Community Structure, Plant Interactions, Seedling
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COMMUNITY STRUCTURE, PLANT INTERACTIONS, SEEDLING PERFORMANCE AND SEED BANK COMPOSITION OF SALT MARSHES ALONG AN ESTUARINE GRADIENT IN COOS BAY, OREGON by HOLLY BARTON KEAMMERER A DISSERTATION Presented to the Department of Biology and the Graduate School of the University of Oregon in partial fulfillment of the requirements for the degree of Doctor of Philosophy March 2011 DISSERTATION APPROVAL PAGE Student: Holly Barton Keammerer Title: Community Structure, Plant Interactions, Seedling Performance and Seed Bank Composition of Salt Marshes along an Estuarine Gradient in Coos Bay, Oregon This dissertation has been accepted and approved in partial fulfillment of the requirements for the Doctor of Philosophy degree in the Department of Biology by: Dr. Scott Bridgham Chairperson Dr. Richard Emlet Advisor Dr. Steven Rumrill Member Dr. Alan Shanks Member Dr. Gregory Retallack Outside Member and Richard Linton Vice President for Research and Graduate Studies/Dean of the Graduate School Original approval signatures are on file with the University of Oregon Graduate School. Degree awarded March 2011 ii © 2011 Holly Barton Keammerer iii DISSERTATION ABSTRACT Holly Barton Keammerer Doctor of Philosophy Department of Biology March 2011 Title: Community Structure, Plant Interactions, Seedling Performance and Seed Bank Composition of Salt Marshes along an Estuarine Gradient in Coos Bay, Oregon Approved: _______________________________________________ Dr. Richard Emlet Salt marshes are intertidal communities dominated by halophytic vascular plants that are subjected periodically to tidal inundation. These species have developed various adaptations to this stress, including tolerances of fluctuating salinity, extended periods of inundation and intervals of anoxic conditions. The marshes are divided into zones of different plant communities based on species’ tolerances of ambient estuarine conditions. Abiotic stresses change along the estuarine salinity gradient (marine to riverine), potentially altering development and composition of plant communities. Abiotic gradients associated with tides are not the only factors that contribute to development of plant community composition in salt marshes. Both negative (competition) and positive (facilitation) biological interactions are also important. Factors that influence community structure in salt marshes, particularly on the eastern North American seaboard, have been well studied. In contrast, salt marshes along the Oregon coast are smaller and more discrete and have received comparatively little attention. The community structure and seed bank composition of six marshes along an estuarine salinity gradient were evaluated. Four major community types dominated iv marshes that varied in the salinity of inundating tidal waters. Community types were relatively consistent throughout the estuary despite the distances between the marshes. Unlike the emergent plant communities, marsh seed bank composition was more similar within a marsh than within a community type. The low and high marsh community types were separated by a distinct boundary in the marine marshes. Although abiotic factors influence the physical separation of communities, competitive interactions commonly determine the upper limit of a species. In Metcalf marsh, however, the upper boundary for two dominant low marsh species was not determined by competition with the high marsh dominant species. Positive biotic interactions between seedlings and existing vegetation in a community are important factors in determining species distributions, particularly in stressful estuarine environments. In salt marshes, where abiotic stress can be harsh, presence of existing vegetation can ameliorate these conditions and enhance germination and seedling establishment. However, interaction between seedlings and the emergent marsh community was highly competitive, though germination of one species was enhanced in the presence of existing vegetation. This dissertation includes un-published co-authored material. v CURRICULUM VITAE NAME OF AUTHOR: Holly Barton Keammerer GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED: University of Oregon, Eugene, OR Pomona College, Claremont, CA DEGREES AWARDED: Doctor of Philosophy, Biology, 2011, University of Oregon Bachelor of Arts, Biology, 2006, Pomona College AREAS OF SPECIAL INTEREST: Plant Community Ecology Coastal Ecology Restoration Ecology Plants in Extreme Environments PROFESSIONAL EXPERIENCE: NSF GK-12 Fellow, Oregon Institute of Marine Biology, 2006-2008 Supervisors: Janet Hodder, Patricia Mace, Alan Shanks Teaching Assistant, University of Oregon, 2008-2010 Supervisors: Richard Emlet, Nora Terwilliger, B. Frank Eames, Ann Petersen Research Assistant, University of