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Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation

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Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange Masters Theses Graduate School 5-1997 Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation George Francis Smith University of Tennessee - Knoxville Follow this and additional works at: https://trace.tennessee.edu/utk_gradthes Part of the Ecology and Evolutionary Biology Commons Recommended Citation Smith, George Francis, "Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation. " Master's Thesis, University of Tennessee, 1997. https://trace.tennessee.edu/utk_gradthes/1339 This Thesis is brought to you for free and open access by the Graduate School at TRACE: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Masters Theses by an authorized administrator of TRACE: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a thesis written by George Francis Smith entitled "Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation." I have examined the final electronic copy of this thesis for form and content and recommend that it be accepted in partial fulfillment of the equirr ements for the degree of Master of Science, with a major in Ecology and Evolutionary Biology. Niki S. Nicholas, Major Professor We have read this thesis and recommend its acceptance: John Rennie, J. Frank McConnick Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official studentecor r ds.) To the Graduate Council: I am submitting herewith a thesis written by George F. Smith entitled "SouthernAppalachian Fir and Fir-SpruceForest CommunityChanges Following Balsam Woolly Adelgid Infestation." I have examined the finalcopy of this thesis for fo rm and content and recommend that it be accepted in partialfulfil lment of the requirements for the degree of Master of Science, with a major in Ecology. Niki S. Nicholas, Major Professor We have read this thesis and recommendits acceptance: Accepted fo r the Council: Southern Appalachian Fir and Fir-Spruce Forest Community Changes Following Balsam Woolly Adelgid Infestation A Thesis Presented fo r the Master of Science Degree The University of Tennessee, Knoxville George Francis Smith May 1997 Acknowledgments Nearly all scientificresearch nowadays is the result of the collaboration of many individuals, and this thesis is no exception. I am especially gratefulfo r the guidance and encouragement provided by my advisor, Niki Nicholas. John Rennie and J. Frank McConnick, members of my Thesis Committee, have made manysugges tions that have greatly improved this research. I have benefitedfr om the knowledge and supportof the graduate students and facultyin the Department of Ecology and EvolutionaryBiology. I thank Paul Durr and the rest of the biological research staffof the Great Smoky Mountains National Park who initiated this study and collected the data. Eric Pauley, Chris Eagar, and Peter White provided helpful comments on partsof this thesis. The National Park Serviceand the Tennessee Valley Authority provided much appreciated financial assistance. I am verythankf ul for the support provided by my parents, George and Ruth Smith, and especially my wife, Rachel. I would not have gotten far without her loving encouragement. ii Abstract Thesouthern Appalachian firand fir-spruce forests are unique glacial relict communities that occupy 26,600 ha at the highest elevations of only seven mountain areas. Over the last three and a half decades an exotic insect, the balsam woolly adelgid (Adelges piceae Ratz.), has caused catastrophic mortalityto populations ofthe endemic Fraser fir (A biesfraseri (Pursh) Poir.) throughout its entire native range. In 1990 and 1991, a set of temporaryand permanent plots were established at the summits of five mountains in the Great Smoky Mountains to study processes of community change in overstory and understory composition and firsize and age distributions. Nearly 70% of standing firbasal areais dead over the entire study area. Patterns of overstory dominance are determined primarily by elevation and time since the major wave of mortality. Overstoryfir mortality and the resulting increases in canopy openness have led to decreases in densities ofO- 10 yr old firseed lings and bryophyte cover. Herbaceous vegetation cover, densities of red spruce (Picea rubens Sarg.) seedlings, and small shrub densities have increased in some stands. Understory firare growing and maturing more rapidly with greater insolation. Understory firrecru itment andselective mortality of larger firare causing overstory size and age classes to become more skewed towards smaller and younger individuals. Future rates and spatial patternsof infestation and mortality will be determined by rates of firrecovery in patches subject to different biotic and abiotic stressors. Poorly reproducing firmay be eliminated in lower elevation mixed stands. Ill Table of Contents