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Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak Forests of the Unglaciated Allegheny Plateau A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science John J. Wiley Jr. May 2013 © 2013 John J. Wiley Jr. All Rights Reserved. 2 This thesis titled Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak Forests of the Unglaciated Allegheny Plateau by JOHN J. WILEY JR. has been approved for the Department of Environmental and Plant Biology and the College of Arts and Sciences by Brian C. McCarthy Professor of Environmental and Plant Biology Robert Frank Dean, College of Arts and Sciences 3 ABSTRACT WILEY, JOHN J., JR., M.S., May 2013, Environmental and Plant Biology Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak Forests of the Unglaciated Allegheny Plateau (94 pp.) Director of Thesis: Brian C. McCarthy Silvicultural treatments are applied to forests to meet a variety of management objectives including timber production and wildlife habitat management. However, such methods may also profoundly affect other non-timber forest resources and negatively impact biodiversity. Modern forest science has largely neglected the role of forest management activities on bryophytes; thus, we have little insight as to how bryophyte communities respond to stand level treatments. The goal of this investigation was to explore changes in mixed-oak forest bryophyte communities associated with the common silvicultural methods of prescribed fire and thinning. Study sites were within the design of the USDA Forest Service Fire and Fire Surrogate (FFS) Research Program located in three southeastern Ohio forests. Each of these forests contain four treatments: untreated control, prescribed fire only, thinning only, and combined prescribed fire and thinning. Bryophyte occurrence and associated environmental variables were estimated in five 2 × 5 m quadrats along nine linear transects stratified by an integrated moisture index (IMI) in each treatment. I found a total of 116 bryophyte taxa (97 mosses, 19 liverworts). Of these, only 65 were found in more than 5% of the transects. Burning in xeric sites clearly altered bryophyte community richness and composition as burned sites had a 50% 4 reduction in mean richness and a disproportionate decrease in corticolous species. Mesic moisture conditions appear to mitigate species loss and community change. Thinning strongly altered woody substrate availability; however, the increased availability of these substrates did not significantly alter the bryophyte community in relation to their abundance. The use of fire on xeric sites may have a profound long-term negative impact on bryophyte diversity. Thinning may have a multi-decadal legacy effect on bryophyte species that does not appear to significantly degrade the bryophyte community. 5 ACKNOWLEDGMENTS I would like to express my gratitude to all those individuals and organizations that were integral to the development of this research. Foremost, I would like to thank my thesis advisor, Dr. Brian C. McCarthy. I gained much from his guidance and experience in academia, and on a several notable occasions, in life. I would also like to thank my committee members Dr. Morgan L. Vis and Dr. Jared L. DeForest for their input and advice during the development of this thesis and my graduate career. Perhaps one of the most challenging and rewarding aspects of this research was immersing myself in the intricate and often cryptic world of bryophytes. My initial introduction was guided by Dr. Nancy Slack at the Humboldt Field Research Institute, Maine. Later collaborations with the Ohio Moss and Lichen Association, especially Dr. Barbara Andreas, Diane Lucas, and Dr. Robert Klipps, and the Kent State Herbarium proved to be invaluable in my understanding of the Ohio bryophyte flora. The field work for this research was extensive and, often times, onerous, but would not have been possible without the access and information provided by the USFS Vinton Furnace Experimental Station. Also, I would like to thank my field assistants, Sarah Gutzwiller and Pualani Wiley. They both endured long days in less than ideal situations, and I am grateful for their companionship and dedication to detail. Funding for this research was provided by the Ohio Biological Survey, Hiram Roy Wilson Fund in Environmental and Plant Biology, Ohio University Graduate Student Senate, Ohio Moss and Lichen Association, and Association of Southeastern Biologists. 6 TABLE OF CONTENTS Page Abstract ................................................................................................................... 3 Acknowledgments................................................................................................... 5 List of Tables .......................................................................................................... 8 List of Figures ....................................................................................................... 12 Introduction ........................................................................................................... 15 Methods................................................................................................................. 22 Study Sites ........................................................................................................ 22 Study Design ..................................................................................................... 23 Field Methods ................................................................................................... 25 Substrates .......................................................................................................... 26 Site Conditions .................................................................................................. 29 Bryophytes ........................................................................................................ 29 Data Analysis .................................................................................................... 30 Results ................................................................................................................... 35 Bryophyte Diversity .......................................................................................... 35 Site Condition and Substrate Changes .............................................................. 37 Bryophyte Community Response ..................................................................... 42 Taxonomic Relationships ................................................................................. 44 Treatments and Substrates ................................................................................ 45 7 Discussion ............................................................................................................. 48 Floristics ............................................................................................................ 48 A Synthesis ....................................................................................................... 50 Management Implications ................................................................................. 55 Conclusions ....................................................................................................... 59 Tables .................................................................................................................... 60 Figures................................................................................................................... 74 References ............................................................................................................. 83 8 LIST OF TABLES Page Table 1. Bryophyte species, general growth form (L, Leafy; T, Thalloid; A, Acrocarpous; P, Pleurocarpous), coefficient of conservatism (C of C; higher is more habitat specific) (Andreas et al. 2004), forest of occurrence (R, Raccoon Ecological Management Area; T, Tar Hollow; Z, Zaleski;*,county record;`,verified historical record), and accession number from the Cooperrider Herbarium, Kent State University collected in three mix-oak forests of southeastern Ohio...................................................................................60 Table 2. Most commonly encountered (> 10% of all transects, n = 108) bryophytes in three mixed-oak forests in southeastern Ohio. .......................................................64 Table 3. Forest diversity metrics: number of species (S), Shannon-Weiner diversity (H’), and evenness (J) for the bryophyte community occurring in three mixed- oak forests in southeastern Ohio. ...........................................................................65 Table 4. Mean (± S.E.) of site condition variables and substrate cover values within 2 × 5 m plots established in xeric, intermediate, and mesic moisture classes under a 2 × 2 factorial of silvilcultural thinning and prescribed fire in three mixed-oak forest in southeastern Ohio. Cover of substrate types represent the 2-D coverage within the plot as visually mapped onto a gridded representation of the plot. Cover types are: exposed soil (SOIL), leaf litter (LEAF), coarse woody debris (CWD, diameter ≥ 10.0 cm, < 45° from the ground), fine woody 9 debris (FWD, diameter ≥ 1 cm and < 10 cm), tree base cover (TREE, all stems >DBH in height), stump base (STUMP, dead standing stems ≥ 45° from the ground and
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