Clemson University TigerPrints All Theses Theses 8-2015 A NEW, LOW-ELEVATION APPALACHIAN SAXIFRAGE (HYDATICA NECKER EX. GRAY 1821) IN SOUTH CAROLINA LAARY CUSHMAN Clemson University, [email protected] Follow this and additional works at:https://tigerprints.clemson.edu/all_theses Recommended Citation CUSHMAN, LAARY, "A NEW, LOW-ELEVATION APPALACHIAN SAXIFRAGE (HYDATICANECKER EX. GRAY 1821) IN SOUTH CAROLINA" (2015).All Theses. 2190. https://tigerprints.clemson.edu/all_theses/2190 This Thesis is brought to you for free and open access by the Theses at TigerPrints. It has been accepted for inclusion in All Theses by an authorized administrator of TigerPrints. For more information, please [email protected]. A NEW, LOW-ELEVATION APPALACHIAN SAXIFRAGE (HYDATICA NECKER EX. GRAY 1821) IN SOUTH CAROLINA A Thesis Presented to the Graduate School of Clemson University In Partial Fulfillment of the Requirements for the Degree Master of Science Plant and Environmental Sciences by Laary Jackson Cushman August, 2015 Accepted by: Patrick D. McMillan, Committee Chair Elena Mikhailova Peter Adler Harry E. Shealy, Jr. ABSTRACT In 2002 a unique population of Hydatica Necker ex Gray 1821 (Appalachian Saxifrage) was discovered by Patrick McMillan in the upper Piedmont of Pickens County, SC, growing on a low-elevation granite dome within the Nature Conservancy’s Nine Times nature preserve. The plants appeared to be similar to the widespread Hydatica petiolaris (Raf.) Small populations of the Blue Ridge Escarpment (BRE). However, the Pickens County population displays an annual life form and flowers during the late winter and early spring, in contrast to the perennial life forms and late spring through early fall flowerings of the BRE population. A study was initiated to determine if the population in Pickens County meets the criteria of the biological, phylogenetic and ecological species concepts and thus constitutes it as a novel species. Morphology comparisons, non-coding DNA analysis, common garden experiments, and ecological comparisons were chosen as species concept criteria to test high-elevation H. petiolaris populations and the low-elevation, Pickens County Hydatica sp. population. The floral form and stature, distinct DNA phylogeny, and unique adaptive zone of the Pickens County population proved distinct from other Hydatica petiolaris populations. Therefore, experimental evidence indicates that the Pickens County population meets three different species concepts, representing a novel species known only to this location. ii DEDICATION Thanks to my wife, Larisa Jean, and my children, Alekzandre, Lili, and Phillip, for their support and putting up with 3 years of fieldwork, and fridges full of plants, bugs, and sometimes dead animals; your love, patience, collaboration, and faith in my ability as a Husband, Father, and Student helped to drive me thus far. iii ACKNOWLEDGEMENTS I would first like to thank my thesis director, advisor, and friend, Dr. Patrick D. McMillan, for mentoring me through this project and for his guidance, wisdom, patience, and unrestrained freedom to pursue the questions answered herein; without these things, I would have never come thus far. I would also like to thank Drs. Elena Mikhailova and Peter Adler for serving on my thesis committee and for all their invaluable advice, suggestions, and insight, as well as Dr. Harry E. Shealy, Jr., who has and still serves as my mentor since my first botany class, many years ago. I would like to thank Dr. Kathy Mathews of Western Carolina University, for her invaluable help in teaching me a foundation in plant genetics through the use of her lab, and Max Lanning for his help with specific plant data.. Dr. Michael Caterino of Clemson University was also very gracious in allowing me lab space and time alongside his entomology students, while I extracted the much needed DNA for my final analysis. And thanks to the Wade T. Batson Grant for Field botany, and its committee for two financial awards that helped to expedite this research. Thanks to my family, who often hiked with me searching for a small glimpses of this plant in the most rugged and beautiful of places, and to my hiking companion, Chad Bates, for making the long, lonely hikes into the granite domes of God's country more enjoyable. Mostly, I thank my Lord and Saviour, Christ the King, who has made all things known and unknown, and can still make Man anew. The hand that has drawn the limits of the granite hills and the boundary of the salted sea, still draws men to the foot of a Cross; Romans 10:9, The Holy Bible. iv TABLE OF CONTENTS Page TITLE PAGE .................................................................................................................... i ABSTRACT ..................................................................................................................... ii DEDICATION ................................................................................................................ iii ACKNOWLEDGMENTS .............................................................................................. iv LIST OF TABLES ......................................................................................................... vii LIST OF FIGURES ...................................................................................................... viii CHAPTER I. INTRODUCTION ......................................................................................... 1 The Blue Ridge Escarpment ................................................................... 1 The case of Hydatica petiolaris (Raf.) Small .......................................... 3 Species concepts ..................................................................................... 5 Systematics ............................................................................................. 6 Objectives of Study ................................................................................. 6 II. MATERIALS AND METHODS ................................................................... 8 The Nine Times Nature Preserve (Study Site) ......................................... 8 Study Sites and Sampling ........................................................................ 8 Biological Analysis .................................................................................. 9 Controlled Garden Experiments ............................................................ 10 Ecological Analysis ............................................................................... 11 Phylogenetic Analysis ............................................................................ 13 III. RESULTS .................................................................................................... 