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Molecular Systematics & Evolution of the CTENOSAURA HEMILOPHA

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Loma Linda University TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works Loma Linda University Electronic Theses, Dissertations & Projects 9-1999 Molecular Systematics & Evolution of the CTENOSAURA HEMILOPHA Complex (SQUAMATA: IGUANIDAE) Michael Ray Cryder Follow this and additional works at: https://scholarsrepository.llu.edu/etd Part of the Biology Commons Recommended Citation Cryder, Michael Ray, "Molecular Systematics & Evolution of the CTENOSAURA HEMILOPHA Complex (SQUAMATA: IGUANIDAE)" (1999). Loma Linda University Electronic Theses, Dissertations & Projects. 613. https://scholarsrepository.llu.edu/etd/613 This Thesis is brought to you for free and open access by TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. It has been accepted for inclusion in Loma Linda University Electronic Theses, Dissertations & Projects by an authorized administrator of TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. For more information, please contact [email protected]. LOMA LINDAUNIVERSITY Graduate School MOLECULARSYSTEMATICS & EVOLUTION OF THECTENOSAURA HEMJLOPHA COMPLEX (SQUAMATA: IGUANIDAE) by Michael Ray Cryder A Thesis in PartialFulfillment of the Requirements forthe Degree Master of Science in Biology September 1999 0 1999 Michael Ray Cryder All Rights Reserved 11 Each person whose signature appears below certifies that this thesis in their opinion is adequate, in scope and quality, as a thesis for the degree Master of Science. ,Co-Chairperson Ronald L. Carter, Professor of Biology Arc 5 ,Co-Chairperson L. Lee Grismer, Professor of Biology and Herpetology - -/(71— William Hayes, Pr fessor of Biology 111 ACKNOWLEDGMENTS I would like to express my appreciation to the institution and individuals who helped me complete this study. I am grateful to the Department of Natural Sciences, Lorna Linda University, for scholarship, funding and assistantship. I wish to thank Ronald L. Carter and L. Lee thismer for their advice and the use of their facilities, and I wish to thank the other member of my guidance committee, William Hayes for his advice and comments. I would also like to thank the remaining members of the faculty and student body for constantly inspiring me and providing an educational environment full of thought and support. I am grateful to Bradford Hollingsworth for providing technical assistance, friendship, and advice. His assistance on sequencing protocol, conducting the computer analyses and his understanding of phylogenetics was invaluable to my project. Many thanks to the Center of Molecular Biology and Gene Therapy at Loma Linda University for generating the DNA sequences needed for this project. Larry Buckley helped me in acquiring tissues and DNA sequences and I sincerely appreciate his efforts, comments and advice concerning spiny-tailed iguanas. I would like to thank the many individuals who helped me complete the field work done for this project, including Lee Grismer, Brad Hollingsworth, Humberto Wong, Jesse Grismer, Gene Kim, Nelson Wong, Victor Velasquez and the numerous Mexican ranchers, fisherman and friends who provided me with food, shelter and knowledge of the surrounding areas. I would also like to give a special thanks to my wife, Elissa, whose tireless support and sacrifice allowed me to complete my graduate studies. To my kids, Zachary and Miranda: never give up and always strive for the best. iv TABLE OF CONTENTS LIST OF FIGURES vi LIST OF TABLES viii ABSTRACT 1 INTRODUCTION ..... 3 Phylogenetic Analysis... ... 13 Biogeographic Topics__ ......... 14 MA i ERIALS AND METHODS ..................................................... 15 Cytochrome b and Cytochrome thddase III ..•••••••••••••.••.•••••.•.• 15 Sample Availability.