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MULTI-SCALE FACTORS RELATED TO ABUNDANCE OF BATS AND INSECT PREY IN SAVANNAS, WOODLANDS, AND FORESTS IN THE OZARK HIGHLANDS, USA _______________________________________________________________________ A Dissertation Presented to The Faculty of the Graduate School At the University of Missouri-Columbia ________________________________________________________________________ In Partial Fulfillment Of the Requirements for the Degree Doctor of Philosophy ________________________________________________________________________ by KATHRYN MARIE WOMACK Dr. Frank R. Thompson III, Dissertation Supervisor MAY 2017 The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled MULTI-SCALE FACTORS RELATED TO ABUNDANCE OF BATS AND INSECT PREY IN SAVANNAS, WOODLANDS, AND FORESTS IN THE OZARK HIGHLANDS, USA Presented by Kathryn Marie Womack a candidate for the degree of Doctor of Philosophy and hereby certify that, in their opinion, it is worthy of acceptance. ____________________________________________ Professor Frank R. Thompson III ____________________________________________ Professor Matthew Gompper ____________________________________________ Dr. Sybill K. Amelon ____________________________________________ Professor Lori Eggert _____________________________________________ Professor Rose-Marie Muzika This would have not been possible without the love and support from my village. Special thanks to Niki, Ed, Jaymi, and Cara. ACKNOWLEDGEMENTS I would like to thank the many individuals without whose efforts this study would not have been possible. First, I would like to thank my advisor, Frank Thompson for all his support, advice, and patience throughout the years. I would also like to extend a special thanks to Sybill Amelon for taking a chance on me over a decade ago, and who brought me to Missouri. I would like to my dissertation committee: Matt Gompper, Sybill Amelon, Lori Eggert, and Rose-Marie Muzika. I would not have completed as many sites without the help and willingness to run trap at sites for 3 nights from Sarah Bradley, Nettie Sittingup, Linda Mills, Megan York-Harris, Clarissa Starbuck, Risa Wright, and Sybill Amelon. In addition, I could not have completed this project without the ArcGIS and statistical assistance from Bill Dijak and Jaymi LeBrun. Tom Bonnot and Julianna Jenkins supported me tremendously in helping me trouble shoot errors in R code, and provided an environment for me to increase my confidence in my statistical analysis skills. I would like to thank my other lab mates: Liz Matseur and Melissa Roach for being a support system, and providing encouragement during the final hours of the degree. I could not have finished with as much grace without their encouragement. Lastly, I would like to thank my family, both born with and chosen, for taking my 4 a.m. phone calls, and believing in me when I did not believe in myself. I love you and I hope you feel that this degree is shared among us. This research was funded by the USDA Forest Service Northern Research Station. ii Table of Contents ACKNOWLEDGEMENTS…………………………………………………...………...ii LIST OF ILLUSTRATIONS…………………………………………………….….…vii DESCRIPTION OF CHAPTERS………………………………………………..……xv DISSERTATION ABSTRACT………………………………………………….…....xvi CHAPTER 1: INTRODUCTION……………………………………………………….1 LITERATURE CITED ……………………………………………………..…………6 CHAPTER 2: PERFORMANCE OF HIERARCHICAL ABUNDANCE MODELS ON SIMULATED BAT CAPTURE DATA…….……………………………………...8 ABSTRACT…………………………………………………………………………8 1 INTRODUCTION…………………………………………………………………...9 2 METHODS……………………………………………………………………….14 2.1 Model descriptions and assumptions………………………………..…...14 2.2 Data simulation……………………………………………….……..…...14 2.3 Model performance………………………………………………….…...17 3 RESULTS……………………………………………………………………..…17 3.1 Scenario 1………………………………..……………………………....17 3.2 Scenario 2…………………………………………….……..…………...18 iii 3.3 Scenario 3………………………………………………….………….....18 3.4 Scenario 4………………………………………………….………….....19 4 DISCUSSION…………………………………………………………………….19 LITERATURE CITED……………………………………………………………..…23 CHAPTER 3: RESTORATION AND HABITAT FACTORS RELATED TO INSECT ABUNDANCE ACROSS A GRADIENT OF SAVANNAS, WOODLANDS, AND NON-MANAGED FORESTS IN THE OZARK HIGHLANDS, USA…….….38 ABSTRACT………………………………………………………………..………..38 1 INTRODUCTION…………………………………………………………………..39 2 METHODS…………………………………………………………………..……45 2.1 Study areas……………………………………………………….…..45 2.2 Study design……………………………………………………….…46 2.3 Insect surveys………………………………………………………...47 2.4 Insect processing protocol…………………………………………...48 2.5 Vegetation surveys and covariates………………………………...…49 2.6 Environmental, management, and temporal covariates……………..49 2.7 Data analysis……………………………………………………...…50 3 RESULTS………………………………………………………………...……….52 iv 3.1 Active plots……………………………..…………………….……....52 3.2 Passive plots…………………………………………….………........54 3.3 Pitfall traps……………………………………….……………….....55 4 DISCUSSION……………………………………………………………………...55 LITERATURE CITED………………………………………………………….…….62 CHAPTER 4: BAT ABUNDANCE IN RELATIONSHIP TO HABITAT FACTORS AT MULTIPLE SCALES ACROSS SAVANNAS, WOODLANDS, AND FORESTS IN THE MISSOURI OZARKS…………………………………………………...……92 ABSTRACT………………………………………………………………………....92 1 INTRODUCTION…………………………………………………………………..93 2 METHODS………………………………………………………………………..98 2.1 Study areas..………………………………………………….………98 2.2 Experimental design……………………………………………...