 16 December 1982 Title: GROWTH AND REPRODUCTIVE ECOLOGY OF LARIX LARICINA IN INTERIOR ALASKA Redacted for Privacy Abstract approved: bonAdB. Zobel Larix laricina in Alaska occurs on sites ranging from productive slopes and riverbottoms to wet bogs underlain by permafrost. It is largest on sites without permafrost, where it is associated with Picea glauca, but is much more common in bogs, where it occurs with Picea mariana. In this study, I describe the environment of different Larix habitats; compare growth and reproduction of Larix on different sites; and compare the ecology of Larix with the associated Picea species. Larix is a deciduous conifer; however, some needles on juvenile Larix, usually on new shoots, are retained during the winter and function the following spring. This habit may conserve carbohydrates and nutrients. Browsing damage to seedlings by snowshoe hares is common. Larix in bogs die younger than those in warm uplands or riverbottoms; the maximum ages observed in each were 71,87, and 127 years, respectively.Maximum height and diameter growth rates of Larix were as great or greater than in uplands or riverbottoms,but the duration of rapid growth was twice as long in upland andfour times longer in riverbottom trees (90-100 years), resultingin their greater size. Cone production and seed viability of Larix vary betweenstands and from year to year. Maximum annual cone production among 10 cone crops observed was 1.97 x105conesha-1; the highest seed viability was 60 percent. When stand cone production was high, coneproduction of individual trees was strongly correlated withtree size. Seeds mature by mid-August and disperse beginning inlate August to early September. Ninety-five percent of viable seed at one stand fellby November 1. Germination in the laboratory is inhibited at or below 10 deg C. At 12 and 15 deg, germination is stimulated bylong photo- period and stratification period; thisstimulation decreases at higher temperatures. Natural germination occurs after soils warmand summer rains begin. In riverbottoms, germination is better on mineral seedbeds than feathermoss, possiblydue to a better moisture regime. In tussocky bogs, low temperatures maydelay germination in troughs;inadequate moisture may delay germinationin higher micro- sites. The best microsites for survival ofgermlings in tussocky bogs appear to be open troughswhich are high enough to escape sub- mergence. Larix 1 aricina appears to be aseral species on warmer sites, establishing soon after P. 11 auca, butdying earlier. Its seral relationship to P. mariana is less clear,in part due to ignorance of bog processes. GROWTH AND REPRODUCTIVE ECOLOGY OF LARIX LARICINA IN INTERIOR ALASKA BY KEVIN R. BROWN A THESIS Submitted to OREGON STATE UNIVERSITY in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Completed December 16, 1982 Commencement June 1983 APPROVED: Redacted for Privacy Professor of Botany and 1%tholgy in Isharge of major Redacted for Privacy Head of Department of otany and Plant Pathology Redacted for Privacy Dean of Graduate hoof 0 Date thesis is presented December 16, 1982 Typed by Mary Ann (Sadie) Airth for Kevin R. Brown ACKNOWLEDGEMENTS This study was funded by Cooperative Agreement 239 with the U.S. Forest Service Pacific Northwest Forest and Range Experiment Station. Their financial and logistical support are greatly appreciated. Support for computing services was provided by the Oregon State Uni- versity Computing Center. John Zasada first gave me the opportunity to work in theboreal Forest,in particular, interior Alaska, and sparked my interests in plant population ecology and silviculture, for which I am most grate- ful. Throughout this study, he was a source of encouragement and good ideas, more of which I should have listened to. My advisor, Don Zobel, deserves much credit.His high stand- ards, ready desire to help, to discuss ideas, and toreview all stages of this project were an invaluable help. The other members of my committee, Denis Lavender, DonMinore, and Jim Green, thoroughly and constructively reviewed this thesis under a severe timeshortage. I appreciate the chances to discuss this project withJoe Antos, who greatly helped to clarify my sometimes muddled thoughts,and the help of Francis Herman in arranging to have the stemanalysis performed. The past three years were made so much more enjoyableby the support (and craziness!) of many friends, includingMeadow, Mark, Marianne, and especially Anne.To all my contra-dancing friends, balance and swing! This thesis is dedicated to my parents, Randall and Helene Brown,and to my grandparents, Bryce and Ruth Kerr, for their stimulation of my interestin the outdoors, and their love, encouragement, (and financial