Structure, Development, and Spatial Patterns in Pinus Resinosa Forests of Northern Minnesota, U.S.A
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Structure, development, and spatial patterns in Pinus resinosa forests of northern Minnesota, U.S.A. A THESIS SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE UNIVERSITY OF MINNESOTA BY Emily Jane Silver IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE Anthony W. D’Amato Shawn Fraver July 2012 © Emily J. Silver 2012 Acknowledgements I would like to acknowledge many sources of inspiration, guidance, and clarity. First, I dedicate this manuscript to my co-advisors Dr. Anthony W. D’Amato and Dr. Shawn Fraver. I thank Tony for supporting my New England nostalgia, for endless patient explanations of statistical analyses, for sending me all over the state and country for enrichment opportunities, and advising me on the next steps of my career. To Shawn, my favorite deadwood-dependent organism, for countless hours in the field, persistent attention to timelines, grammar, details, and deadlines, and for career advice. I would like to thank Dr. Robert Blanchette, the most excited and knowledgeable third committee member I could have found. His enthusiasm and knowledge about mortality agents and fungi of the Great Lakes region helped me complete a very rewarding aspect of this research. Second, to the colleagues and professionals who provided ideas, snacks, drinks, and field work assistance: Miranda and Jordan Curzon, Paul Bolstad, Salli and Ben Dymond, Paul Klockow, Julie Hendrickson, Sascha Lodge, Justin Pszwaro, Eli Sagor, Mike Reinekeinen, Mike Carson, John Segari, Alaina Berger, Jane Foster, and Laura Reuling. Third to Dr. Tuomas Aakala (whose English is better than my own) for good humor, expertise, and patience coding in R and conceptualizing the spatial statistics (initially) beyond my comprehension. I would like to extend thanks to the people that made this work possible. To Brian Palik and the Forest Service Northern Research Station for the funding and support to i complete this project. To Rebecca Marty for re-discovering the Allison plot, and Dr. Mark Fulton for recognizing this project for its intrinsic value. I would like to recognize Chris Gronewold for providing missing data and inspiration when all seemed lost. I would like to thank John Bradford for access to red pine chronosequence data that strengthened the analyses presented here. I appreciated the ease with which I arranged field lodging, thanks to the tireless work of Erin Fider and Jon Ross at the Itasca Biological Station as well as Pat and Dave at Bert’s Cabins. I would like to thank the University of Minnesota’s Forest Resources Department, as well as the Graduate School for a one-year fellowship and a community that made me feel well-supported. I would like to honor and recognize the original scientists who initiated both the Allison plot and the Growing-Stock-Levels experiment: J.H. Allison, S. Spurr, J. Zasada, J. Benzie, and P. Zehngraff. Their foresight and wisdom has led to two amazing datasets and research yet to come. Thank you to Doug Kastendick for maintaining the Growing- Stock-Levels study today. Finally, I would like to thank my family members and friends, from Massachusetts to Minnesota, for feeding me and reminding me why I am a researcher. Specifically to Scott Silver, Hannah Ramer, and Andrew Huff for reading drafts of this manuscript. I would like to thank Andrew for patiently watching me fill our at-home, wall-sized white-board with lists, grids, R and SAS code, and occasional expletives. His love and support were an unexpected, but joyful part of this research process. And to the managers and recreationalists of forests: I hope this research continues to support the conservation, protection, and stewardship of the world’s rich forest resources, whether they be wildlands or woodlands. ii Abstract The developmental dynamics and structure of old-growth forests often serve as important baselines for forest management. Nonetheless, long-term information on natural patterns of development and mortality is rare for many commercially and ecologically important forest types. This study uses a long-term dataset from a large (2 hectare), stem-mapped plot within an old-growth Pinus resinosa (red pine) forest in Itasca State Park to quantify patterns of forest structural development and mortality events over an 87-year period. These patterns were compared with those found in other old-growth red pine -dominated forests in northern Minnesota, as well as with unmanaged second-growth and extended rotation red pine forests to quantify the range of variation in structural conditions and to evaluate the effectiveness of extended rotation forestry at promoting the development of late-successional structural conditions. Old-growth forests were more structurally complex than unmanaged second- growth and extended rotation red pine systems, due in large part to the significantly higher volumes of coarse woody debris, higher snag densities, and spatially random patterns of mortality found in old growth. Nonetheless, extended rotation thinning treatments, by accelerating advancement to larger tree diameter classes, generated diameter distributions closely approximating those found in old growth. These results suggest that extended rotation treatments may accelerate the development of old-growth structural characteristics, provided that random patterns of mortality are emulated and coarse woody debris and snags are deliberately retained and created on site. These and other developmental characteristics of old-growth systems can inform red pine iii management when management objectives include the restoration of late-successional forest conditions. iv Table of Contents Acknowledgements .............................................................................................................. i Abstract .............................................................................................................................. iii List of Tables .................................................................................................................... vii List of Figures .................................................................................................................. viii Chapter One: Introduction .................................................................................................. 1 Chapter Two: The spatial pattern of mortality in an old-growth Pinus resinosa - dominated forest in northern Minnesota, U.S.A. ................................................................ 4 1. Introduction ................................................................................................................. 4 2. Methods....................................................................................................................... 7 2.1 Study area ............................................................................................................. 7 2.2 Long-term permanent plot sampling ..................................................................... 8 2.3 Field and laboratory methods ............................................................................... 9 2.4 Data analysis ...................................................................................................... 10 2.4.1 Mortality rates ................................................................................................. 10 2.4.2 Spatial pattern analyses ................................................................................... 11 3. Results ....................................................................................................................... 14 3.1 Mortality rates .................................................................................................... 14 3.2 Spatial patterns of mortality .............................................................................. 16 3.3 Mortality agents .................................................................................................. 20 4. Discussion ................................................................................................................. 20 4.1 Mortality rates .................................................................................................... 21 4.2 Spatial pattern of mortality ................................................................................. 22 4.3 Mortality agents .................................................................................................. 24 4.4 Conservation and management implications ...................................................... 25 4.5 Conclusions ......................................................................................................... 27 Chapter Three: Structure and stand-development of old-growth, unmanaged second- growth, and extended rotation Pinus resinosa forests in northern Minnesota, U.S.A. ..... 29 1. Introduction ............................................................................................................... 29 2. Methods..................................................................................................................... 32 2.1 Study sites............................................................................................................ 32 2.2 Field methods ...................................................................................................... 36 2.3 Statistical analysis .............................................................................................. 37 v 3. Results ......................................................................................................................