Examining the Coexistence of Splachnaceae Mosses in Newfoundland Peatlands
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Examining the Coexistence of Splachnaceae Mosses in Newfoundland Peatlands by © Christopher Hammill A thesis submitted to the School of Graduate Studies in partial fulfillment of the requirements for the degree of Master of Science Biology Memorial University of Newfoundland April 2016 St. John’s Newfoundland and Labrador Abstract Coexistence amongst highly similar species has captured the imagination of many ecologists. In my work I've endeavoured to answer the question of how Splachnum ampullaceum and Splachnum pensylvanicum coexist in peatlands in Newfoundland. I've employed individual based simulation modelling in conjunction with machine learning to answer if coexistence is facilitated most by dispersal differences or by competitive similarity. Experimental work examined dispersal differences and competitive ability directly. I found that coexistence is facilitated primarily by temporal niche separation via differing phenology, and substrate availability. My experimental work shows spore dispersal is dependent on distance to nearest moss population with no evidence of species-specific differences. The competitive ability of the two species was shown to vary according to moisture with potential facilitation effects at high population densities. Together the results show that the two Splachnaceae are more dissimilar than they appear, with niche separation along temporal and hydrological axes. ii Acknowledgements I’d like to start off by thanking all of the people who made my work possible. I’d like to thank Nicole McMurray for being there with me every step of the way, and my friends and family for supporting me throughout the process and helping me to maintain my sanity. I’d like to thank Dr. Paul Marino for being an excellent guide, editor, and friend. To my supervisory committee, Drs. Suzanne Dufour, Amy Hurford, and Shawn Leroux: thanks for pushing me to accomplish more than I thought I could, and providing guidance all along the way. I couldn’t have done it without your help. I’d like to thank Dr. Tom Chapman for providing vital services: caffeine, mentorship, and encouragement when I was ready to give up. To the rotating cast of characters in the Chapman lab: Kate Basset, Holly Caravan, Andrew Chaulk, and Jake Coates, thanks for all the advice, assistance, comradery, friendship, laughs, and beer. I’m grateful to Dr. Ted Miller, Paul Regular, Alejandro Buren, and Dr. Lourdes Pena-Castillo for providing the right nudges to statistical computing, and helping me find my interests. I’m also grateful to the kind employees of Salmonier Nature Park for helping me get moose dung and allowing me use of their facilities throughout my field work. iii Contents Abstract ........................................................................................................................................................ 2 Acknowledgements ..................................................................................................................................... 3 Contents ........................................................................................................................................................ 4 List of Tables ............................................................................................................................................... 6 List of Figures .............................................................................................................................................. 9 List of Appendices ..................................................................................................................................... 11 Chapter 1: Grand Intro .............................................................................................................................. 1 References ................................................................................................................................................ 6 Co-Authorship Statement .......................................................................................................................... 8 Chapter 2: Modelling the coexistence of Splachnum ampullaceum and Splachnum pensylvanicum in Newfoundland Peatlands ............................................................................................................................ 9 Abstract ..................................................................................................................................................... 9 Intro........................................................................................................................................................... 9 Methods .................................................................................................................................................. 14 Ecology of Splachnum ampullaceum and Splachnum pensylvanicum: ............................................... 14 Empirical Parameter Estimates: .......................................................................................................... 15 Modelling Ideology: ............................................................................................................................ 16 Model Overview: ................................................................................................................................. 17 Spatial Considerations: ....................................................................................................................... 17 iv Space Competition .............................................................................................................................. 18 Spore Production and Transfer: .......................................................................................................... 19 Relative spore transfer rate: ............................................................................................................... 21 Phenology: .......................................................................................................................................... 22 Parameter Influence on Coexistence: ................................................................................................. 23 Influence of Conspecific Aggregation: ................................................................................................ 24 Role of Phenological Offset: ................................................................................................................ 27 Results ..................................................................................................................................................... 27 Empirical Parameter Estimates: .......................................................................................................... 27 Role of Attraction: ............................................................................................................................... 28 Simulation Outcomes: ......................................................................................................................... 28 Machine Learning and Variable Importances ..................................................................................... 29 Effect of Phenological Offset: ............................................................................................................. 31 Discussion................................................................................................................................................ 31 References .............................................................................................................................................. 37 Figures ..................................................................................................................................................... 40 Chapter 3: Examining Dispersal and Competitive Abilities of Splachnum ampullaceum and S. pensylvanicum............................................................................................................................................. 50 Abstract ................................................................................................................................................... 50 Introduction ............................................................................................................................................ 50 Methods .................................................................................................................................................. 53 v Splachnaceae Ecology: ........................................................................................................................ 53 Competition Experiment: .................................................................................................................... 54 Spore Movement: ............................................................................................................................... 56 Results ..................................................................................................................................................... 59 Spore Movement ................................................................................................................................ 59 Competition Experiment ..................................................................................................................... 60 Discussion...............................................................................................................................................