Oregon, 2009-2010 Supervisor: Richard Emlet Administrative Outreach Coordinator, Oregon Institute of Marine Biology, 2010 Supervisors: Joyce Croes, Janet Hodder, Craig Young vi GRANTS, AWARDS, AND HONORS: Graduate Research Fellowship, National Estuarine Research Reserve System, 2010-2011 Best Graduate Student Presentation, Pacific Estuarine Research Society Conference, 2010 vii ACKNOWLEDGMENTS First, I want to thank my advisor, Dr. Richard Emlet, not only for guidance and encouragement along this path, but for allowing me to follow my interests, wherever they led me. I am also extremely grateful to Dr. Sally Hacker for providing a botanical sounding board, for her advice and for her confidence in me. Thanks also to Dr. Steve Rumrill, who was always willing to talk and was always interested to hear about the current trials and tribulations of my research, Dr. Scott Bridgham for the use of his laboratory equipment and his advice, particularly regarding ecological analysis and Drs. Alan Shanks and Greg Retallack for their support and guidance through this process. I am also appreciative of the continual encouragement of Dr. Jonathan Wright. I also have to thank the faculty, staff, and graduate students at OIMB. In particular, Shirley Pedro who always cheered me up, Paul Dunn, who jumped with me through the hoops of the first two years of graduate school and Stephanie Schroeder, who has been my dissertation-writing confederate over the past few months. I am grateful also to Craig Cornu who contributed considerable time and to SSNERR for lending equipment while establishing marsh plot elevations and water level logger elevations. I am indebted to Dr. Dave Buckner who generously loaned me an optical sighting device, Dr. Laurel Pfeiffer-Meister for her statistical advice, Heidi Harris for her amazing map-making abilities, and J.K. Rowling and Jim Dale, who kept me company and kept me entertained though many solitary hours in the field and laboratory. I am so thankful for my amazing friends, both here and elsewhere, who were always willing to listen and who had the utmost confidence in me; in particular Stacy Galleher, Ali Helms, Jarret Roberts, Tushani Illangasekare, Angela Nierman Bahns, Mei viii Groth, JD Barton, Jane Valerius and Ann Kuenstling. Thanks also to Angie Kemp and Jonathan Bates who kept me fit and sane. And especially Kay Collins who was always happy to go for a run or share a glass, or two, of wine; she made my life here infinitely happier. And, of course, I am eternally grateful to my amazing parents; my mother who helped me in the field and my father who talked me through the rough patches of writing this dissertation. They always listened, encouraged and believed in me. This research was partially funded by a National Science Foundation GK-12 grant to Drs. Jan Hodder and Alan Shanks at OIMB, by a grant from the National Science Foundation, OCE 0527139 to Dr. Craig Young (PI; OIMB), and co-PIs Drs. Richard Emlet (OIMB), Michelle Wood (University of Oregon) and Will Jaeckle (Illinois Wesleyan University). The research was also partially funded by a graduate research fellow award from the Estuarine Reserves Division, Office of Ocean and Coastal Resource Management, National Ocean Service, National Ocean and Atmospheric Administration. ix This dissertation is dedicated to my parents, who instilled in me a love of botany. And to my sister, Linnaea, who was always willing to let me tag along on her rambling, woodland adventures. x TABLE OF CONTENTS Chapter Page I. GENERAL INTRODUCTION ................................................................................ 1 Abiotic Influence on Salt Marsh Plant Distribution .............................................. 1 Competition and Facilitation in Salt Marshes ........................................................ 3 Seed Banks in Salt Marshes ................................................................................... 4 Scope and Objectives ............................................................................................. 5 II. ENVIRONMENTAL FACTORS AND SALT MARSH COMMUNITY STRUCTURE ALONG AN ESTUARINE SALINITY GRADIENT ........................ 8 Introduction ............................................................................................................ 8 Methods.................................................................................................................. 10 Site Descriptions .............................................................................................. 10 Vegetation and Environmental Sampling ........................................................ 12 Statistical Analyses .......................................................................................... 15 Community Analysis ......................................................................................