In Riding Mountain National Park, Manitoba, Canada
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Landscape-Level Vegetation Dynamics in Riding Mountain National Park, Manitoba, Canada by Richard Caners thesis presented to the University of Manitoba in partial fulfillment of the requirements for the degree of Master of Science in the Faculty of Graduate Studies Departrnent of Botany University of Manitoba Winnipeg, Manitoba, Canada R3T 2N2 @ Richard Caners 2001 NationalLibrarY Bibliothèque nationale l*l du Canada Acquisitions and Acquisitions et Bibliographic Services services bibliographiques 395 Wellington Street 3!15, rue Wellington Ottawa ON KlA 0|,¡4 OttawaON KlA0N4 Canada Canada Yùt frle Voús rólèmæ Oufrle ì'loÚetélétw The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loatr, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sr¡r format électonique. 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Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts from it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sâns son permission. autorisation. 0-612-76901-1 Canad'ä THE T]NTYERSITY OF MANITOBA F'ACULTY OF GRADUATB STIIDIES ¡trr**tr COPYRIGHT PERMISSION LANDSCAPE-LEVEL VEGETATION DYNAMICS IN RIDING MOI]NTAIN NATIONAL PARIÇ MANITOBA, CANADA BY RICHARD CANERS A ThesisÆracticum submitted to the Faculty of Graduate Studies of The University of Manitoba in partial fulfillment of the requirement of the degree of MASTER OF'SCIENCE RICHARD CANERS O 2OO1 Permission has been granted to the Library of the University of Manitoba to lend or sell copies of this thesis/practicum, to the National Library of Canada to microfilm this thesis and to lend or sell copies of the film, and to University Microfilms Inc. to publish an abstract of this thesis/practicum. This reproduction or copy of this thesis has been made available by authority of the copyright o\üner solely for the purpose of private study and research, and may only be reproduced and copied as permitted by copyright laws or with express written authorization from the copyright owner. ABSTRACT The cornplex physiography and disturbance history of Riding Mountain National Park have resulted in a complex assemblage of aspen parkland, eastern deciduous and boreal forest communities on the landscape. Few studies have examined forest composition, structure and dynamics in the eastern extension of the mixedwood forest region. This study uses detailed vegetation, edaphic and environmental data collected from 202-100 mz forest plots distributed throughout the Park to elucidate landscape-level trends in forest structure and dynamics. Stands were flrrst classified into 8 dominant stand types, and described in terms of their biotic and abiotic characteristics. Factors affecting patterns of understory tree regeneration and the tirning of their recruitment were examined on the landscape, and growth of understory green ash (Fraxinus pennsylvanica Marsh.) and white spruce (Picea glauca (Moench) Voss) were analyzed in different habitats to infer future changes in stand composition. Successional trajectories were created for each of the 8 stand types using a static size-class analysis of tree species, and a comprehensive concepfual model of stand dynamics was created. Sections of this thesis have previously been published in Caners & Kenkel (1998). Results indicate that successional trajectories for stand types in the Park do not converge towards a single self-perpetuating 'climax' communify. Instead, vectors diverge, converge and remain cyclical, with multiple potential pathways recognized for each stand type. This study demonstrates that species assemblages, and the propensity for change, are governed by the cumulative and synergistic effects of climate, topography, disturbance frequency, size and intensity, edaphic conditions and the proximity of parental seed sources. These factors have resulted in a patchwork mosaic of forest stands on the landscape of varying structure, composition and seral stage. Overall, results are in general agreement with studies from central and eastern regions of the boreal forest. Post-fire stands are dominated by pioneering hardwoods such as aspen, balsam poplar and paper birch. Mid-succession stands show an increasingly greater proportion of white spruce in the canopy, whereas late-succession stands are dominated by white spruce and balsam fir and are driven by gap dynamic processes. The oldest stands are commonly open and shrub- dominated, especially by beaked hazelnut and mountain maple. In areas of dense shrub cover and/or areas without a proximate seed source, regeneration of tree species is dramatically reduced. Ungulate herbivores selectively browse trees and shrubs, impacting the long-terms dynamics of forest systems. ACKNOWLEDGEMENTS This