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Taxonomy and Phylogeny of the Genus Lappula Moench (Boraginaceae) in North America

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Taxonomy and Phylogeny of the Genus Lappula Moench (Boraginaceae) in North America TAXONOMY AND PHYLOGENY OF THE GENUS LAPPULA MOENCH (BORAGINACEAE) IN NORTH AMERICA by SUSAN J. ROLFSMEIER B.A., University of Montana, 1997 M.S., University of Nebraska, Lincoln, 2000 AN ABSTRACT OF A DISSERTATION submitted in partial fulfillment of the requirements for the degree DOCTOR OF PHILOSOPHY Division of Biology College of Arts and Sciences KANSAS STATE UNIVERSITY Manhattan, Kansas 2013 Abstract The genus Lappula Moench is a diverse group of herbaceous plants in the large, cosmopolitan family Boraginaceae. Over sixty species are recognized in Eurasia; many are found in cold deserts, steppes, and semi-deserts of Central Asia. Fewer species were described from western North America, and compared with the Asian species they are poorly known. Various North American taxa have been placed into synonomy under Eurasian species, and complex patterns of variation have made species circumscription challenging. The goal of this dissertation was to explore phylogenetic relationships between North American and Eurasian species and to revise the taxonomy of the North American species. A molecular phylogenetic study was initiated in order to infer patterns of relationships among the North American species relative to Eurasian diversity. Samples were collected from throughout the western United States and from Siberia. Additional samples of Eurasian species were taken from herbarium specimens. Sequences were generated for three DNA regions (the ITS region of the nuclear ribosomal DNA, and chloroplast intergenic spacers trnS-trnG and rpl32-trnL) and phylogenies were generated using parsimony and Bayesian analyses. Results were in general agreement among all analyses. The genus Lappula was recovered as a monophyletic group, with the exception of the morphologically anomalous L. sessiliflora (Boiss.) Gürke, which was sister to the genus Rochelia Rchb. The native North American species of Lappula and L. redowskii (Hornem.) Greene formed a clade. Samples of L. squarrosa (Retz.) Dumort., a Eurasian steppe plant with a wide introduced range in North America, grouped with samples from Eurasia. Sampled species of Hackelia Opiz, a genus sometimes treated as part of Lappula, formed a clade separate from the Lappula species sampled here. Herbarium and field studies resulted in a revised taxonomy for the North American Lappula. Nomenclatural problems were resolved; nine native species and four varieties were recognized along with the introduced Eurasian species L. squarrosa. TAXONOMY AND PHYLOGENY OF THE GENUS LAPPULA MOENCH (BORAGINACEAE) IN NORTH AMERICA by SUSAN J. ROLFSMEIER B.A., University of Montana, 1997 M.S., University of Nebraska, Lincoln, 2000 A DISSERTATION submitted in partial fulfillment of the requirements for the degree DOCTOR OF PHILOSOPHY Division of Biology College of Arts and Sciences KANSAS STATE UNIVERSITY Manhattan, Kansas 2013 Approved by: Major Professor Carolyn J. Ferguson Abstract The genus Lappula Moench is a diverse group of herbaceous plants in the large, cosmopolitan family Boraginaceae. Over sixty species are recognized in Eurasia; many are found in cold deserts, steppes, and semi-deserts of Central Asia. Fewer species were described from western North America, and compared with the Asian species they are poorly known. Various North American taxa have been placed into synonomy under Eurasian species, and complex patterns of variation have made species circumscription challenging. The goal of this dissertation was to explore phylogenetic relationships between North American and Eurasian species and to revise the taxonomy of the North American species. A molecular phylogenetic study was initiated in order to infer patterns of relationships among the North American species relative to Eurasian diversity. Samples were collected from throughout the western United States and from Siberia. Additional samples of Eurasian species were taken from herbarium specimens. Sequences were generated for three DNA regions (the ITS region of the nuclear ribosomal DNA, and chloroplast intergenic spacers trnS-trnG and rpl32-trnL) and phylogenies were generated using parsimony and Bayesian analyses. Results were in general agreement among all analyses. The genus Lappula was recovered as a monophyletic group, with the exception of the morphologically anomalous L. sessiliflora (Boiss.) Gürke, which was sister to the genus Rochelia Rchb. The native North American species of Lappula and L. redowskii (Hornem.) Greene formed a clade. Samples of L. squarrosa (Retz.) Dumort., a Eurasian steppe plant with a wide introduced range in North America, grouped with samples from Eurasia. Sampled species of Hackelia Opiz, a genus sometimes treated as part of Lappula, formed a clade separate from the Lappula species sampled here. Herbarium and field studies resulted in a revised taxonomy for the North American Lappula. Nomenclatural problems were resolved; nine native species and four varieties were recognized along with the introduced Eurasian species L. squarrosa. Table of Contents List of Figures ............................................................................................................................... vii List of Tables ................................................................................................................................. ix Acknowledgements ......................................................................................................................... x Dedication .................................................................................................................................... xiii Chapter 1 - A molecular phylogenetic study of the genus Lappula Moench ................................. 1 Introduction ................................................................................................................................. 2 Materials and Methods ................................................................................................................ 9 Results ....................................................................................................................................... 23 Discussion ................................................................................................................................. 43 Conclusions ............................................................................................................................... 48 Literature Cited ......................................................................................................................... 49 Acknowledgements ................................................................................................................... 57 Chapter 2 - A taxonomic revision of Lappula (Moench) in North America ................................ 59 Introduction ............................................................................................................................... 60 Morphology .............................................................................................................................. 62 Cytology .................................................................................................................................... 67 Taxonomic background ............................................................................................................ 67 Taxonomic Treatment ............................................................................................................... 73 Genus description (as applicable to the North American species)........................................ 73 Key to Species ....................................................................................................................... 74 1. Lappula coronata Greene ................................................................................................. 80 2. Lappula cucullata A. Nelson ............................................................................................ 84 3. Lappula desertorum Greene ............................................................................................. 92 4. Lappula fremontii (Torr.) Greene ..................................................................................... 98 Key to varieties: ................................................................................................................ 98 4a. Lappula fremontii (Torr.) Greene var. fremontii. ........................................................ 98 4b. Lappula fremontii (Torr.) Greene var. nelsonii var. novem ..................................... 110 v 5. Lappula heterosperma Greene ........................................................................................ 113 6. Lappula longispina ......................................................................................................... 120 7. Lappula montana Greene ................................................................................................ 124 8. Lappula occidentalis (S. Watson) Greene ...................................................................... 129 Key to varieties ............................................................................................................... 129 8a. Lappula occidentalis (S. Watson) Greene var. occidentalis. .................................... 129 8b. Lappula occidentalis var. stricta (S. Watson) comb. nov......................................... 147 9. Lappula squarrosa (Retz.) Dumort. ...............................................................................
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