Smooth Brome Invasion Influences Nitrogen Cycling and Soil Bacterial Community Structure in a Fescue Grassland
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Smooth brome invasion influences nitrogen cycling and soil bacterial community structure in a fescue grassland A Thesis Submitted to the College of Graduate Studies and Research In Partial Fulfillment of the Requirements For the Degree of Master of Science In the Department of Plant Sciences University of Saskatchewan Saskatoon By Candace Lynn Piper Copyright, Candace Piper, May 2013. All Rights Reserved. Permission to Use In presenting this thesis in partial fulfillment of the requirements for a graduate degree from the University of Saskatchewan, I agree that the Libraries of this University may make it freely available for inspection. I further agree that permission for copying of this thesis in any manner, in whole or in part, for scholarly purposes may be granted by the professors who supervised my thesis work or, in their absence, by the Head of the Department or the Dean of the College in which my thesis work was done. It is understood that any copying or publication of use of this thesis or parts thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given to me and to the University of Saskatchewan in any scholarly use which may be made of any material in my thesis. Requests for permission to copy or to make other use of material in this thesis in whole or part should be addressed to: Head of the Department of Plant Sciences 51 Campus Drive University of Saskatchewan Saskatoon, Saskatchewan S7N 5A8 i Abstract Exotic plant invasions represent a significant threat to the integrity of native grasslands. Across the Northern Great Plains, grasslands invaded by smooth brome (Bromus inermis Leyss) support lower plant diversity, potentially resulting in important consequences for ecosystem function. Previous research on smooth brome has primarily focused on aboveground changes in plant communities, but there is growing evidence that the soil ecosystem can be significantly altered with invasion. The two objectives of this thesis were to examine whether smooth brome invasion alters soil nitrogen cycling, and to determine if changes in plant community diversity or productivity influence soil bacterial communities. Relationships between smooth brome and the soil ecosystem were assessed using data collected from a Festuca hallii Vasey (Piper) (plains rough fescue) grassland located near Macrorie, SK. Gross rates of nitrogen cycling and community productivity from smooth brome invaded and native grassland sites were compared to determine the potential influence of smooth brome invasion on the soil nitrogen cycle. The relationship between increasing smooth brome abundance and soil bacterial structure and composition was also studied. Gross mineralization rates and total soil nitrogen were significantly higher in smooth brome‐invaded areas relative to native grassland. Bacterial and archaeal amoA, used as indicators of ammonia‐ oxidizer population sizes, were altered by smooth brome cover. Higher gross mineralization rates were likely due to stimulated microbial activity caused by increased litter and root production in areas invaded by smooth brome. Smooth brome decreased plant species richness through increased litter production, but had ii the opposite effect on bacterial communities. Bacterial communities had higher species richness and evenness in soils invaded by smooth brome, and smooth brome invasion was also associated with bacteria important for soil nitrogen cycling. As bacteria dominate microbial biomass and are important for decomposition processes, a more even bacterial community may have supported increased mineralization rates in smooth brome‐invaded soils. Specifically, a more even bacterial community may have increased mineralization rates through greater resource utilization and niche partitioning. The responses observed in these studies suggest that belowground changes with smooth brome invasion have the potential to have important consequences for ecosystem processes. iii Acknowledgements Thank you to my supervisors, Dr. Eric Lamb and Dr. Steven Siciliano, for your positive support, patient guidance, and for your ability to see the big picture. Your continued enthusiasm and commitment to science have not only improved my project but also my experience as a graduate student. Thank you to my advisory committee, Dr. Jim Romo, Dr. Yuguang Bai, and Dr. Bruce Coulman for your advice and feedback, and for asking the tough questions that ultimately improved this project. Special thank you to Dr. Romo for his help in selecting my field site and for sharing his knowledge and experience of smooth brome with me. This project required many helping hands, so a special thank you to Sarah Hardy, Amanda Guy, Sherri Friedrich and Curtis Wiens for assistance and companionship in both the lab and the field. Thank you to my fellow lab mates, Digit Guedo, Christiane Catellier, Jenalee Mischkolz and Mike Lavender for providing feedback and companionship as this project evolved. Thank you to the private landowner, James Thorpe, for allowing me access to the land that became my field site. Thank you to the Department of Plant Sciences, NSERC, and various scholarships provided through the Department and College of Agriculture for financial support. And finally, much thanks goes to my family and especially my husband, John, for their continued support and understanding throughout this academic pursuit. iv Table of Contents Smooth brome invasion influences nitrogen cycling and soil bacterial community structure in a fescue grassland ................................................................................................................................. 1 Permission to Use ..................................................................................................................................... i Abstract ....................................................................................................................................................... ii Acknowledgements ............................................................................................................................... iv Table of Contents ..................................................................................................................................... v List of Tables .......................................................................................................................................... vii List of Figures ........................................................................................................................................... ix List of abbreviations .............................................................................................................................. xi 1 Introduction ..................................................................................................................................... 1 1.1 Grassland Communities and Plant‐Soil Interactions .................................................................. 2 1.2 Invasive Species ............................................................................................................................................ 6 1.2.1 Invasive Species .............................................................................................................................. 6 1.2.2 Ecosystem susceptibility to invasion ..................................................................................... 7 1.2.3 Invasion Mechanisms ................................................................................................................... 9 1.3 Smooth brome (Bromus inermis Leyss) .......................................................................................... 13 1.4 Objectives ...................................................................................................................................................... 15 1.5 References ..................................................................................................................................................... 16 Chapter Preamble ................................................................................................................................ 24 2 Smooth brome alters soil nitrogen cycling processes in a Fescue Grassland ....... 25 2.1 Abstract .......................................................................................................................................................... 25 2.2 Introduction ................................................................................................................................................. 26 2.3 Methods .......................................................................................................................................................... 28 2.3.1 Field site .......................................................................................................................................... 28 2.3.2 Ammonia‐oxidizer and plant biomass study ................................................................... 29 2.3.3 Nitrogen cycling study .............................................................................................................. 31 2.3.4 Statistical analyses ...................................................................................................................... 34 2.4 Results ............................................................................................................................................................