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SEED COVER Internet L A N D M A N A G E M E N T H A N D B O O K 40 Field Studies of Seed Biology 1997 Ministry of Forests Research Program Field Studies of Seed Biology Carole L. Leadem, Sharon L. Gillies, H. Karen Yearsley, Vera Sit, David L. Spittlehouse, and Philip J. Burton Ministry of Forests Research Program Canadian Cataloguing in Publication Data Main entry under title: Field studies of seed biology (Land management handbook ; ) “Tree seed biology.”--Cover. --- . Trees – British Columbia – Seeds – Experiments. Trees – Seeds – Experiments. Reforestation – British Columbia – Experiments. Reforestation – British Columbia – Experiments. I. Leadem, Carole Louise Scheuplein, – II. British Columbia. Ministry of Forests. Research Branch. III. Series .. .’’ -- © Province of British Columbia Published by the Research Branch B.C. Ministry of Forests Bastion Square Victoria, BC Copies of this and other Ministry of Forests titles are available from Crown Publications Inc. Fort Street Victoria, BC Please address any comments or suggestions to the senior author: Carole L. Leadem Glyn Road Research Station B.C. Ministry of Forests PO Box Stn Prov Govt Victoria, BC ii CREDITS Carole L. Leadem B.C. Ministry of Forests, Research Branch, Glyn Road Research Station, PO Box Stn Prov Govt, Victoria, BC Sharon L. Gillies University College Fraser Valley, King Road, Abbotsford, BC H. Karen Yearsley B.C. Ministry of Forests, Research Branch, PO Box Stn Prov Govt, Victoria, BC Vera Sit B.C. Ministry of Forests, Research Branch, PO Box Stn Prov Govt, Victoria, BC vw c David L. Spittlehouse B.C. Ministry of Forests, Research Branch, PO Box Stn Prov Govt, Victoria, BC vw c Philip J. Burton Symbios Research and Restoration, PO Box , Smithers, BC Editor, indexer: Fran Aitkens Typesetting: Dynamic Typesetting Graphic production: Lyle Ottenbreit Proofreading: Rosalind C. Penty Steve Smith Publication design: Anna Gamble Original photos and illustrations: Figures ., ., . Dr. David Spittlehouse Figure . Dr. John Owens, Dep. Biol., Univ., Victoria, B.C. Figures . and . Paul Nystedt, B.C. Min. For. Figures ., ., . H. Karen Yearsley Figures . and . Dr. D.G.W. Edwards, Can. For. Serv., (retired) Pac. For. Cent., Victoria, B.C. iii INTRODUCTION I like trees because they seem more resigned to the way they have to live than other things do. (Willa Cather “O Pioneers!”) Except in limited areas where there is enough methods for conducting field studies of tree seeds. A advance regeneration, establishment of forest cover recent assessment of ecosystem management needs on harvested lands continues to depend on seedling stressed the importance of standardized sampling planting programs or on natural regeneration by and monitoring techniques, and the lack of consist- seeds. Whereas successful plantation programs ent methods for archiving, accessing, and updating depend primarily on plant competition and site databases (U.S. Dep. Agric. For. Serv. a). Tech- variables at the time of planting, successful natural niques gleaned from agriculture literature are generally regeneration depends not only on the availability not applicable, and traditional ecological studies of seeds, but on favourable environmental condi- (e.g., of seed banks) tend to be primarily descriptive tions throughout the processes of seed production, with little emphasis on experimental approaches. dispersal, germination, and seedling establishment. The primary objective of this manual is to detail Site preparation and other silvicultural treat- methods that have been gleaned from the literature ments can improve the suitability of the seedbed and from personal experience of the authors. It is a and its micro-environment, but there is still much manual of methods with some general guidelines we do not understand about how various factors and interpretation. Relevant background papers are contribute to successful forest establishment. We cited where appropriate, but it is not a literature have gained some insights, under controlled condi- review. The manual is intended for use by researchers tions, about the influence of major factors such as in public and private forest resource management light and temperature, but we have limited experi- agencies, universities, and colleges. Although specifi- ence with biological responses under actual cally directed to tree seed research in forested conditions in the field. ecosystems, many of the methods described can Anyone who has conducted research in the field be used to study seeds of graminoid, herb, and quickly comes to realize the complexity of the sys- shrub species in both forest and non-forest plant tems chosen for study. An immense number of communities. The extensive background informa- external and internal factors that affect living tion