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Spatial and Temporal Dynamics of Batesian Mimicry Between Adelpha University of the Pacific Scholarly Commons University of the Pacific Theses and Dissertations Graduate School 2018 Spatial and temporal dynamics of Batesian mimicry between Adelpha californica and Limenitis lorquini Louis Albert Prusa University of the Pacific, [email protected] Follow this and additional works at: https://scholarlycommons.pacific.edu/uop_etds Part of the Biology Commons Recommended Citation Prusa, Louis Albert. (2018). Spatial and temporal dynamics of Batesian mimicry between Adelpha californica and Limenitis lorquini. University of the Pacific, Thesis. https://scholarlycommons.pacific.edu/uop_etds/3133 This Thesis is brought to you for free and open access by the Graduate School at Scholarly Commons. It has been accepted for inclusion in University of the Pacific Theses and Dissertations by an authorized administrator of Scholarly Commons. For more information, please contact [email protected]. SPATIAL AND TEMPORAL DYNAMICS OF BATESIAN MIMICRY BETWEEN ADELPHA CALIFORNICA AND LIMENITIS LORQUINI by Louis A. Prusa A Thesis Submitted to the Office of Research and Graduate Studies In Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE Department of Biological Sciences College of the Pacific University of the Pacific Stockton, CA 2018 2 SPATIAL AND TEMPORAL DYNAMICS OF BATESIAN MIMICRY BETWEEN ADELPHA CALIFORNICA AND LIMENITIS LORQUINI by Louis A. Prusa APPROVED BY: Advisor: Ryan I. Hill, Ph.D, Committee Member: Tara Thiemann, Ph.D. Committee Member: Marcos Gridi-Papp, Ph.D. Department Chair: Craig Vierra, Ph.D. Dean of Graduate Studies: Thomas Naehr, Ph.D. 3 SPATIAL AND TEMPORAL DYNAMICS OF BATESIAN MIMICRY BETWEEN ADELPHA CALIFORNICA AND LIMENITIS LORQUINI Copyright 2018 by Louis A. Prusa 4 DEDICATION This thesis is dedication to my mentor Dr. Ryan I. Hill. He had faith in me and my abilities to conduct this long term project and to see it through to completion under his mentorship. Though it seemed daunting, Dr. Hill was there for support every step of the way. 5 ACKNOWLEDGMENTS My gratitude goes to Dr. Ryan Hill who gave me a chance and my lab mates Cassidi Rush and Erin Thompson and the rest of the Hill lab. I also want to thank my family, especially my parents, my brother and sister, my friends, and my thesis committee as well as the Biology Department of The University of the Pacific. This work was supported by National Science Foundation grant DEB-1342706 to R. I. Hill. Furthermore I want to thank Lindsay Wourms for her help with the field work in Sunol and Camp Ohlone. Thanks also go to Joe DiDonato, Gordon Wiley and the staff of East Bay Regional Parks and Sunol Regional Wilderness. 6 Spatial and temporal dynamics of Batesian mimicry between Adelpha californica and Limenitis lorquini Abstract by Louis A. Prusa University of the Pacific 2018 Conspicuous coloration is one of the main ways that animals communicate. The use of eye-catching color patterns to warn predators of an unprofitable trait is referred to as aposematism. Once predators learn to recognize the color pattern, a new signaling niche becomes available where other species can share the same signal. This mimicry niche can involve a “hide in plain sight” strategy by mimicking or parasitizing this signal, with mimics lacking the defense and associated costs that make them unprofitable. This is termed Batesian mimicry, and it decreases predation by taking advantage of the memory and learning of the predator community. Thus, a primary prediction in Batesian mimicry systems is that the model and mimic are found in sympatry. Another, fundamental prediction of Batesian mimicry is that the model outnumbers the mimic and that models emerge before the mimics to educate the predator guild. Some of these patterns were not significant in the California Coast Ranges as seen in Long et al. (2015), and no study has 7 estimated population sizes for this temperate Batesian mimicry system. Furthermore, compared with community studies of mutualistic Müllerian mimicry in the tropics, no studies have tested predictions of parasitic Batesian mimicry on small scale patterns of habitat use and movement patterns. If mimicry is as an important part of the biology of these temperate species, as it is for their tropical counterparts, we predict that in addition to emerging first and being more abundant, the model and mimic will overlap strongly in habitat but the model will be more abundant in each habitat, and will move more and be more widespread among available habitats. Our results confirm these predictions and indicate that A. californica is effectively educating habitat specialist and generalist predators providing an umbrella of protection for the mimic L. lorquini. 