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Ecology and Evolution of Southeastern United States Yucca Species

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Ecology and Evolution of Southeastern United States Yucca Species ECOLOGY AND EVOLUTION OF SOUTHEASTERN UNITED STATES YUCCA SPECIES by JEREMY DANIEL RENTSCH (Under the Direction of JIM LEEBENS-MACK) ABSTRACT The genus Yucca contains approximately 40 species with most diversity found in Mexico and the southwestern United States. The southeastern United States is home to three well- described yucca species: the fleshy-fruited Y. aloifolia, the capsular-fruited Y. filamentosa, and Y. gloriosa – with a fruit type that does not follow convention. Yucca species are perhaps best known for the obligate pollination mutualism they share with moths in the genera Tegeticula and Parategeticula. Such interactions are thought to be highly specialized, restricting gene flow between species and even make evolutionary reversions to generalist life history characterizes impossible. Here, we show that Y. gloriosa is an intersectional, homploid, hybrid species produced by the crossing of Y. aloifolia and Y. filamentosa. We go on to show that Y. aloifolia has escaped from the obligate pollination mutualism and is being pollinated diurnally by the introduced European honey bee, Apis mellifera – an observation that directly refutes the idea that highly specialized species interactions lead to evolutionary dead ends. Finally, we utilized high throughput sequencing a biotinylated probe set in order to sequence many genes of interest in Y. aloifolia, laying the ground work to better understand its introduction history and pattern of pollinator association. INDEX WORDS: Yucca, hybrid speciation, population genetics, obligate mutualism ECOLOGY AND EVOLUTION OF SOUTHEASTERN UNITED STATES YUCCA SPECIES by JEREMY DANIEL RENTSCH BS, Kent State University, 2007 A Dissertation Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY ATHENS, GEORGIA 2013 © 2013 Jeremy D. Rentsch All Rights Reserved ECOLOGY AND EVOLUTION OF SOUTHEASTERN UNITED STATES YUCCA SPECIES by JEREMY DANIEL RENTSCH Major Professor: Jim Leebens-Mack Committee: Michael Arnold Shu-mei Chang Jim Hamrick Richard Lankau Wendy Zomlefer Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia May 2013 TABLE OF CONTENTS Page LIST OF TABLES ....................................................................................................................................................................... vi LIST OF FIGURES ................................................................................................................................................................... vii CHAPTER I INTRODUCTION AND LITERATURE REVIEW ........................................................................................................... 1 The Yucca Genus ........................................................................................................................................ 1 Obligate Pollination Mutualism........................................................................................................... 1 Interspecific Gene Flow and Hybridization .................................................................................... 2 Pollination Biology of Yucca species ................................................................................................. 4 Exceptions to the Obligate Mutualism.............................................................................................. 3 Population biology of Yucca species ................................................................................................. 5 References .................................................................................................................................................... 5 II HOMOPLOID HYBRID ORIGIN OF YUCCA GLORIOSA: INTERSECTIONAL HYBRID SPECIATION IN YUCCA (AGAVOIDEAE, ASPARAGACEAE) ............................................................................................... 8 Abstract ......................................................................................................................................................... 9 Introduction ............................................................................................................................................. 10 Materials and Methods ........................................................................................................................ 13 Results ........................................................................................................................................................ 17 Discussion ................................................................................................................................................. 18 References ................................................................................................................................................. 22 iv III YUCCA ALOIFOLIA (ASPARAGACEAE) OPTS OUT OF AN OBLIGATE POLLINATION MUTUALISM. .................................................................................................................................................... 30 Abstract ...................................................................................................................................................... 31 Introduction ............................................................................................................................................. 32 Materials and Methods ........................................................................................................................ 34 Results ........................................................................................................................................................ 38 Discussion ................................................................................................................................................. 40 References ................................................................................................................................................. 43 IV POPULATION GENETICS OF YUCCA ALOIFOLIA ................................................................................ 48 Abstract ...................................................................................................................................................... 49 Introduction ............................................................................................................................................. 50 Materials and Methods ........................................................................................................................ 52 Results ........................................................................................................................................................ 56 Discussion ................................................................................................................................................. 57 References ................................................................................................................................................. 59 V CONCLUSION AND DISCUSSION .............................................................................................................. 65 References ................................................................................................................................................. 67 v LIST OF TABLES Page Table 2.1: Microsatellite loci found to be variable between Y. aloifolia and Y. filamentosa. .................. 25 Table 3.1: Fruit composition statistics ......................................................................................................................... 46 Table 3.2: Diurnal pollination fruit composition statistics .................................................................................. 46 Table 4.1: Summary of sequencing and assembly statistics ................................................................................ 63 Table 4.2: Summary of SNP location and quality ..................................................................................................... 63 Table 4.3: Population genetics statistics ...................................................................................................................... 64 vi LIST OF FIGURES Page Figure 2.1: Location of field sites, species collected per site, and overall range of Y. aloifolia, Y. filamentosa, and Y. gloriosa in the United States. ..................................................................................... 26 Figure 2.2: Principal Coordinates Analysis (hybrid origin) ................................................................................. 27 Figure 2.3: STRUCTURE analysis .................................................................................................................................... 28 Figure 2.4: HINDEX analysis ............................................................................................................................................. 29 Figure 3.1: Experimental and control fruit sets ........................................................................................................ 46 Figure 3.2 Honeybee with fluorescent pollen analog ............................................................................................. 47 Figure 4.1: Yucca aloifolia collection sites ..................................................................................................................
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