A Study of Epixylic Bryophyte Ecology on Fraser Fir Logs in the Great Smoky Mountains National Park
Total Page:16
File Type:pdf, Size:1020Kb
University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange Masters Theses Graduate School 12-1998 A Study of Epixylic Bryophyte Ecology on Fraser Fir Logs in the Great Smoky Mountains National Park Erica Choberka University of Tennessee - Knoxville Follow this and additional works at: https://trace.tennessee.edu/utk_gradthes Part of the Botany Commons Recommended Citation Choberka, Erica, "A Study of Epixylic Bryophyte Ecology on Fraser Fir Logs in the Great Smoky Mountains National Park. " Master's Thesis, University of Tennessee, 1998. https://trace.tennessee.edu/utk_gradthes/2383 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 Erica Choberka entitled "A Study of Epixylic Bryophyte Ecology on Fraser Fir Logs in the Great Smoky Mountains National Park." 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 Botany. David K. Smith, Major Professor We have read this thesis and recommend its acceptance: Ken McFarland, Sally P. Horn 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: 1 am submitting herewith a thesis written by Erica Choberka entitled "A study of epixylic bryophyte ecology on Fraser fir logs in the Great Smoky Mountains National Park." I have examined the finalcopy of this thesis for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Master of Science, with a major in Botany. l5aVidK. Smith, Major Professor We have read this thesis And recommend its acceptance: ! ) ;'!_!_I , l /,. ./ . Accepted for the Council: Associate Vice Chancellor and Dean of Thc Graduate School A STUDY OF EPIXYLIC BRYOPHYTE ECOLOGY ON FR.A.SER FIR LOGS IN THE GREAT SMOKY MOUNTAINS NATIONAL PARK A Thesis Presented for the Master of Science Degree The University of Tennessee, Knoxville Erica Renee Grimm Choberka December 1998 DEDICA. TI ON This thesis is dedicated to my husband, David, and all of my closest friends, whose smiles and love have made this thesis possible. II ACKNOWLEDGMENTS There are many people that have generously helped and encouraged me throughout the course of this research. I would particularly like to thank Dr. David K. Smith, who oversaw this research. He provided me with many valuable skills and insights. I appreciate that he shared with me his wisdom about field work. I would like to thank Dr. Sally Hom and Dr. Ken Mcfarland fo r serving on my committee and providing helpfulcomments, Dr. Paul Davidson fo r his help with liverwort identification, Bernice Stevens for providing accommodations and inspiration, Keith Langdon and Janet Rock for their issuing of a bryophyte collecting permit fo r the Great Smoky Mountain National Park, Jamie Estill for computer help, and Timeka Newson, Jennifer \Vinkler. David Sime, Mark Whited, Theresa Lange, Dr. Smith, and Dr. McFarland fo r helping with field work. I would like to give a special thank you to my husband David and my extended family. Without their emotional support and encouragement, this study would not have been possible. Lastly, I would like to thank my friends fo r providing me with humor throughout my graduate career. This research was supported by the Aaron J. Sharp Fund and a Graduate Teaching Assistantship, the Department of Botany, The University of Tennessee, Knoxville. Ill Abstract The SouthernAppalach ian spruce-fir fo rest is experiencing the chaotic conditions of ecosystem destruction resulting from the balsam woolly adelgid (Adelges piceae (Ratz)) infestation. In the present study, I have examined the community structure of bryophytes on fir logs in the high elevation spruce-firfo rest of the Great Smoky Mountains National Park (GSMNP) to learn about the responses ofbryophytes to the sudden change in forest structure. This study has four primary objectives: 1) to provide an updated list of the epixylic bryophytes on fir logs in the spruce-firfo rest ofthe GSMNP; 2) to compare results with previous epixylic bryophyte studies performed in the Great Smoky Mountains National Park; 3) to describe bryophyte communities on logs and describe the environmental factors that control community structure; 4) to create a quantitative method fo r sampling bryophyte species cover on logs. Epixylic bryophytes on 79 Fraser fir (Abiesfraseri (Pursh) Poir.) logs were sampled. Relative frequency values for the species were scored within the upper surface of a 60 degree arc on the log, in em x 100 em quadrats. Environmental variables such