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Ecological and Genetic Status of the Purple Pitcher Plant, Sarracenia Purpurea L., in Maryland and Virginia

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Ecological and Genetic Status of the Purple Pitcher Plant, Sarracenia Purpurea L., in Maryland and Virginia ECOLOGICAL AND GENETIC STATUS OF THE PURPLE PITCHER PLANT, SARRACENIA PURPUREA L., IN MARYLAND AND VIRGINIA by Philip M. Sheridan B.S. Biology, May 1994, Virginia Commonwealth University M.S. Biology, August 1996, Virginia Commonwealth University A Dissertation Submitted to the Faculty of Old Dominion University in Partial Fulfillment of the Requirement for the Degree of DOCTOR OF PHILOSOPHY ECOLOGICAL SCIENCES OLD DOMINION UNIVERSITY May 2010 Approved by: ________________________ Dr. Frank Day (Director) ________________________ Dr. Kneeland Nesius (Member) ________________________ Dr. Robert Griesbach (Member) ________________________ Dr. Lytton Musselman (Member) ABSTRACT ECOLOGICAL AND GENETIC STATUS OF THE PURPLE PITCHER PLANT, SARRACENIA PURPUREA L., IN MARYLAND AND VIRGINIA Philip M. Sheridan Old Dominion University, 2010 Director: Dr. Frank Day Sarracenia purpurea is a rare wetland plant in Virginia and a threatened species in Maryland, with two potential subspecies in the region. I utilized restriction fragments from the intron of the chalcone synthase gene to compare S. purpurea populations and determine whether the subspecies concept was supported. I performed a census of existing populations, compiled all known historical data on the species, and investigated the reasons for the species demise and predicted dates of extinction. Bloom phenology was examined to see if climate change may have influenced bloom period. Soil, vegetation, and climatic information was obtained to determine if taxonomic differences correlated with environmental variables. I found no genetic difference in the intron of the chalcone synthase gene in mid-Atlantic S. purpurea populations while I did find differences with other Sarracenia species and S. purpurea varieties. These results suggest that a single taxon of S. purpurea occurs in Maryland and Virginia. Only 31% (4 of 13)of the sites are extant on the western shore of Maryland and District of Columbia while 33% (14 of 42) of the sites remain in Virginia with respective populations of 46 and 513 clumps. Causes of regional extirpation include beaver flooding, succession, and development. Predicted pitcher plant population extinction dates, based on trend line from130 years of data, are 2015 (Maryland) and 2055 (Virginia). Disturbance, especially natural fire, played an essential role in maintaining purple pitcher plant historically in Maryland and Virginia. Sarracenia purpurea blooms May 8 – June 12 in Maryland and Virginia with a peak May 18-20. Peak bloom period of S. purpurea may have shifted as much as a week from historical dates, perhaps due to climate change. Purple pitcher plant soils in Maryland and Virginia met expected conditions of low pH (3.5–4.9) and were low in almost all macro- and micro-nutrients. Perturbed or polluted sites exhibited elevated levels of exchangeable cations magnesium, calcium, and sodium. Climatic data disclosed that southern Virginia purple pitcher plant sites are both warmer and wetter than those in Maryland. Maryland pitcher plant bogs had greater species richness than Virginia bogs but the latter had more state rare plants. iv Copyright, 2010, by Philip M. Sheridan, All Rights Reserved. The author at Wakefield bog, Sussex County, VA in 1985 v This dissertation is dedicated to Margie Sheridan. “Indeed, it has been quite a pleasure to have experienced and studied these [cedar swamps and bogs] ecologically significant sites. Local and State governments should immediately institute a truly effective effort, perhaps acquisition, including appropriate buffer areas, to preserve their natural integrity. Indeed, I am convinced that society should demand it.” Bill Sipple, Days Afield. vi ACKNOWLEDGMENTS Thanks to Drs. Peter and Genevieve Sheridan for all their help and to my loving wife Margie Sheridan. My committee has also provided the support and encouragement to see this project through and I thank them for their assistance. I would also especially like to thank the USDA, Dr. Robert Griesbach, and USDA staff for help, supplies, and equipment to complete the molecular work in the lab in Beltsville, Maryland. I would like to thank the landowners who gave me permission to access their property for field surveys and to those individuals who helped procure samples of Sarracenia flowers from distant locations including Louise Kern (US Forest Service Florida), Adam Knatkovich (West Virginia and Pennsylvania), Dave Evans (New Jersey), Matt Opel (Connecticut), Jamie Dozier (South Carolina). Additional thanks are due to the following: Jim Solomon of the Missouri Botanical Garden; Rob Naczi of the New York Botanical Garden; Beth Chambers of William and Mary Herbarium; John Hayden of the University of Richmond; Emily Wood and Carolyn Beans of Harvard University Herbaria; Bill McAvoy of the Delaware Natural Heritage Program; Darren Loomis, Johnny Townsend, and Chris Ludwig of the Virginia Natural Heritage Program; Kathryn McCarthy, Chris Frye, and Jason Harrison of the Maryland Natural Heritage Program; Ron Beck and Charity James of USDA; Dr. Paul Chu of A&L Labs Inc.; Larry Brown of NOAA; Robert Wright; Jim Robinson; Joan Walker. Bill Scholl and his parents, James and Betty Scholl, provided invaluable assistance in my early botanical work looking for Sarracenia in Virginia. Keith and Mary Underwood provided logistical support for Maryland efforts. I would also like to thank anyone whom I may have forgotten to mention who helped with this research. Thank you all very much! vii TABLE OF CONTENTS LIST OF TABLES ................................................................................................................... ix LIST OF FIGURES .................................................................................................................. x INTRODUCTION .................................................................................................................... 1 Ecology ......................................................................................................................... 2 Taxonomy and Genetics ............................................................................................... 4 Biogeography................................................................................................................ 5 Goals ............................................................................................................................. 6 STUDY AREAS ....................................................................................................................... 8 Site Characterization – Maryland ................................................................................. 8 Site Characterization – Virginia ................................................................................. 10 A CENSUS OF PURPLE PITCHER PLANT, SARRACENIA PURPUREA L., IN MARYLAND AND VIRGINIA ................................................................................. 12 Introduction ................................................................................................................. 12 Materials and Methods ............................................................................................... 13 Results ......................................................................................................................... 16 Discussion ................................................................................................................... 23 Extinction Watch ............................................................................................ 23 Mechanisms for purple pitcher plant persistence in the wild ........................ 25 Climate change effects and the subspecies question ..................................... 28 GENETIC ANALYSIS OF PURPLE PITCHER PLANT, SARRACENIA PURPUREA L., SUBSPECIES UTILIZING THE CHALCONE SYNTHASE INTRON ........................................................................................................... 32 Introduction ................................................................................................................. 32 Materials and Methods ............................................................................................... 35 Plant Material ................................................................................................. 35 DNA Isolation ................................................................................................ 37 PCR amplification .......................................................................................... 37 Restriction analysis ........................................................................................ 38 Results ......................................................................................................................... 39 Comparison of outgroups ............................................................................... 39 Comparison within S. purpurea ..................................................................... 44 Discussion ................................................................................................................... 47 SOILS AND VEGETATION OF PURPLE PITCHER PLANT, SARRACENIA PURPUREA L., SITES IN MARYLAND AND VIRGINIA ....................... 54 Introduction ................................................................................................................. 54 Materials and Methods ............................................................................................... 57 viii
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