CHAPTER PAGE I. Introduction and Objectives 1.1 Introduction 1 1.2 Objectives 3 II. Literature Review 5 2.1 The SouthernAppalachian Spruce-Fir Forest Community 5 2.2 Disturbance in Spruce-Fir Forests 12 2.3 The Balsam Woolly Adelgid 20 2.4 Current Status of the Spruce-Fir Ecosystem 26 m. Patterns of Overstory Composition 35 3.1 Abstract 35 3.2 Introduction 35 3.3 Methods 38 3.4 Results 41 3.5 Discussion 52 3.6 Conclusions 60 IV. Understory Responses to Overstory Change 62 4.1 Abstract 62 4.2 Introduction 63 4.3 Methods 65 4.4 Results 68 4.5 Discussion 75 iv CHAPTER PAGE 4.6 Conclusions 87 v. Size and Age Class Distributionsof Overstoryand UnderstoryFras er Fir 88 5.1 Abstract 88 5.2 Introduction 89 5.3 Methods 91 5.4 Results 94 5.5 Discussion 102 5.6 Conclusions 114 VI. Summary 115 Literature Cited 119 Vita 129 v List of Tables TABLE PAGE 3.1 Temporary plot data-percent of standing basal area that is dead for fivelocations in GSMNP 43 3.2 Temporary plot data-logistic regressions of live and dead overstory fir, spruce, and hardwood frequencies on elevation, years since major 2 mortality, and (years since major mortality) for five locations in GSMNP 44 3.3 Permanent plot data-multivariate regression of live fir, spruce, and hardwood basal area and density on elevation, years since major mortality, percent slope, and linearly transformed aspect for five locations in GSMNP 51 3.4 Permanent plot data-logistic regressions of live and dead overstory fir, spruce, and hardwood frequencies on plot elevation, years since major mortality, percent slope, and transformed aspect for five locations in GSMNP 53 4.1 Multivariate regression of understory firin size classes on understory spruce, hardwoods, and shrubs for five locations in GSMNP 73 4.2 Mean percent cover for herbs, twig/leaf litter, and bryophytesin four overstory stand types for five locations in GSMNP 74 4.3 Multivariate regression of understory fir,spruce, hardwoods, and shrubs on elevation, years since major mortality, transformedaspect, and overstory fir, spruce, and hardwood basal area for five areas in GSMNP 77 5.1 Mean dbh and age of cone-bearing fir for five locations in GSMNP 97 vi List of Figures FIGURE PAGE 2.1 Southern Appalachian spruce-fir forest areas 6 3.1 Temporary plot data-live and dead fir, spruce, and hardwood basal area for five locations in GSMNP 42 3.2 Temporary plot data-predictions of the proportion of standing Fraser fir stems that would be dead on an 1891 m elevation mountain over time since major mortality based on logistic regression and data from five locations in GSMNP 46 3.3 Permanent plot data-liveand dead fir, spruce, and hardwood basal area for five locations in GSMNP 47 3.4 Permanent plot data-live and dead fir, spruce, and hardwood basal area by stand type for five locations in GSMNP 48 3.5 Permanent plot data-liveand dead fir, spruce, and hardwood density for fivelocations in GSMNP 49 3.6 Permanent plot data-liveand dead fir, spruce,and hardwood density by stand type for five locations in GSMNP 50 4.1 Live fir, spruce, and hardwood sapling and large shrub densities for fivelocations in GSMNP 69 4.2 Live fir, spruce, and hardwood sapling and large shrub densities by stand type for fivemountains in GSMNP 70 4.3 Live fir, spruce, and hardwood seedling and small shrub densities for fivelocations in GSMNP 71 vii FIGURE PAGE 4.4 Live fir, spruce, and hardwood seedling and small shrub densities by stand type for five locations in GSMNP 72 4.5 Fir (in 4 size classes), spruce, and hardwood seedlings and shrub species < 1.37 m tall. Mean frequency by percentage cover of three ground cover types 76 5.1 Mean age of Fraser fir in six size classes for five locations in GSMNP 95 5.2 Number of dominant and codominant overstory fir individuals in 10 yr age classes for fivelocations in GSMNP 96 5.3 Densities of overstory fir in 1 em dbh size classes for fivelocations in GSMNP- power function model 99 5.4 Densities of overstory fir in 1 em dbh size classes for five locations in GSMNP- sine wave model 100 5.5 Cumulative proportion distributions of overstory fir in I em dbh size classes for five locations in GSMNP 101 5.6 Densities of understory fir in 5 yr age classes for five locations in GSMNP 103 5.7 Cumulative proportion distributions of understory fir in 5 yr age classes for five locations in GSMNP 104 5.8 Densities of understory fir in 5 yr age classes by stand type for five locations in GSMNP 105 5.9 Cumulative proportion distributions of understory fir in 5 yr age classes by stand type for five locations in GSMNP 106 viii Chapter I Introduction and Objectives 1.1 Introduction The southern Appalachian fir and spruce-fir forests are physiognomically and floristically unique ecosystems found only in an island-like distribution at the peaks of the seven highest mountain areas in NorthCarolina, Tennessee, and southwestern Virginia (Oosting and Billings 195 I; Ramseur 1960).
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