…..99 2.3 Mist net survey and bat capture protocol……………………..……100 2.4 Site scale measurements and covariates……………………………101 2.5 Patch scale covariates…………………………………………...…104 2.6 Landscape scale covariates……………………………………..….104 2.7 Data analysis…………………………………………….….............105 v 3 RESULTS……………………………………………………………………..…106 3.1 Northern long-eared bats……………..………………………………...107 3.2 Tri-colored bats……………………………………….……..……..…...108 3.3 Evening bats…………………………………………….………...….....109 3.4 Eastern red bats…………………………………….………………......110 4 DISCUSSION……………………………………………………………….........111 5 MANAGEMENT IMPLICATIONS…………………………………………….........117 LITERATURE CITED………………………………………………………….…...119 CHAPTER 5: CONCLUSION…………………………………………………..……152 VITA………………………………………………………………………………...…156 vi List of Illustrations LIST OF TABLES Chapter 2 Tables Page 1. Model scenarios and parameters used to generate simulated datasets to evaluate the performance of n-mixture and removal models fit in the UNMARKED package. We varied the number of sites, number of visits to a site, the known population size (Ń) and the probability of detection (p) to create four scenarios………………………………………………………………..……...…27 2. Mean estimated abundance (N), standard error (SE), relative bias (RB), mean absolute error (MAE) and mean absolute percent error (MA%E) in abundance estimates by the n-mixture and removal model from simulated data for different numbers of sites and Ń=70, p=0.5, and 3 visits to a site………………...………29 3. Mean estimated abundance (N), standard error (SE), relative bias (RB), mean absolute error (MAE) and mean absolute percent error (MA%E) in abundance estimates by the n-mixture and removal model from simulated data for different numbers of visits and Ń=70, p=0.5, and 80 sites………………………………...30 4. Mean estimated abundance (N), standard error (SE), relative bias (RB), mean absolute error (MAE) and mean absolute percent error (MA%E) in abundance estimates by the n-mixture and removal model from simulated data for different known population sizes and detection probabilities and 3 visits to a site…...…...31 5. Mean estimated abundance (N), standard error (SE), relative bias (RB), mean absolute error (MAE) and mean absolute percent error (MA%E) in abundance estimates by the n-mixture and removal model from simulated when detection probability changed from 0.5 to 0.1 after individuals first capture (p, p ) and 1 known population =70, number of sites=80, and 3 visits to a site……………….33 Chapter 3 Tables 1. Hypotheses and candidate models used for effects on insect response variables based on passive and active plot surveys across a gradient of savannas, woodlands, and non-managed forests in the Ozarks of Missouri, 2014-2016. Site vii was used as a random effect and year (YR) and Julian date (JUL) were included as fixed effects in all models.…………………………………………………….66 2. Descriptions of model covariates used in generalized linear mixed models with a negative binomial distribution to predict abundances of insects across a gradient of actively managed savanna-woodlands and non-managed forests in the Ozark Highlands of Missouri, 2014-2016. ………………………………………..…....77 3. Response variables used in generalized linear mixed models with a negative binomial distribution examining the effects of restoration management, habitat, and climate covariates on insect abundance across a gradient of savanna, woodland, and forests in the Ozarks of Missouri, 2014-2016. Response variables represent the mean captures at plots over the 3 survey days. We combined malaise and panel trap yields for each plot for analysis. ……………………..…………..70 4. Descriptive statistics for covariates measured at plots where active methods were used to sample insects in a study of relationships between insect abundances and savanna woodland restoration in the Ozarks of Missouri, 2014-2016. ………….71 5. Descriptive statistics for covariates measured at plots where passive methods were used to sample insects in a study of relationships between insect abundances and savanna woodland restoration in the Ozarks of Missouri, 2014-2016..………….72 6. Support for generalized linear mixed models of different insect responses including the number of model parameters (k), log likelihood (LogLik), Akaike’s Information Criterion for small sample size (AICc), delta AICc (∆AICc), and AICc weight (w). Models were fit to data from 179 active plots collected across i a gradient of savanna, woodland, and forest in the Missouri Ozarks in summers, 2014-2016. ………………………………………………………………………73 7. Support for post hoc and a priori candidate generalized linear mixed models for mean Tricopteran response variable including the number of model parameters (k), log likelihood (LogLik), Akaike’s Information Criterion for small sample size (AICc), delta AICc (∆AICc), and AICc weight (w). Models were fit to data from i 179 active plots collected across a gradient of savanna, woodland, and forest in the Missouri Ozarks in summers, 2014-2016……..……………………………..76 viii

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Presented by Kathryn Marie Womack a candidate for the 8. CHAPTER 2: Performance of hierarchical abundance models on simulated bat capture data. Kathryn M. Womack. 1,3. , Sybill K. Amelon. 2. , Frank R. Thompson III. 2 2013, Womack 2008). Nocturnal insect activity is negatively related to.
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