Finding Niebla Homalea: Predicting the Distribution of a Fog Lichen
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FINDING NIEBLA HOMALEA: PREDICTING THE DISTRIBUTION OF A FOG LICHEN A Thesis Presented to the faculty of the Department of Biological Sciences California State University, Sacramento Submitted in partial satisfaction of the requirements for the degree of MASTER OF SCIENCE in Biological Science (Ecology, Evolution, and Conservation) by Jamie Marie LeFevre SUMMER 2019 © 2019 Jamie Marie LeFevre ALL RIGHTS RESERVED ii FINDING NIEBLA HOMALEA: PREDICTING THE DISTRIBUTION OF A FOG LICHEN A Thesis by Jamie Marie LeFevre Approved by: ________________________, Committee Chair Ronald M. Coleman, Ph.D. ________________________, Second Reader Jamie M. Kneitel, Ph.D. ________________________, Third Reader Benjamin H. Becker, Ph.D. ________________________ Date iii Student: Jamie Marie LeFevre I certify that this student has met the requirements for format contained in the University format manual, and that this thesis is suitable for shelving in the Library and credits to be awarded for the thesis. ________________________, Graduate Coordinator _______________________ Jim Baxter Ph.D. Date Department of Biological Sciences iv Abstract of FINDING NIEBLA HOMALEA: PREDICTING THE DISTRIBUTION OF FOG LICHEN by Jamie LeFevre The fundamental niche is the set of conditions within which a species would be successful in the absence of biotic factors that might limit success. When a species or taxa is not well studied, as in the case with many lichens, understanding the environmental conditions that form its fundamental niche is a useful starting to point to predict species distribution. In recent years, ecological niche models have been used to successfully estimate fundamental niches and to predict species distribution. Ecological niche models are based on species presence and absence data and combinations of environmental variables. Once the environmental requirements for a species have been identified, predictive models can be used to extrapolate where the species would be expected to occur elsewhere in the landscape, allowing the fundamental niche to be mapped. The fundamental niche is reflected on a map based on the areas where abiotic conditions are right for a given species to occur. This lays the foundation to v explore larger questions such as the role of biotic interactions, predicting species responses to climate change, or developing conservation strategies. In this study, I have applied ecological niche modeling to Niebla homalea, a fog lichen endemic to the coastal zones of California and Baja California. The distribution and habitat requirements of N. homalea are largely unknown and the goal of this study is to expand our understanding of these attributes. Presence/absence data of N. homalea were collected at Bodega Bay, Point Reyes National Seashore, Half Moon Bay/San Francisco, and Monterey; in California, United States of America. Sampling sites were restricted to public lands with rock outcrops and stratified by average summer day-time fog density. Data on nine independent variables were also collected. Non-parametric multiplicative regression (NPMR) was used to find which independent variables are the best predictor variables for the presence of N. homalea. NPMR uses a local multiplicative smoothing function with leave-one-out cross-validation to estimate the response variable. The best predictor variables were found to be precipitation, habitat type, temperature, and fog density. The model was then used to extrapolate the probability that N. homalea exists in similar environments to those where the data were collected, based on independent variable characteristics. A distribution map representing the relative likelihood (high to vi low) of suitable habitat, based on the inter-relationships of the predictor variables was created using ArcGIS. The mapping allowed a comparison between the predicted habitat suitability map and the known distribution of N. homalea. A comparison of predicted habitat suitability and known locations of N. homalea found that N. homalea tended to be in high probability of occurrence areas on the habitat suitability map. ________________________, Committee Chair Ronald M. Coleman, Ph.D. ________________________ Date vii ACKNOWLEDGEMENTS I would like to thank Pam Kirkbride for introducing me to the wonderful world of lichens and Shelly Benson for introducing me to fog lichens, and sharing her knowledge and goals. I hope the information from this study can benefit your ongoing research on fog lichens. I would like to thank Susie Bennett from the National Park Service for letting me collect data at Rancho Corral de Tierra on Montara Mountain and Montara State Beach and for your enthusiasm for my research. I would also like to thank Bruce McCune for answering my questions when I got stuck using HyperNiche. I would like to thank my husband, Joe Griffin, and my daughter, Alyssa Egbert, for their support and going with me on the lichen hunts. Without their constant support and encouragement, I would not have been able to complete my research. Finally, I would like to thank my committee members Jamie Kneitel, and Benjamin Becker for their guidance during my research and manuscript preparation. I would like to express my sincere gratitude to my advisor, Ron Coleman, for allowing me freedom to explore my own ideas and at same time giving guidance to complete my research and “get it done”. viii TABLE OF CONTENTS Page Acknowledgements .......................................................................................................... viii List of Tables .. ................................................................................................................ xii List of Figures ................................................................................................................. xiii Chapter 1. INTRODUCTION ........................................................................................................ 1 Predictive Habitat Modeling ................................................................................ 3 Lichen Distribution .............................................................................................. 6 Fog Lichens .......................................................................................................... 8 Hypothesis.......................................................................................................... 10 2. MATERIALS AND METHODS ................................................................................ 12 Study Sites ......................................................................................................... 12 Data Collection .................................................................................................. 17 Response variables ............................................................................................. 17 Predictor variables ............................................................................................. 21 Analysis.............................................................................................................. 23 Model Strategy and Evaluation .......................................................................... 29 3. RESULTS…. .............................................................................................................. 33 Data Collection .................................................................................................. 33 NPMR Model ..................................................................................................... 33 Sensitivity of Individual Predictors ................................................................... 41 ix Predictive Sensitivity of Occurrence ................................................................. 45 4. DISCUSSION ............................................................................................................. 49 Predictive Sensitivity of Occurrence ................................................................. 52 Ecological Niches and Conservation ................................................................. 59 Model Implications ............................................................................................ 61 Model Limitations .............................................................................................. 63 Suggestions for Future Research ....................................................................... 65 5. CONCLUSION ........................................................................................................... 67 Appendix A. Field data collected at Bodega Bay: sampling site location, rock group number and location, species recorded at each rock group and percent cover of each lichen species on each rock group .................. 69 Appendix B. Representitive photographs of lichens observed at Bodega Bay .............. 86 Appendix C. Field data collected at Point Reyes National Seashore: sampling site location, rock group number and location, species recorded at each rock group and percent cover of each lichen species on each rock group ......................................................................................................... 91 Appendix D. Representitive photographs of lichens observed at Point Reyes National Seashore ................................................................................... 102 Appendix E. Field data collected at Half Moon Bay and San Francisco: sampling site location, rock group number and location, species recorded at each rock group and