support). TABLE OF CONTENTS Page I. INTRODUCTION 1 II. ENVIRONMENT 3 REGIONAL SETTING 3 STUDY SITES 5 Methods 8 Results 11 III. VEGETATION 25 INTRODUCTION 25 METHODS 27 RESULTS 29 Riverbottom 29 Shaw Creek Upland 34 Bogs 35 DISCUSSION 48 IV. GROWTH OF LARIX LARICINA 51 INTRODUCTION 51 METHODS 52 RESULTS 53 DISCUSSION 61 V. KEPRODUCTIVE ECOLOGY 68 INTRODUCTION 68 Reproductive Cycle of Larix spp. 68 PHENOLOGY OF FLOWERING AND POLLEN RELEASE 70 Methods 70 Results 71 CONE PRODUCTION AND SEED VIABILITY 74 Methods 74 Results 75 Discussion 78 SEED DEVELOPMENT 82 Methods 82 Results 84 Discussion 86 SEED DISPERSAL 92 Methods 92 Results 93 Discussion 96 GERMINATION RESPONSE TO TEMPERATURE, PRE- CHILLING, AND LIGHT 97 Methods 97 Results 101 Discussion 108 FIRST-YEAR GERMLING EMERGENCE AND SURVIVAL 117 Methods 117 Results 119 Discussion 125 GENERAL DISCUSSION OF REPRODUCTIVE ECOLOGY 136 VI. SYNTHESIS 140 BIBLIOGRAPHY 145 LIST OF FIGURES Figure Page 1. Map of Alaska showing range of Larix laricina and general locations of study sites 7 2. Air temperatures during a clear and cloudy day, July 4-5, 1981, at BCO, SCF, and SCU 15 3. Substrate temperatures at 0, -5, and -15 cm in the sedge meadow transect, SCF, May-August, 1981 16 4. Precipitation and water table fluctuation, Bonanza Creek, in 1981 20 5. Age structure of Larix laricina and Picea mariana at BCO, BCF, and BCF-N 40 6. Age structure of Larix laricina and Picea mariana at Goldstream (GS-1, GS-4) 42 7. Age structure of Larix laricina and Picea mariana at Smith Lake and SCF 43 8. Height growth of young Larix laricina and Picea mariana in bog sites and young L. laricina, P. glauca, and P. mariana at T-Tield, 1981 54 9. Overwintering needles on Larix laricina seedlings in a) late September 1980 and b) May 1981 56 10. Percentage of Larix laricina from bog sites bearing overwintering needles 57 11. Height growth of Larix laricina with age in bog, upland, and riverbottom sites 59 12. Seasonal patterns of needle and shoot elongation of Larix laricina at BCO, 1981 60 13. Branch of mature Larix laricina in May, 1981, showing an erect female cone and vegetative shoot morphology 72 14. Timing of pollen flight for a single tree at BCF-N, 1981 73 15. Mean lengths of megagametophytic tissue, embryo cavity, embryo, and cotyledons of Larix lariciana seed collected at SCF, 1981 85 16. Germination of L. laricina seed collected on 3 days in August 1981 at SCF 88 17. Total and viable seedfall of Larix laricina under the BCF stand, September 1980-November 1981 94 18. Dispersal patterns of viable Larix seed 9/80-5/81 based on samples from 30 traps 95 19. Germination of seed from Alaska 11 and 19 days after beginning of test; experiment 1 104 20. Germination versus temperature after 17 days, Alaskan seed 107 21. Germination vs. time; NS32, Minnesota, and SCF, Experiment 3 110 22. Germination rate, midday surface temperature, and surface volumetric moisture , riverbottom site, (BCR) seedbed plots, 1981 121 23. Total germling emergence and survival, treated plots, upland site (SCU) in 1981 123 24. Total germling emergence and survival at bog (BCO), microsites - 1981 126 25. Germination rate, midday surface temperature and surface volumetric moisture, BCO bog microsites, 1981 128 LIST OF TABLES Table Page 1 Location and description of study sites 6 2 Air temperature summaries from study sites and nearby weather stations, 1980-81 13 3 Monthly precipitation (cm), May-August 1980 and 1981, at two study sites and the nearest weather stations. 18 4 Soil matric potential at T-Field, 1981 21 5 Xylem pressure potentials "p" measured at midday 22 6 Basal area and density of tree species on study sites 31 7 Diameter class distribution of tree species on riverbottom (BCR, Willow Island) and upland (SCU) study sites 32 8 Age class distribution of Larix laricina and Picea glauca on riverbottom (BCR, Willow Island) and upland (SCU) study sites 33 9 Size structure of Picea mariana and Larix laricina at BCO, 1981 36 10 Tree age by size class of Picea mariana and Larix laricina at BCO 38 11 Stem diameter-age correlations of mature Larix laricina in bog study sites 38 12 Percent cover of vegetation (excluding trees) and substrate types at bog study sites 45 13 Relationship of Larix laricina seedling density to total density of saplings and mature trees in bog sites 46 14 Occurrence of natural Larix seedlings by type of substrate in bogs 47 15 Total elongation (mm) of leaders and terminal first-order long shoots on mature Larix laricina in 1980 and 1981 62 16 Cone production by Larix laricina at five study sites in 1980 and1787 76 17 Correlations of cone production per tree with tree size and age at bog study sites, 1980-81
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