as general location, longitude and latitude, slope. slope aspect. elevation. dominant tree and shrub species, and canopy cover were recorded for each plot. Three different multivariate tech..'liques were used in this study, TWINSPAN, Detrended Correspondence Analysis (DCA), and Direct Gradient Analysis (DGA). TWINSPAN separated all the sites sampled into three different bryophyte communities. No wellia curvifolia, Brutherella rccurvans, and Brotherella recurvansl Hyp num IV imponens. These communities were separated into nine unions that describe diffe rent combinations of species and environmental conditions. TWINSPAN also separated all the species that perform similarly into six diffe rent clusters. These clusters are directly related to environmental conditions such as available light and decay stage. DCA provided a two-dimensional scatterplot of each species's overall average response within sampled sites. Finally, DGA provided insight into the environmental factors that significantly influence species distribution. The results of this study suggest that each bryophyte species responds uniquely to environmental factors and that species replacement occurs in a unidirectional pattern. There seem to be five factors that most significantly influence the presence of a species on a log: species life strategy, species ability to colonize optimal substrate, the amount of bryophyte cover on the log, the decay class of a log, and the canopy conditions. The epixylic bryophyte communities on Fraser firlogs in the GSMNP have drastically changed since the health of the spruce-fir fo rests have declined. A total of 1 9 species that were present on Fraser firlogs in the past are now missing completely, and many other species that were once abundant are declining. Three main union types have been lost: unions fo und on very wet logs, unions of corticolous species found on recently fallen fi r trees. and unions of soil species fo und on completely decayed logs. Furthermore, there seems to have been a shiftfr om species rich communities that used to be fo und on moist logs in a healthy spruce-fir fo rest. to less species rich communities that are currently found on dryer logs in the decimated fo rest. v TABLE OF CONTENTS CHAPTER PAGE I. INTRODUCTION .......................................................... II. B"<\CKGROUND.... ........................................................ 2.1 The Spruce-Fir Forest ofthe SouthernAppalachians ......... 2.2 Fraser Fir ........................................................... 6 Bryophyte Ecology ............................................... 10 III. LITERATUREREVIEW .................................................. 13 3.1 Epixylic Bryophyte Research.................................... 13 Epixylic Bryophyte Research in the Great Smoky Mountains National Park......................................... 23 IV. MATERIALSAND METHODS......................................... 26 4.1 Field Site Description............................................. 26 4.2 Field Methodology................................................ 26 4.3 Analytical Methods................................................ 31 4.4 Multivariate Techniques.......................................... 32 4.4.1 Direct Gradient Analysis................................. 32 4.4.2 Ordination (Indirect Gradient Analysis). .............. 33 4.4.3 Classification (Cluster Analysis). ....................... 35 V. RESULTS I DISCUSSION ................................................ 37 5.1 Species List . ........................................................ 37 VI 5.2 TWINSPAN Results............................................. 37 5.3 TWlNSPAN Discussion........................................ 39 5.3.1 TWlNSPAN Bryophyte Unions ...................... 39 )- . .).� ., ') TWINS PAN Species Groups .......................... 42 5.4 DCA Results ...................................................... 44 5.5 DCA Discussion ................................................. 45 5.6 Direct Gradient Analysis Results .............................. 47 5.7 Direct Gradient Analysis Discussion .... ...................... 54 5. 7.1 Species Relative Frequency vs. Decay. .............. 54 5.7.2 Species Relative Frequency vs. Canopy Class...... 55 5.7.3 Species Relative Frequency vs. Amount of Bryophyte Cover......................................... 55 5.7.4 Species Relative Frequency vs. Log Position........ 56 VI. CONCLUSION ............................................................ 57