Analysis of the Composition and Structure of Eastern
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CURRENT COMPOSITION AND STRUCTURE OF EASTERN HEMLOCK ECOSYSTEMS OF NORTHEASTERN OHIO AND IMPLICATIONS OF HEMLOCK WOOLLY ADELGID INFESTATION Thesis Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Thomas Daniel Macy, B.S. Graduate Program in Environment and Natural Resources The Ohio State University 2012 Thesis Committee: Dr. David M. Hix, Advisor Dr. P. Charles Goebel Dr. Stephen N. Matthews Copyright by Thomas Daniel Macy 2012 ABSTRACT Ohio’s eastern hemlock (Tsuga canadensis (L.) Carr.) ecosystems provide important ecological and economic benefits to the state. As a foundation species, eastern hemlock has a major influence on forest structure and functional processes. Eastern hemlock is also dominant in several popular outdoor recreation and tourist areas in Ohio. Since its accidental introduction in 1951 hemlock woolly adelgid (Adelges tsugae Annand; HWA), an invasive insect native to Japan, has been causing widespread mortality of eastern hemlock in an expanding portion of its range. We sampled two types of hemlock forests that occur in northeastern Ohio: hemlock swamp forests of the Huron-Erie Lake Plain physiographic region (ELP) and stream-ravine forests of the Glaciated Allegheny Plateau physiographic region (GAP). Vegetation and environmental data were collected from seven mature stands in northeastern Ohio. Information on the current composition and structure of these forests will be critical for predicting potential pathways of stand development in response to the possible decline of eastern hemlock and for informing management and restoration plans. The objectives of our study were to: 1) compare the community composition and structure of eastern hemlock forest ecosystems of the ELP and GAP; 2) identify environmental factors influencing plant composition and structure; and 3) model HWA- ii induced eastern hemlock mortality and predict changes in forest composition and structure using the Forest Vegetation Simulator (FVS) and the Hemlock Woolly Adelgid Event Monitor. Principal components analysis (PCA) showed clear separation between stands of the two physiographic regions based on slope percent and slope position, with stands of the ELP occupying flat bottomlands and stands of the GAP occupying middle- and upper-hillslope positions. Hierarchical cluster analysis failed to group trees, saplings, seedlings, or ground-flora by physiographic region, while multi-response permutation procedures (MRPP) detected a significant difference only in the seedling stratum (p = 0.03) between physiographic regions. The plant community composition and structure of stands in the ELP and GAP were similar due to the influence of eastern hemlock, which was dominant in both the tree layer (> 55% importance value for both physiographic regions) and the sapling layer (> 47% relative density for both physiographic regions), but was absent from the seedling layer. Redundancy analysis (RDA) revealed species- environment relationships consistent with species life-history traits and habitat requirements. FVS analysis forecast significant decreases (p < 0.05) in the total basal area and basal area of eastern hemlock following HWA infestation in both physiographic regions. FVS predicted decreases in eastern hemlock basal area up to 80% thirty years after infestation. The potential loss of this foundation species is forecast to drastically alter forest composition, structure, and functional processes. iii ACKNOWLEDGEMENTS I would like to thank my advisor, Dr. David Hix, for his guidance and advice during both my undergraduate and graduate careers at Ohio State. I would also like to thank committee member Dr. Charles Goebel for providing very useful insights that improved this research. Their generosity in helping me to attend and present at various professional conferences has greatly increased my knowledge and broadened my understanding of natural systems. I would also like to thank committee member Dr. Stephen Matthews, for his interest in this research and helpful comments. I am grateful to the School of Environment and Natural Resources for the opportunity to be a teaching associate, as well as the faculty and staff for their assistance. I would like to thank the Ohio Agricultural Research and Development Center for awarding me a SEEDS Grant, without which, this research would not have been possible. Thank you to The Cleveland Museum of Natural History, Friends of Wooster Memorial Park, The Holden Arboretum, The Nature Conservancy, and Ohio State Parks for granting me access to their properties to conduct this research. Finally, I would like to thank Sarah Adams and my family for their love and support. iv VITA 2010........................................B.S. Forestry, Fisheries, and Wildlife, specialization in Forestry and Wildlife Management, School of Environment and Natural Resources, The Ohio State University 2010 to 2012 ..........................Graduate Teaching Associate, School of Environment and Natural Resources, The Ohio State University Fields of Study Major Field: Environment and Natural Resources Area of Specialization: Forest Science v TABLE OF CONTENTS Abstract ............................................................................................................................... ii Acknowledgements ............................................................................................................ iv Vita .......................................................................................................................................v List of Tables ................................................................................................................... viii List of Figures ......................................................................................................................x Chapter 1: Introduction ........................................................................................................1 Background ....................................................................................................................1 Biology and Characteristics of Hemlock .......................................................................2 Hemlock Woolly Adelgid ..............................................................................................6 Research Justification and Objectives ...........................................................................9 References ....................................................................................................................12 Chapter 2: Plant Community Composition and Structure of Eastern Hemlock Forest Ecosystems of Northeastern Ohio ......................................................................................19 Introduction ..................................................................................................................19 Study Area ...................................................................................................................21 Methods........................................................................................................................23 Vegetation ..............................................................................................................23 Environmental Attributes .......................................................................................24 Data Analyses ........................................................................................................25 Results ..........................................................................................................................29 Principal Components Analysis .............................................................................29 Plant Community Composition and Structure .......................................................29 Plant Community Diversity ...................................................................................31 vi Downed Woody Debris..........................................................................................31 Hierarchical Cluster Analysis ................................................................................32 Multi-response Permutation Procedure ..................................................................32 Redundancy Analysis.............................................................................................32 Discussion ....................................................................................................................34 References ....................................................................................................................41 Chapter 3: Modeling Hemlock Woolly Adelgid-Induced Mortality and Predicting Stand Development Using the Forest Vegetation Simulator .......................................................70 Introduction ..................................................................................................................70 Study Area ...................................................................................................................73 Methods........................................................................................................................75 Vegetation ..............................................................................................................75 Forest Vegetation Simulator ..................................................................................75 Data Analyses