8 TABLE OF CONTENTS LIST OF TABLES .............................................................................................................. 9 LIST OF FIGURES .......................................................................................................... 10 CHAPTER 1: Introduction ..................................................................................................... 11 2: Materials and Methods ................................................................................... 20 Study Sites and Sampling ........................................................................... 20 Temporal, Relative Abundance and Population Size Analysis .................. 24 Habitat, Topography, Flight Height, and Movement Analyses .................. 25 3: Results .............................................................................................................. 28 Temporal Patterns, Relative Abundance and Population Size................... 28 Spatial Analyses ......................................................................................... 32 Habitat and Topography ................................................................ 32 Travel Distances (Single movement distances vs. Lifetime cumulative distances) .............................................................. 36 Flight Height .................................................................................. 38 4: Discussion ......................................................................................................... 40 5: Conclusion ........................................................................................................ 49 LITERATURE CITED ..................................................................................................... 50 APPENDIX A. .................................................................................................................. 60 9 LIST OF TABLES Table .............................................................................................................................. Page 1. Camp Ohlone MRR summary data. ........................................................................ 31 2. Jolly-Seber POPAN (JSP) models. ......................................................................... 31 3 Habitat use breakdown for the model and mimic species in Camp Ohlone. ........... 33 4. Movement probabilities for model and mimic for their best fit model................... 37 10 LIST OF FIGURES Figure ............................................................................................................................. Page 1. Study sites ................................................................................................................ 21 2. Weekly abundance of model and mimic over time ................................................. 29 3 Topographical use breakdown for model and mimic in Camp Ohlone .................... 33 4. Map of Sunol with marks/sightings for model A. californica (A) and mimic L. lorquini (B) ............................................................................................................... 34 5 Map of Camp Ohlone with marks/sightings for model A. californica (A) and mimic L. lorquini (B) ........................................................................................................... 35 6. Single movement distances for model and mimic. .................................................. 36 7. Flight height histograms for model A. californica (A) and mimic L. lorquini (B). 39 11 Chapter 1: Introduction Conspicuous coloration in animals is seen mainly in intra- and intersexual interactions, and in predator avoidance (Cott 1940, Edmunds 1974). Outside of sexual interactions, conspicuous color patterns were counter-intuitive to early naturalists that focused on the survival value of traits. For example, cryptic coloration (e.g. camouflage) was relatively intuitive because it reduces the probability of predator attack (Cott 1940). However, the benefit of using conspicuous coloration that seemingly advertises to predators was paradoxical (Joron and Mallet 1998). In time, naturalists accumulated evidence that these conspicuous color signals are used to warn predators of unprofitable defenses such as spines or chemical deterrents. E. B. Poulton clarified these syndromes and referred to the association between conspicuous coloration and unprofitability as aposematism (Poulton 1887). Aposematic signals are thought to evolve in prey species as a defensive mechanism that efficiently educates predators (Cott 1940, Huheey 1964, Huheey and Brandon 1974, Huheey 1976, Kikuchi 2010, Kikuchi 2013, Kikuchi and Sherratt 2015, Joron and Mallet 1998, Mappes, Marples and Endler 2005, Mallet and Joron 1999). The more memorable and detectable the signals are, the
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