••••••.•••••••••••••••.•.••••••••••••••••••• ...... •.••••.•• 15 Technical Protocols.••••.....•••••.••••••••••.•••••••••••.•••.•.•••••..•.••••••• 17 Alignments_ ...... •••••••. ..... .••••••••.•••••••••••.•••••••••• ••••••••• 21 Tree Reconstruction... ............ ......... ......... ... ... ... ... ... ............ 22 Outgroup Selection... ... ......... 23 Confidence and Support... ...... •••••••••••••••••• 23 RESULTS • •• ••• • • • ••• ••• t•• ••• ••• ••• ••• ••• ••• •• • ••• • 25 Phylogenetic Signal... 25 Combined DNA Sequences... .. 25 Maximum Parsimony Analysis__ ............ 25 DISCUSSION 33 C. nolascensis + C. conspicuosa... .................................... 33 Insular Populations__ ......... ............... 36 C. conspicuosa of Isla Cholludo......................••••••••••••••••••••••• 38 C. nolascensis + C. macrolopha.....................•••••••••••••.•••••••••• 38 C. hemilopha + C. h. insulana........................ ....................... 39 (C. macrolopha I C. nolascensis )+(C. hemilopha I C. h. insulana)... 39 LITERATURE CITED 40 APPENDIX ....... 46 Appendix I: Results of the computer program 46 Appendix II: Habitat and Specimen Photos of the C. hemilopha complex . 49 LIST OF FIGURES Figure 1. Proposed relationships of Ctenosaura: a) de Queiroz (1.987a,b) strict b) Hollingsworth (1998) strict consensus tree from a consensus tree; frequency bin analysis 8 Figure 2. Relationships based on congruence between de Queiroz (1987a,b) and Hollingsworth (1998) and the addition of three more recently described species Figure 3. Distribution of the Ctenosaura hemilopha complex ... ..... 11 Figure 4. Mammalian mitochondrial DNA gene map ... ...... 16 Figure 5. Strict consensus tree of the unweighted parsimony analysis using 22 ingroup samples and 2 outgroup taxa ... 27 Figure 6. Map showing the relative distance between; Islas San Esteban and San Pedro Nolasco in the Gulf of California_ . 28 Figure 7. Map of relative geographic distances between all populations within the Ctenosaura hemilopha complex._ ..... 31 Figure 8. Phylogenetic tree from witweighted analysis depicting the placement of the anomalous associations within Ctenosaura nolascensis 34 Figure 9. Map showing the geographic intermediacy of Isla Tiburon to Isla San Esteban and Mainland Sonora 37 Figure 10. Ctenosaura nolascensis basking in the morning sun on. tor) of a cactus on Isla San Pedro Nolasco, Sonora, Mexico._ .... 49 Figure 11. Rocky habitat of Ctenosaura nolascensis on Isla San Pedro Nolasco, Sonora, Mexico (top); Large male C. nolascensis basking on a granite boulder at the top of Isla San Pedro Nolasco, Sonora, Mexico (bottom)... ... 50 Figure 12. Large male Ctenosaura nolascensis sitting above nesting site on Isla San Pedro Nolasco, Sonora, Mexico (top); Green juvenile from Isla. San Pedro Nolasco, Sonora, Mexico (bottom)... ..... 51 Figure 13. Female Ctenosaura nolascensis basking on granite outcrop on Isla San Pedro Nolasco, Sonora, Mexico (top); Male C. nolascensis foraging in plants for food on Isla San Pedro Nolasco, Sonora, Mexico (bottom)... ... 52 Figure 14. Isla San Esteban, Sonora, Mexico, as seen from a southern shore of Isla Tiburon (top); Arroyo Limantur on Isla San Esteban, Sonora, Mexico 53 Figure 15. Large male Ctenosaura conspicuosa basking in the sun on top of a giant cactus in Arroyo Limantur on Isla San Esteban, Sonora, Mexico (top); Small male C. conspicuosa seeks refuge amongst the rocks and dense foliage on Isla San Esteban, Sonora, Mexico (bottom)... 54 Figure 16. Ctenosaura conspicuosa male seeking shelter in a rock crevice of Arroyo Limantur on Isla San Esteban, Sonora, Mexico (top); Juvenile C. conspicuosa from Isla San Esteban, Sonora, Mexico (bottom)... 55 Figure 17. Ctenosaura conspicuosa basking in the sun on a cactus within Arroyo Limantur on Isla Esteban, Sonora, Mexico... 