Factors Affecting the Abundance of a Declining Grassland Bird: Implications for Recovery Strategy Planning and Implementation by Danielle Marie Ethier A Thesis presented to The University of Guelph In partial fulfillment of the requirements for the Degree of Doctor of Philosophy in Integrative Biology Guelph, Ontario, Canada © Danielle Marie Ethier, August 2016 ABSTRACT FACTORS AFFECTING THE ABUNDANCE OF A DECLINING GRASSLAND BIRD: IMPLICATIONS FOR RECOVERY STRATEGY PLANNING AND IMPLEMENTATION Danielle M. Ethier Advisors: Dr. Thomas Nudds University of Guelph, 2016 Dr. Steve Crawford Uncertainty is pervasive in all conservation decisions, the systematic treatment of which is necessary when evaluating the causes of species endangerment and deciding on appropriate actions to better ensure persistence. In Canada, a guiding principal of endangered species legislation for reducing uncertainty is adaptive management; however, adaptive management is not explicit in the development and application of most recovery strategies. To set the context for adaptive management recovery programming for at-risk species in Ontario, Canada, I synthesized existing knowledge, articulate and test critical uncertainties as hypotheses, and begin model development to predict expected outcomes of management alternatives, using the geo- politically defined population of Bobolink (Dolichonyx oryzivorus) in Ontario as a case study. Specifically, in Chapter 1, I combine evidence from empirical studies and expert judgement to model causal mechanisms driving Bobolink abundance dynamics and characterize the structural complexity of the management system. In Chapter 2, I spatially resolved Bobolink abundance trends to determine which areas of the province are contributing most substantially to overall abundance decline, and which presently satisfy recovery targets. In Chapter 3, I examined spatio- temporal variability in landscape pattern and processes to determine by which mechanism(s), and to what extent, changes in quantity and quality of agricultural grassland habitats have contributed to regional changes in abundance. Finally, in Chapter 4, I developed regionally scaled habitat suitability index (HSI) models to facilitate development of predictive population viability models. HSI models were used to evaluate to what extent lower-order proxies for processes of habitat selection influence patterns in abundance regionally. My research highlights how a science-based approach to recovery strategy planning can more effectively identify and address uncertainties in knowledge, and be used as an avenue to set the foundation of an adaptive management program. iv Dedication In loving memory of my Old Man, Richard L. Ethier 1945-2015 v Acknowledgments Foremost, I thank my husband, Josh Sayers, for his unwavering support throughout my seemingly endless academic pursuits, and our daughter, Annica, who reminds me every day that there is much more to life than research. My deepest gratitude to my advisor Dr. Thomas Nudds, who agreed to take on one last PhD student at the beginning of his retirement. He has taught me the importance of asking the first question first, and being an advocate for science-based decision making for species-at-risk. My co-advisor, Dr. Steve Crawford, offered timely advice and helpful comments to improve the clarity of my thesis. My committee member, Dr. Nicola Koper, offered statistical guidance, editorial support, and cheerful encouragement throughout my PhD. I thank the past and current members of the Nudds, Norris, Newman, McAdam, McCann, and Fryxell labs for their friendship and research support. I am especially grateful to my office mates (Laura Trout, Rachael Derbyshire, Grace Pittman, Alex Sutton, and Samantha Knight) and lab mates (Van La, Allan Derbertin, Zan Zhang, and Kevin Reid) for making every day at the office fun and inspiring. I acknowledge the countless volunteers involved in collecting North American Breeding Bird Survey and Ontario Breeding Bird Atlas data, including their official sponsors. I thank the anonymous stakeholders that participated in the elicitation workshop and online questionnaire. Mike Brienesse and Kevin Reid provided guidance on stakeholder elicitation and development of Bayesian Belief Networks. This research was funded by the Ontario Ministry of Natural Resources and Forestry, Natural Sciences and Engineering Research Council of Canada (NSERC) post-graduate scholarship, University of Guelph Dean’s Tri-council Scholarship, and an I.O.D.E. War Memorial Doctorate Scholarship. vi Table of Contents Abstract ........................................................................................................................................... ii Acknowledgments........................................................................................................................... v List of Tables ................................................................................................................................. ix List of Figures ................................................................................................................................. x Prologue .......................................................................................................................................... 1 Thesis Goal .................................................................................................................................. 3 Background ................................................................................................................................. 5 Agricultural Grasslands ........................................................................................................... 5 Grassland Birds........................................................................................................................ 6 Bobolink (Dolichonyx oryzivorus) .......................................................................................... 6 Endangered Species Act (2007) .............................................................................................. 8 Exemption Regulation ............................................................................................................. 9 Recovery Goals, Objectives and Actions ................................................................................ 9 Literature Cited ......................................................................................................................... 11 Figures ....................................................................................................................................... 15 Co-authorship and Chapter Publication Status ............................................................................. 