thesis would not have been possible without the support received from my family, friends, colleagues and advisors. My family has provided me with motivation, self-confidence and financial support during my years of study. My fellow colleagues and labmates have listened intently, counseled and provided much-needed advice. Dedicated and unremitting field assistance was provided by Rod Lastra, Boyan Tracz and Heidi Wiebe; a sincere thank-you for all of your much-appreciated work away from home. Many hours of volunteer assistance was provided by Rod Lastra and Dave Walker. Additional field assistance and direction was welcomingly provided by the staff of Riding Mountain National Park, especially Roger Baird, Stephen Cornelson, Sean Fry, Terry Hoggins, Clem Labbe, Steve Malcolm, Gord Pylypiuk and Jacques Saquet. I would like to extend my sincere thanks to Wybo Vanderschuit for his guidance and friendship during my work in the Park. Gregg Morden (Department of Geology) gave technical advice and assistance in the processing of soil samples, and Dr. Les Fuller (Department of Soil Science) provided guidance in the identification of soil profiles. I would like to thank the members of my advisory committee, Dr. Bruce Ford (Department of Botany), Dr. Stephen McCanny (Canadian Heritage, Parks) and Dr. Rene Van Acker (Department of Plant Science), for their advice and direction. I would especially like to thank my advisor, Dr. Norm Kenkel (Department of Botany), for his patience, support, guidance and tireless efforts with this project. My passion for ecology stems from his teachings and enthusiasm. This project was funded by a grant from Heritage Canada, Parks, and an Engineering Research Councíl of Canada individual operating grant to N.C. Kenkel. LIST OF FIGURES Fig 2.1. The location of Riding Mountain National Park in relation to surrounding vegetation regions. --__66 Fig2.2. Total precipitation (June - Sept.; mm) and maximum July temperature ("C) in relation to elevational changes along the Manitoba Escarpment. ____ 61 Fig 3.1. Location of plots sampled within Riding Mountain National Park between 1996-1991 for stand types I-VIII 102 Fig 3.2. Height ranges for each tree canopy, and quadrat design. __________-__-____-___ 104 Fig 3.3 a-d. Residual correspondence analysis ordinations of ¡z:196 (a), n:164 (b) n:133 (c), and n:115 (d) sampled stands. -.-........'.--105 Fig 3.4. Minimum stand age frequency histograms for each of the eight stand types I- vtrr. ......-... ....107 Fig 3.5 a-h. Canonical correspondence analysis biplots of environmental variables (a), trees (b), shrubs (c), herbs (d), graminoids (e), ferns and fern allies (f), bryophytes (g), and plots (h) for n:179 stands and 6 environmental variables, utilizing n:l7l species. _______-- 108 Fig 4.1. Measurement of ring widths for green ash(Fraxinus pennsylvanica Marsh.). --------160 Fig4.2. Calculation of mean annual incremental radial growth (mm.yr-r) for green ash. .__________ ...._.. 160 Fig 4.3. Multiple discriminant analysis of 3 groups using n:75 stands and p:14 variables. ______ 16l Fig 4.4. Mean density and height class of tree species for n:7 aspen-dominated boreal forest and n:3 mixed eastern deciduous forest stands from which understory green ash were sampled. _.__...__162 Fig 4.5. Age (years), basal diameter (cm) and height (m) relationships for ri:44 understory (<10 m in height) green ash sampled from n:7 aspen-dominated boreal forest stands and n:3 mixed eastem deciduous stands. -___-___163 Fig 4.6. Age of n:44 understory (<10 m in height) green ash in relation to the minimum age of the stand (years) from which they were sampled. ......__________164 Fig 4.7. Comparison of incremental radial growth (mm . yr-'; for ¡z:21 harvested understory green ash from n:J aspen-dominated boreal stands, as well as n:8 canopy green ash from n:3 mixed eastern deciduous stands and n:14 understory green ash from the same iz:3 mixed eastern deciduous stands. 165 1il Fig 4.8. Mean incremental radial growth (mm . yr-'¡ for understory and canopy green ash sampled from n:J aspen-dominated boreal and n:3 mixed eastern deciduous stands. t66 Fig 4.9. The effects of Dutch elm disease on two mixed eastern deciduous forest canopies. 161 Fig 5.1. Measurement of ring widths for white spruce (Picea glauca (Moench) Voss). .------ 185 Fig 5.2. Calculation of mean annual incremental radial growth (rnm . yr-r) for white spruce. _- -_- - 185 Fig 5.3. Specimen age (years) against square root of height for n:67 harvested understory (<10 m in height) white spruce ___--___-186 Fig 5.4. Multiple discriminant analysis of 4 groups using rz:119 stands and p:14 variables. 187 Fig 5.5. Age of n:92 understory (<10 m in height) white spruce in relation to the minimum age of the stand (years) from which they were sampled.