included in the text also provides valuable organisms must be taken into account—with lim- reference material for many who have an interest ited possibilities to control these factors. A major in tree seeds, but who are not directly involved in constraint, particularly in a forest environment, is research activities. The detailed examples from the difficulty inherent in conducting field studies previous studies are included, not to prescribe how involving seeds. Infrequent seed production, preda- such studies should be done, but to assist in plan- tion by animals, difficulty locating small seeds, ning by providing reference values on which to estimating the numbers of buried seeds, measuring base measurements, sample sizes, and other germination, and monitoring survival pose myriad experimental details. challenges for the field researcher. Added to these Since the manual is directed primarily to re- difficulties is the lack of information about effective searchers working in the province of British v Columbia (B.C.), Canada, many examples (forest types, species, research topics), procedures (the biogeoclimatic ecosystem classification system), and regulatory policies are specific to this geographic and political jurisdiction. Nevertheless, it is hoped that the underlying principles are self-evident and will be generally applicable to the conduct of field research elsewhere. vi ORGANIZATION OF THE HANDBOOK Following a discussion of planning and organizing a H. Karen Yearsley, M.Sc., R.P.Bio., earned her field study (Section ) and setting up an environ- graduate degree from the Faculty of Forestry, Uni- mental monitoring program for the experimental versity of British Columbia, and is a member of the site (Section ), the manual is arranged by subject Association of Professional Biologists of British areas most often associated with field studies of tree Columbia. Her years of research experience in seeds: natural seed production (Section ), seed dis- B.C. include work on ecosystem classification, persal (Section ), seed predation (Section ), seed forest succession, and forest soil seed banks. Karen banks (Section ), assessing seed quality and viabil- Yearsley wrote the sections on seed predation ity (Section ), and effects of silvicultural practices (Section ) and soil seed banks (Section ), and on emergence (Section ). Each section was written contributed to the sections on planning field by one or more experts as follows: studies (Table .) and seed dispersal (Section ). Carole Leadem, Ph.D., R.P.Bio., earned her Vera Sit, M.Sc., earned her graduate degree from degree in plant physiology from the Botany Depart- the Statistics Department, Dalhousie University, and ment, University of British Columbia, and is a is a member of the Statistical Society of Canada. She member of the Association of Professional Biolo- has been with Biometrics Section, Research Branch, gists of British Columbia. She has been in charge of B.C. Ministry of Forests, since . Vera Sit wrote the tree seed biology research program with the B.C. the sections on experimental design and data analy- Ministry of Forests in Victoria since . Carole sis (Sections ., ., ., ., ., ., .) and the Leadem wrote the sections on planning and case studies (Section .), and reviewed and contrib- organizing field studies (Section ), natural seed uted to all the statistical sections. production (Section ), seed responses to the envi- David Spittlehouse, Ph.D., P.Ag., earned his ronment (Section .), seed testing in the laboratory graduate degree in forest climatology from the De- (Section .), seedbed preferences (Section ..), partment of Soil Science, University of British and contributed to the sections on seed dispersal Columbia. His research includes modifying site and silvicultural practices. microclimate to improve seedling regeneration, and Sharon Gillies, Ph.D., earned her degree in plant determining how forest harvesting and regrowth of physiology from the Department of Biological Sci- the forest affects forest hydrology. He has worked ences, Simon Fraser University. She has been a for the B.C. Ministry of Forests in Victoria since biology instructor at the University College Fraser . Dave Spittlehouse wrote most of the section Valley since . Sharon Gillies coordinated compi- on designing an environmental monitoring program lation of the original manuscript, was responsible (Section ). for creating the handbook structure and adhering Philip Burton, Ph.D., R.P.Bio., earned his degree to Ministry of Forests style manual, edited author in plant biology from the University of Illinois at submissions for the first complete draft, wrote the Urbana-Champaign. An independent researcher and section on seed dispersal (Section ), and provided consultant, he has been investigating
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