16 Chapter 1: Complexity of Factors Affecting Bobolink Population Dynamics Communicated with Directed Acyclic Graphs ............................................................................................................... 17 Abstract ..................................................................................................................................... 17 Introduction ............................................................................................................................... 18 Methods ..................................................................................................................................... 20 Results ....................................................................................................................................... 21 1. Habitat Quantity ............................................................................................................. 21 1.1 Breeding habitat quantity ......................................................................................... 22 1.2 Non-breeding habitat quantity .................................................................................. 23 2. Habitat quality ................................................................................................................ 24 2.1 Breeding habitat quality ........................................................................................... 24 2.1.1 Landscape condition .......................................................................................... 24 2.1.2 Patch condition .................................................................................................. 25 2.1.3 Social information ............................................................................................. 26 2.2 Non-breeding habitat quality .................................................................................... 27 vii 3. Fecundity........................................................................................................................ 27 3.1 First hay harvest date................................................................................................ 28 3.2 Livestock trampling.................................................................................................. 29 3.3 Nest predation and parasitism .................................................................................. 30 3.4 Pesticides .................................................................................................................. 31 3.5 Prey availability........................................................................................................ 31 4. Mortality ........................................................................................................................ 32 4.1 Breeding period mortality ........................................................................................ 32 4.1.1 Date of first harvest ........................................................................................... 33 4.1.2 Pesticides ........................................................................................................... 33 4.1.3 Collisions with tall structures ............................................................................ 33 4.2 Non-breeding period mortality ................................................................................. 33 4.2.1 Pesticides ........................................................................................................... 34 4.2.2 Persecution......................................................................................................... 34 States of Published and Stakeholder Knowledge ...................................................................... 34 Discussion ................................................................................................................................. 35 Tables ........................................................................................................................................ 48 Figures ....................................................................................................................................... 49 Chapter 2: Scalar considerations in population abundance trend estimates: implications for recovery strategy planning for species of conservation concern .................................................. 53 Abstract ..................................................................................................................................... 53 Introduction ............................................................................................................................... 54 Methods ..................................................................................................................................... 57 Study area........................................................................................................................... 57 Spatially explicit trend estimation ..................................................................................... 57 Results ....................................................................................................................................... 61 Discussion ................................................................................................................................. 62 Literature cited .......................................................................................................................... 67 Figures ....................................................................................................................................... 71 Chapter 3: Spatio-temporal variation in mechanisms driving regional-scale abundance patterns of Bobolink (Dolichonyx oryzivorous) .............................................................................................. 74 Abstract ..................................................................................................................................... 74 Introduction ............................................................................................................................... 76 viii Methods ..................................................................................................................................... 78 Study area........................................................................................................................... 78 Response variable .............................................................................................................. 78 Predictor variables ............................................................................................................. 80 Analysis.............................................................................................................................. 