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PLANT-POLLINATOR NETWORK ASSEMBLY AFTER WILDFIRE by Michael Peter Simanonok A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Ecology & Environmental Science MONTANA STATE UNIVERSITY Bozeman, Montana August 2018 ©COPYRIGHT by Michael Peter Simanonok 2018 All Rights Reserved ii DEDICATION To Stacy for your endless love and patience through this entire ordeal. And to Geoff, I’ll see what I can do about carrying that fire. iii ACKNOWLEDGEMENTS First I need to thank Laura for giving me the freedom and range to do just about anything I wanted with developing and running these projects over the years. You’ve made this PhD process as smooth as it can be, and I don’t think I would have even made it to this point under anyone else’s guidance. Everyone who’s helped on this project: data collection, entry, ID, etc. There are entirely too many to list on this single page, but especially “Zab” Reese for the absolutely unbelieveable amount of work you’ve put in behind the scenes in the lab, you’ve saved us all, and Dylan Cole for saving one of my field seasons. Also, thanks to Casey Delphia for additional identification help and for lending the nesting tubes, and to Jane Klaussen for your chemistry wizardry. All my friends who’ve passed through the basement of AJMJ, it’s been an honor to have been a part of this community for so long. Especially those of y’all from the Burkle lab over the years: Stacy Davis, Anthony Slominski, Laura Heil, Will Glenny, and Simone Durney. And, our founding office cohort: Stacy Davis, Dan Bachen, Adam Mitchell, and Erin Kenison. Funding for these projects was provided by the National Science Foundation, Montana State University, the Joint Fire Science Program, and the Montana Native Plant Society. iv TABLE OF CONTENTS 1. INTRODUCTION ...........................................................................................................1 2. PLANT-POLLINATOR NETWORK MOTIFS AND SPECIES’ ROLES DO NOT CHANGE WITH BURN SEVERITY OR TIME-SINCE-BURN ................................................................................................4 Introduction ......................................................................................................................4 Methods..........................................................................................................................11 Field Sampling .......................................................................................................11 Analyses .................................................................................................................12 Results ...........................................................................................................................15 Discussion .....................................................................................................................17 3. HIGH-SEVERITY WILDFIRE LIMITS BUMBLEBEE NUTRITION AND AVAILABLE FLORAL POLLEN QUALITY COMPARED TO MIXED-SEVERITY BURNS .......................................33 Introduction ....................................................................................................................33 Methods..........................................................................................................................37 Study System .........................................................................................................37 Field Sampling .......................................................................................................38 Lab Methods ..........................................................................................................39 Statistical Analyses ................................................................................................39 Results ...........................................................................................................................40 Quality of Available Floral Pollen .........................................................................40 Quality of Corbicular Pollen ..................................................................................41 Bumblebee Nutrition ..............................................................................................41 Discussion .....................................................................................................................41 Quality of Available Pollen ...................................................................................42 Bumblebee Nutrition ..............................................................................................43 Disturbance & The Nutritional Landscape ............................................................44 4. NESTING SUCCESS OF WOOD-CAVITY-NESTING BEES AFTER WILDFIRE DECLINES WITH INCREASING TIME-SINCE-BURN BUT IS UNAFFECTED BY BURN SEVERITY ........................................................................................................51 Introduction ....................................................................................................................51 Methods..........................................................................................................................56 Study Site ...............................................................................................................56 Field Sampling .......................................................................................................56 v TABLE OF CONTENTS CONTINUED Statistical Analysis .................................................................................................58 Results ...........................................................................................................................58 Discussion .....................................................................................................................60 5. CONCLUSIONS............................................................................................................70 REFERENCES CITED ......................................................................................................73 APPENDICES ...................................................................................................................80 APPENDIX A: Species Lists .................................................................................80 vi LIST OF TABLES Table Page 1. Fire Locality Information ...................................................................................25 2. Bee Species Composition .................................................................................25 3. Floral Species Composition ...............................................................................26 4. Network Motif Identity and Frequency Analysis .............................................26 5. Species’ Roles of Shared Species Analysis .......................................................27 6. High-Severity Species’ Roles Analysis ............................................................28 7. Mixed-Severity Species’ Roles Analysis ...........................................................29 8. Structural Equation Modelling Nesting Results ...............................................66 A1. Bee Species Sampled Via Hand Netting ............................................................81 A2. Floral Species Sampled Along Floral Census Transects ..................................94 A3. Bumble Bees Used for Nutritional Analyses ...................................................108 A4. Visitation for Bumble Bee Nutrition ...............................................................110 A5. Floral Pollen Samples ......................................................................................114 A6. Emerged Nesting Tubes by Species ................................................................116 vii LIST OF FIGURES Figure Page 1. Conceptual Network Motif Figure .....................................................................29 2. Bee and Floral Abundance and Species Richness .............................................30 3. Bee and Floral Species Turnover .......................................................................31 4. Interaction Turnover ..........................................................................................32 5. Available Floral Pollen Quality .........................................................................47 6. Corcibuclar Pollen Quality ................................................................................48 7. Available Floral Pollen Quality vs. Corbicular Pollen Quality .........................49 8. Bumblebee Nutrition ..........................................................................................50 9. Nesting Box Example ........................................................................................67 10. Structural Equation Modeling Hypothetical Pathways .....................................68 11. Structural Equation Modeling Results ..............................................................69 viii ABSTRACT Plant-pollinator networks are threatened by anthropogenic influence due to habitat loss, changing fire regimes, climate change and other factors. Furthermore, we have little current knowledge for how species interactions and processes like pollination assemble and recover post-disturbance. Studying the mechanisms by which plant-pollinator interactions assemble in a post-disturbance landscape, particularly across gradients of disturbance