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Scorpion Phylogeography in the North American Aridlands

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Scorpion Phylogeography in the North American Aridlands UNLV Theses, Dissertations, Professional Papers, and Capstones 8-1-2012 Scorpion Phylogeography in the North American Aridlands Matthew Ryan Graham University of Nevada, Las Vegas Follow this and additional works at: https://digitalscholarship.unlv.edu/thesesdissertations Part of the Biology Commons, Desert Ecology Commons, and the Population Biology Commons Repository Citation Graham, Matthew Ryan, "Scorpion Phylogeography in the North American Aridlands" (2012). UNLV Theses, Dissertations, Professional Papers, and Capstones. 1668. http://dx.doi.org/10.34917/4332649 This Dissertation is protected by copyright and/or related rights. It has been brought to you by Digital Scholarship@UNLV with permission from the rights-holder(s). You are free to use this Dissertation in any way that is permitted by the copyright and related rights legislation that applies to your use. For other uses you need to obtain permission from the rights-holder(s) directly, unless additional rights are indicated by a Creative Commons license in the record and/or on the work itself. This Dissertation has been accepted for inclusion in UNLV Theses, Dissertations, Professional Papers, and Capstones by an authorized administrator of Digital Scholarship@UNLV. For more information, please contact [email protected]. SCORPION PHYLOGEOGRAPHY IN THE NORTH AMERICAN ARIDLANDS by Matthew Ryan Graham Bachelor of Science Marshall University 2004 Master of Science Marshall University 2007 A dissertation submitted in partial fulfillment of the requirements for the Doctor of Philosophy in Biological Sciences School of Life Sciences College of Sciences The Graduate College University of Nevada, Las Vegas August 2012 Copyright by Matthew R. Graham, 2012 All Rights Reserved THE GRADUATE COLLEGE We recommend the thesis prepared under our supervision by Matthew R. Graham entitled Scorpion Phylogeography in the North American Aridlands be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biological Sciences School of Life Sciences Brett Riddle, Ph.D., Committee Co-Chair Jef Jaeger, Ph.D., Committee Co-Chair Allen Gibbs, Ph.D., Committee Member Matthew Lachniet, Ph.D., Graduate College Representative Tom Piechota, Ph.D., Interim Vice President for Research & Dean of the Graduate College August 2012 ii ABSTRACT Scorpion Phylogeography in the North American Aridlands by Matthew Ryan Graham Dr. Brett R. Riddle, Examination Committee Co-Chair Professor University of Nevada, Las Vegas Dr. Jef R. Jaeger, Examination Committee Co-Chair Research Assistant Professor University of Nevada, Las Vegas Understanding the geographic, geologic, and climatic forces responsible for generating current patterns of biodiversity has been a central objective of phylogeography. To develop a better understanding of these processes in the North American aridlands, I used DNA sequence data and species distribution modeling to conduct three phylogeographic assessments incorporating four species of arid-adapted scorpions: Hadrurus arizonensis, H. jedediah, H. spadix, and Paruroctonus becki. In an assessment of H. arizonensis, phylogeographic patterns indicate that Pleistocene climate cycles and associated glacial refugia played a central role in structuring the genetic diversity of this species in the Mojave and Sonoran deserts, mostly supporting predictions from a recent model of historical biotic assembly for these regions. However, the phylogeography of H. arizonensis also revealed a potential glacial refugium along the Lower Colorado River Valley that had not been considered in iii previous evaluations. To assess the impact that Pleistocene climate fluctuations had in other North American aridlands, I then compared phylogeographic patterns from H. jedediah and H. spadix, to those from H. arizonensis. Since these three species are closely related and morphologically similar, effects from phylogenetic signal and divergent phenotypes should have been minimal, so differences in phylogeographic patterns should reflect the relative influence of Pleistocene climates in different regions. Under this assumption, comparative phylogeography of these three species suggest that the impact of glacial climates was most pronounced for the biotas of the Great Basin and Snake River Plain, over those in the Colorado Plateau, and Mohave and Sonoran deserts. Finally, I conducted a phylogeography of P. becki, an unrelated species that spans the Mojave Desert and western Great Basin. Phylogenetic analyses identified five mitochondrial lineages in P. becki. The timing and geographic arrangement of these lineages supports a vicariant origin associated with the tectonically dynamic Eastern California Shear Zone. In association with these deeper