82 Results ....................................................................................................................................... 84 Spatio-temporal Bobolink trend estimates ......................................................................... 84 Spatio-temporal landscape change ..................................................................................... 85 Bayesian variable selection ................................................................................................ 86 Discussion ................................................................................................................................. 87 Literature cited .......................................................................................................................... 93 Tables ........................................................................................................................................ 98 Figures ..................................................................................................................................... 100 Chapter 4: Local habitat selection by grassland birds scales to regional extents: implications for landscape-level conservation planning ....................................................................................... 105 Abstract ................................................................................................................................... 105 Introduction ............................................................................................................................. 106 Methods ................................................................................................................................... 108 Results ..................................................................................................................................... 112 Discussion ............................................................................................................................... 112 Literature cited ........................................................................................................................ 117 Figures ..................................................................................................................................... 121 Epilogue ...................................................................................................................................... 123 Literature cited ........................................................................................................................ 127 Supplemental Material ................................................................................................................ 128 Appendix A (Chapter 1).............................................................................................................. 128 Appendix B (Chapter 2) .............................................................................................................. 131 Appendix C (Chapter 2) .............................................................................................................. 135 Appendix D (Chapter 3).............................................................................................................. 136 Appendix E (Chapter 4) .............................................................................................................. 143 ix List of Tables Chapter 1 Table 1: Eight vegetation variables were identified as important predictors of Bobolink (Dolichonyx oryzivorus)................................................................................................................ 48 Chapter 3 Table 1: Agricultural census divisions in Ontario, Canada which display strong evidence of Bobolink (Dolichonyx oryzivorous) population growth or decline over the entire study period (1986-2011) and five sub-periods.. ............................................................................................. 988 x List of Figures Prologue Figure 1: Relative abundance and distribution map for Bobolink (Dolichonyx oryzivorous) based on North American Breeding Bird Survey (BBS) data from 1987-2006 (Blancher 2009).. ........ 15 Chapter 1 Figure 1: Schematic diagram of a conceptual two-season life cycle model for Bobolink (adapted from Pulliam and Danielson 1991). . ............................................................................................ 49 Figure 2: Directed acyclic graphs (DAGs) showing the causal relationships (links and arrows) among variables (ovals) hypothesized to be driving changes in Bobolink habitat quantity on the breeding and non-breeding grounds based on stakeholder elicitation and empirical evidence from the literature.. ................................................................................................................................ 52 Figure 3: Directed acyclic graphs (DAGs) showing the causal relationships (links and arrows) among variables (ovals) hypothesized to be driving changes in Bobolink habitat quality on the breeding and non-breeding grounds based on stakeholder elicitation and empirical evidence from the literature.. ................................................................................................................................ 53 Figure 4: Directed acyclic graphs (DAGs) showing the causal relationships (links and arrows) among variables (ovals) hypothesized to be driving changes in Bobolink fecundity and mortality on the breeding and non-breeding grounds based on stakeholder elicitation and empirical evidence from the literature.. ........................................................................................................ 54 Chapter 2 Figure 1: Relative abundance and distribution map for Bobolink (Dolichonyx oryzivorous) based on North American Breeding Bird Survey (BBS) data from 1987-2006 (Blancher 2009).. ........ 71 Figure 2: Posterior mean population trend estimates for Bobolink (Dolichonyx oryzivorous) from A. 1967-2011 and B. 1998-2011 mapped to agricultural census divisions in southern Ontario, Canada, using spatially explicit hierarchical models in a Bayesian statistical framework.. ......... 72 Figure 3: Box plots of posterior mean trends (solid centre line in box) for Bobolink (Dolichonyx oryzivorous) from A. 1967-2011 and B. 1998-2011 shown across 35 agricultural census divisions in Ontario, Canada (ordered from west to east).. .......................................................................... 73 Chapter 3 Figure 1: Directed acyclic graph showing causal relationships (link and arrows) among variables (ovals) hypothesized to account for population declines in Bobolink (Dolichonyx oryzivorous) (diamond) based on evidence in the published literature and/or stakeholder consultation. ....... 100
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