56 Figure 18. Isla Cholludo, Sonora, Mexico (top), Northern shore of Isla Cholludo, Sonora, Mexico (bottom)... 57 vi Figure 19. Dense forest of cacti covering Isla Choiludo, Sonora, Mexico (top); Ctenosaura conspicuosa perched on a rock on Isla Cho'ludo, Sonora, Mexico ..•••.• 58 Figure 20. Arroyo above Rancho Santuaria in El Chorro, Baja California Stir, Mexico (top); Rocky mountainside above El Chorro, Baja California Sur, Mexico that contains crevices that serve as habitat for Ctenosaura hemilopha (bottom)... 59 Figure 21. Ctenosaura hemilopha habitat found on rocky mountainsides, south of San Bartolo, Baja California Sur, Mexico (top); Male C. hemilopha basking in the sun on a tree branch just south of San Bartolo, Baja California Sur, Mexico (bottom)... ..... 60 Figure 22. Green (unicolor) juvenile Ctenosaura hemilopha from Rancho Santuario in El Chorro, Baja California Sur, Mexico... 61 Figure 23. Northwest facing arroyo on Isla Cerralvo, Baja California, Mexico that provides habitat for Ctenosaura hemilopha (insulana) (top); Southwest shore- line habitat of C. hemilopha (insulana) on Isla Cerralvo, Baja California, Mexico (bottom)... ... ......... ...... 62 Figure 24. Arroyo on the northwest facing side of Isla Cerralvo, Baja California, Mexico, that provides habitat for Ctenosaura hemilopha (insulana) (top); Large male C. hemilopha (insulana) basking in the sun on Isla Cerralvo, Baja California, Mexico (bottom)... 63 Figure 25. Ctenosaura conspicuosa asleep on top of a cactus in Arroyo Limantur at sundown on Isla San Esteban, Sonora, Mexico... 64 vii LIST OF TABLES Table 1. Iguanid genera . ...... ............ ... ............ ... ... ... ... ..., • . •• .. • • . ...... 4 Table 2. Species and subspecies of the genus Ctenosaura......... ... ... ......... ..... 5 Table 3. Specimen localities with their corresponding codes and subspecies/ species designations ...... ...........
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  • An Overlooked Pink Species of Land Iguana in the Galápagos Gabriele Gentilea,1, Anna Fabiania, Cruz Marquezb, Howard L

    An Overlooked Pink Species of Land Iguana in the Galápagos Gabriele Gentilea,1, Anna Fabiania, Cruz Marquezb, Howard L

    An overlooked pink species of land iguana in the Galápagos Gabriele Gentilea,1, Anna Fabiania, Cruz Marquezb, Howard L. Snellc, Heidi M. Snellc, Washington Tapiad, and Valerio Sbordonia aDipartimento di Biologia, Universita` Tor Vergata, 00133 Rome, Italy; bCharles Darwin Foundation, Puerto Ayora, Gala´ pagos Islands, Ecuador; cDepartment of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131; and dGalápagos National Park Service, Puerto Ayora, Gala´ pagos Islands, Ecuador Edited by Francisco J. Ayala, University of California, Irvine, CA, and approved November 11, 2008 (received for review July 2, 2008) Despite the attention given to them, the Galápagos have not yet finished offering evolutionary novelties. When Darwin visited the Galápagos, he observed both marine (Amblyrhynchus) and land (Conolophus) iguanas but did not encounter a rare pink black- striped land iguana (herein referred to as ‘‘rosada,’’ meaning ‘‘pink’’ in Spanish), which, surprisingly, remained unseen until 1986. Here, we show that substantial genetic isolation exists between the rosada and syntopic yellow forms and that the rosada is basal to extant taxonomically recognized Galápagos land igua- nas. The rosada, whose present distribution is a conundrum, is a relict lineage whose origin dates back to a period when at least some of the present-day islands had not yet formed. So far, this species is the only evidence of ancient diversification along the Galápagos land iguana lineage and documents one of the oldest events of divergence ever recorded in the Galápagos. Conservation efforts are needed to prevent this form, identified by us as a good species, from extinction. Fig. 1. Galápagos Islands.