patterns, demographic analyses indicated that a lineage in the Great Basin had undergone a recent post-glacial expansion, which according to predictions from climate-based models and a landscape interpolation of genetic distances, probably occurred from refugial areas in the northwest Great Basin. In general, phylogeographic assessments of North American aridland scorpions support phylogeographic inferences from co-distributed organisms, but add a novel glacial refugium in the Mojave Desert and a unique pattern of post- glacial expansions from an area within the Great Basin. iv ACKNOWLEDGEMENTS First, I thank my wife, Jessica Graham, for her continuous support of my sometimes crazy ambitions. Jessica and I have shared many grand adventures together and I look forward to many more; it is fun to be married to someone I also consider my best friend. I owe colossal thanks my parents, George and Penny Graham, who have been incredibly supportive, to the point that they accompanied me on numerous collecting trips in often harsh and downright miserable desert conditions. The following people deserve thanks for their willingness to stumble through the desert under the haze of purple lights in search of venomous arachnids: Joshua Ingoldt, Melinda Graham, Stephen Parks, Jef Jaeger, Tereza Jezkova, Michael Webber, Joshua Greenwood, Rebeca Rivera, Mallory Eckstut, Adam Leland, and Ryan Valverde. I thank Tereza Jezkova, my friend and unofficial mentor, for always being there to help in any way possible, even though it usually came with a good dose of sarcasm. I thank Michael Webber for her dedication to helping others and for being a great friend, despite claiming to hate systematics (we all know that she really loves it). I owe thanks to Robert Bryson Jr. for his encouragement and motivation, and for showing me how to “kick ass and take names”. Victor Fet, Rich Ayrey, Robert Bryson Jr., and Michael Wall kindly donated specimens. I thank Viktória Oláh-Hemmings for teaching me valuable skills in the lab, for help with analyses, and for showing the rest of us how to stay organized. Lisa Hancock kindly assisted with georeferencing. Victor Fet deserves many thanks for his guidance over the years and for his enthusiasm towards anything related v to scorpions; he is a constant reminder that science should be fun and rewarding. Michael Wall at SDNHM and Vince Lee, Anthea Carmichael, Stan Williams, and Charles Griswold at CAS provided assistance accessing museum collections. Lorenzo Prendini graciously provided important specimens and encouragement. I am grateful to everyone who has served on my advisory committee: Jef Jaeger, Brett Riddle, Matthew Lachniet, Allen Gibbs, Victor Fet, and Kathy Longshore. In particular, I thank Jef Jaeger and Brett Riddle for serving as my co-chairs and for their enormous contributions to my development as a scientist. Brett has been a constant role model and source of inspiration. His success is no doubt linked to his affable personality, enthusiasm for science, and his ability to completely change your scientific perceptions with a single sentence. I owe thanks to Jef for putting up with me over the years, for his good humor, support in and out of academia, and for showing me by example that you can achieve great things if you are willing to quit whining and do the work. Samples collected from Death Valley National Park were collected under permits issued by the National Park Service. A portion of this research was conducted under a task agreement with the National Park Service (J8R07080009) administered through the Great Basin Cooperative Ecosystem Studies Unit. vi TABLE OF CONTENTS ABSTRACT ............................................................................................................................ iii ACKNOWLEDGEMENTS ........................................................................................................ v LIST OF TABLES .................................................................................................................... ix LIST OF FIGURES ................................................................................................................... x CHAPTER 1 INTRODUCTION .......................................................................................... 1 CHAPTER 2 PHYLOGEOGRAPHY OF THE ARIZONA HAIRY SCORPION (HADRURUS ARIZONENSIS) SUPPORTS A MODEL OF BIOTIC ASSEMBLY IN THE MOJAVE DESERT AND ADDS A NEW PLEISTOCENE REFUGIUM ................................ 4 Abstract ........................................................................................................................... 4 Introduction ..................................................................................................................... 5 Materials and Methods ..................................................................................................
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