16 Controlled Garden Experiments ............................................................ 16 Climatic Data ......................................................................................... 16 Carolina Vegetative Survey ................................................................... 17 Morphological Analysis ......................................................................... 19 Phylogenetic Analysis ............................................................................ 20 v Table of Contents (Continued)................................................................................... Page IV. DISCUSSION AND CONCLUSIONS ...................................................... 22 Species Concept Support ....................................................................... 22 V. TAXONOMIC TREATMENT AND KEY ................................................. 26 TABLES ........................................................................................................... 30 FIGURES ........................................................................................................... 36 REFERENCES ........................................................................................................... 62 APPENDICES ........................................................................................................... 67 APPENDIX A ........................................................................................................... 68 APPENDIX B ........................................................................................................... 94 vi LIST OF TABLES Table Page 1. List of vouchers examined from the Clemson University Herbarium (CLEMS). ........................................................................... 28 2. First open flower months of NTP and BMT open garden plots vs NTP and BMT field study plots. ............................................. 29 3. Summary of NTP and BMT monthly weather data segregated into High, Low, and Average temperatures, with ANOVA of NTP and BMT monthly weather data following. .......................................................... 30 4. Rank Abundance tables with top ten rankings of CVS survey depicting community composition .......................................... 31 5. ANOVA testing of environmental explanatory variables used in CCA analysis. .......................................................................... 32 vii LIST OF FIGURES Figure Page 1. The Blue Ridge Escarpment traversing northeastern Georgia through South and North Carolina, into Virginia ................................................................................ 34 2. The Nine Times Nature Conservancy Site, located in Pickens, County, SC. ................................................................... 35 3. Representatives of the family Saxifragaceae depicting variable floral morphology and arrangements.A, Boykinia, B, Chrysosplenium, C, Micranthes, D, Ozomelis. ..................................................................................... 36 4. A) High-elevation Hydatica petiolaris B) low-elevation Hydatica sp.1. .............................................................................. 37 5. Study site locations: Nine Times Preserve (NTP), SC, 440 meters (m) above sea level (ASL); Blood Mountain (BM), GA 1036m ASL; Tallulah Gorge (TG), GA, 428m ASL; Big Creek Trail (BCT), GA, 670m ASL; and the Balsam Mountains region (BMT), NC 1584m ASL. ............................... 38 6. Typical 1m2 study plot at the NTP study site. The Hydatica sp.1 population is in flower. ........................................ 39 7. Hydatica sp.1 petals mounted on slides for measurement of length and width data with microscope. .................................. 40 8. Controlled Garden Experiment plantings at the South Carolina Botanical Garden greenhouse. Ambient garden planting trays shown with orange tags representing 2013 plantings, white tags representing 2014 plantings. .................................... 41 9. Patrick D. McMillan performing portion of vegetative survey at NTP. Author is taking picture. ................................... 42 10. Comparative weather data of the NTP and BMT sites for study cycle years 2014 – 2015. NTP and BMT flowering start times indicated by shaded bars. ................................................................................. 43 11. National Weather Service’s National Digital Forecast Database (NDFD) Annual weather data for Pickens, SC. Range is 01/13/2014 – 01/16/2015 ................................................................................... 44 viii LIST OF FIGURES (CONT.) Figure Page 12. National Weather Service’s National Digital Forecast Database (NDFD) Annual weather data for Pickens, SC. Range is 01/13/2014 – 01/16/2015 ................................................................................... 45 13. Rank abundance profiles of vegetative survey at NTP site. .............................................................................................. 46 14. Rank abundance profiles of vegetative survey at BMT site. .............................................................................................. 47 15. This plot shows us that soil depths has a great influence on the presence of species, and that the elevation is the second factor that explains distributions. Many of the species distributions are centered near the soil depths vector. Some environmental factors were also nulled due to identification as useless constraints. They are completely removed from the estimation, and no biplot scores or centroids are calculated for these constraints. (R Core Team 2015). ................................................................... 48 16. This plot shows us that the elevation is the greatest factor that explains distributions. A lot of the species distributions are centered near the soil depths vector. Also note that some environmental factors were nulled due to identification of useless constraints. They are completely removed from the estimation, and no biplot scores or centroids are calculated for these constraints. (R Core Team 2015). Eigen value for Axis 1, 0.7926, Axis 2, 0.1894. ....................................................................................... 49 17. PCA ordination of NTP and BMT CVS data. The vectors drawn are for species Hydatica petiolaris and Hydatica sp1. The projections for the sites indicate a ranking of sites from low to high along each individual vector. Eigen value for Axis 1, 61.4657, Axis 2, 30.6999 ........................................................................................ 50 ix