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Enemy of My Enemy: Can the Rhizosphere Biota of Vincetoxicum Rossicum Act As Its “Ally” During Invasion?

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Enemy of My Enemy: Can the Rhizosphere Biota of Vincetoxicum Rossicum Act As Its “Ally” During Invasion? Enemy of my Enemy: Can the Rhizosphere Biota of Vincetoxicum rossicum Act as its “Ally” During Invasion? by Angela Dukes A Thesis presented to The University of Guelph In partial fulfillment of the requirements for the degree of Master of Science in Environmental Biology Guelph, Ontario, Canada © Angela Dukes, November 2017 ABSTRACT Enemy of my Enemy: Can the Rhizosphere Biota of Vincetoxicum rossicum Act as its “Ally” During Invasion? Angela Dukes Advisors: Dr. Pedro Antunes University of Guelph, 2017 Dr. Kari Dunfield The ‘Enemy of my enemy’ (EE) is a major hypothesis in invasion ecology. It states that a non-native invader ‘accumulates generalist pathogens, which limit competition from indigenous competitors’. Few empirical studies have tested the EE hypothesis in plant invasions, especially on biotic rhizosphere interactions. Here, the EE hypothesis was tested by applying rhizosphere biota from the invasive plant Vincetoxicum rossicum (VIRO) to five co-occurring native plant species, and four native legume species, respectively. Each of the native plant species, and VIRO were grown under controlled conditions for three months, either in presence or absence of soil biota from VIRO invaded and non-invaded soils. Rhizosphere biota from invaded areas had variable effects among native plants (including legumes). It was concluded that the accumulation of rhizosphere enemies that ‘spill’ onto native plants may not be a major factor in the invasive success of VIRO. The EE hypothesis was not supported. iii ACKNOWLEDGEMENTS I deeply appreciated the patience of my supervisors: Dr. Kari Dunfield and Dr. Pedro Antunes. I worked in the Plant and Soil Ecology Lab at Algoma University with Dr. Pedro Antunes. He stood by me, affording me the time to take this journey. I had a steep learning curve, learning the world of soil biota, learning to write at a higher level, and to think more critically. But throughout I had his confidence – and that meant a lot. Pedro also allowed me time to take some tangents, working as a TA, and volunteering with several departmental workshops providing environmental education for children. He held me to a high level of achievement, but also encouraged me, guided me, was a confidant and friend – for this he was one of the best teachers I have ever had. Thank you for your time and confidence Pedro. This work would not have been possible without the assistance of Dr. Akihiro Koyama, our Post Doc in the Plant and Soil Ecology Lab. He conducted the bootstrap analysis in R for this thesis. With these outputs I was able to analyse net biotic effect of Vincetoxicum rossicum on native plant species. Aki also conducted the phylogenetic analysis component of this thesis. I am overwhelmingly grateful to my long-suffering family. They allowed me to defer my full responsibilities as a mom, they stood by me, and encouraged me. I am blown away at how fast my little boys have grown up! And honored to have a husband that so thoroughly nourished our children through my countless weekends away. Many thanks to the Soil and Plant Ecology Lab at Algoma University: Undergraduate students (Kyle Snape, Alecia Finateri, Derissa Vincentini), and Lab coordinator Meaghan Sutherland. Your help in planting and harvest were much appreciated. I am grateful to the staff at the Ontario Forest Research Institute, particularly Darren Derbowka, and Keven Maloney for their suggestions and support in the use of the growth facilities. I deeply appreciate the assistance of Susan Meades, curator of the Northern Ontario Plant Database, for her help with plant identification. Finally, I owe a big thank-you to the staff of the Central Lake Ontario Conservation Authority (Jamie Davidson), Ganaraska Region Conservation Authority, (Ken Towle), Orono Crown Lands (John Standeven), Toronto Zoo (Elizabeth Heighington) and Ontario Ministry of Natural Resources (Bohdan Kowalyk), for your support and access to field sites. The work described in the following pages is the result of funding through Ontario Graduate Scholarship awarded to myself, the NSERC Discovery Grant awarded to Dr. Pedro Antunes, and bursaries from the University of Guelph. iv TABLE OF CONTENTS ABSTRACT ................................................................................................................................................. ii ACKNOWLEDGEMENTS ........................................................................................................................ iii TABLE OF CONTENTS ........................................................................................................................... iv LIST OF TABLES ...................................................................................................................................... vi LIST OF FIGURES .................................................................................................................................. viii Chapter 1 Introduction 1.1 Invasion Biology ......................................................................................................................... 1 1.2 Plant-Biota Interactions and Their Role in Plant Invasions .................................................. 3 1.3 Facilitation of Invasion by Pathogen Accumulation............................................................... 7 1.4 Pathogen Accumulation in Vincetoxicum rossicum .............................................................. 9 1.5 Aim and Objectives .................................................................................................................. 10 Chapter 2 Response of Native Plants to Vincetoxicum rossicum Rhizosphere Biota 2.1 Abstract ...................................................................................................................................... 11 2.2 Introduction ................................................................................................................................ 13 2.3 Materials and Methods 2.3.1 Collection of Biotic Soil Communities ............................................................................ 17 2.3.2 Experimental Design and Growth Conditions .............................................................. 18 2.3.3 Harvest and Response Variables .................................................................................. 21 2.3.4 Statistical Analysis............................................................................................................ 22 2.4 Results 2.4.1 Effect of Invasion on Native Plant Species Biomass .................................................. 26 2.4.2 Analysis of Phylogenetic Signal in Net Biotic Effect ................................................... 30 2.4.3 Effect of Invasion on Native Plant Reproduction ......................................................... 32 2.4.4 Effect of Invasion on Nodule Formation in Desmodium canadense ........................ 33 2.4.5 Effects of Invasion on Incidence of Root Lesions ....................................................... 34 2.4.6 Effects of Invasion on Colonization by Arbuscular Mycorrhizal Fungi ..................... 36 2.5 Discussion 2.5.1 Does ‘Enemy of my Enemy’ Influence Vincetoxicum rossicum Invasion? .............. 39 2.5.2 Does Phylogenetic Relatedness Influence Response to Invasion? ......................... 44 2.5.3 Conclusions ....................................................................................................................... 45 v Chapter 3 Response of Native Legumes to Vincetoxicum rossicum Rhizosphere Biota 3.1 Abstract ...................................................................................................................................... 46 3.2 Introduction ................................................................................................................................ 48 3.3 Materials and Methods 3.3.1 Collection of Biotic Soil Communities ............................................................................ 52 3.3.2 Experimental Design and Growth Conditions .............................................................. 53 3.3.3 Harvest and Response Variables .................................................................................. 56 3.3.4 Statistical Analysis............................................................................................................ 56 3.4 Results 3.4.1 Effect of Invasion on Native Plant Species Biomass .................................................. 58 3.4.2 Effect of Invasion on Desmodium canadense Reproductive Potential .................... 63 3.4.3 Effect of Invasion on Nodule Formation ........................................................................ 65 3.5 Discussion 3.5.1 The Effect of Vincetoxicum rossicum Rhizosphere Biota on Native Legumes Productivity ....................................................................................................................................... 67 3.5.2 The Role of ‘Enemy of my Enemy’ in Vincetoxicum rossicum Invasion .................. 72 3.5.3 Conclusions ....................................................................................................................... 73 CONCLUSION .......................................................................................................................................... 74 REFERENCE LIST .................................................................................................................................. 77 APPENDICES Appendix A Coordinates for Soil Collection
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