A CONTRIBUTION TO THK TAXONOMY AND PHYLOGENY OF OXYTROPIS SECTION ARCTOBIA (FABACEAE) IN NORTH AMERICA B\ Zachary J Ylevers RFCOY1MF.NDFD ' <r Falk Huttmann. Ph.D. 't - Naoki Takebavashi. Ph D Stetanie Ickert-Bond. Ph D Advisory Committee ( hair Christa Yfulder. Ph D Chair. Department of Bioloyv and Wildlife ✓ APPROVED Paul Laver, Ph I) Dean. College ofJ^iatural Science and Vlathematics Itawfence Duffy. Pit D Interim Dean of the Graduate School S ', Date A CONTRIBUTION TO THE TAXONOMY AND PHYLOGENY OF OXYTROPIS SECTION ARCTOBIA (FABACEAE) IN NORTH AMERICA A THESIS Presented to the Faculty of the University of Alaska Fairbanks in Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE By Zachary J. Meyers, B.S. - ' - - » ■ - RASMUSON LIBnr Fairbanks, Alaska UNIVERSITY OF AUSKA-iF^RBANidi August 2012 Abstract North American species of Oxytropis section Arctobia are reviewed through herbarium vouchers, scanning electron microscopy of seed coats, and phylogenetic analysis of sequence data to test the validity of the circumscription of section Arctobia as previously proposed. Morphologically, section Arctobia is characterized by few- flowered inflorescences, few leaflets per leaf, and a mainly cushion-like habit. The micro-morphological data found considerable infraspecific variation in seed coat patterns. Sequence data from the nuclear TRPT, CNGC5, GA3oxl genes and the plastid matK gene were used to infer phylogenetic relationships among members of section Arctobia. Analyses using maximum parsimony, maximum likelihood and Bayesian inference of these sequences resulted in a phylogeny that unambiguously supports the monophyly of Oxytropis. None of the infrageneric subdivisions of Oxytropis were supported by our genetic analyses. Section Arctobia was found nested within species of section Orobia, Glaeocephala, and Baicalia. However, TRPT and matK sequence data support Oxytropis arctobia as a distinct species, which has previously been included under Oxytropis nigrescens under the North American treatment. Here eight species are recognized in section Arctobia for North America and a lectotype of Oxytropis arctobia is selected based on herbarium specimens. iv TABLE OF CONTENTS Page Signature Page ................................................................................................................... i Title Page............................................................................................................................ii Abstract ............................................................................................................................... iii Table of Contents ...............................................................................................................iv List of Figures .................................................................................................................... vi List of Tables ..................................................................................................................... viii l ist of Appendices ............................................................................................................ viii Introduction ........................................................................................................................ 1 Literature Cited..................................................................................................... 4 Chapter 1 A survey of seed coat morphology in Oxytropis DC. sections Arctobia, Baicalia, Glaeocephala, Mesogeae, and Orobia (Fabaceae) from Alaska. Abstract ..................................................................................................................9 Introduction ........................................................................................................... 10 Material and Methods ..........................................................................................13 Results....................................................................................................................15 Discussion ............................................................................................................. 27 Acknowledgements ..............................................................................................29 Literature Cited.....................................................................................................30 Chapter 2 Taxonomy and phylogeny of Oxytropis section Arctobia (Fabaceae) in North America. Abstract ................................................................................................................. 34 Introduction 35 V Material and Methods .......................................................................................... 41 Results.................................................................................................................... 47 Discussion ..............................................................................................................60 Taxonomic Treatment .......................................................................................... 65 Key to the North American species of Oxytropis section Arctobia.............. 66 Literature Cited............................................................ ......................................... 135 Appendix ................................................................................................................145 Conclusion ..........................................................................................................................148 Literature Cited 151 vi LIST OF FIGURES Page Figure 1.1 Seed shapes of Oxytropis in Alaska using scanning electron microscopy. 16 Figure 1.2 Seed coat patterns of Oxytropis in Alaska and the Russian Far East using scanning electron microscopy ........................................................................................... 23 Figure 1.3 Seed anatomy of selected species of Oxytropis in Alaska using scanning electron microscopy .......................................................................................................... 25 Figure 1.4 Frequency of the three main seed coat patterns found in each of the five sections of Oxytropis in Alaska .............................................................26 Figure 2.1 A tentative scheme of phylogeny of Oxytropis section Arctobia modified from Yurtzev (1997) ........................................................................................................... 39 Figure 2.2 Bayesian inference 50% majority consensus tree for the representative North American species of Oxytropis sect. Arctobia using TRPT sequence data ....54 Figure 2.3 Bayesian inference 50% majority consensus tree for the representative North American species of Oxytropis sect. Arctobia using CNGC5 sequence data .55 Figure 2.4 Bayesian inference 50% majority consensus tree for the representative North American species of Oxytropis sect. Arctobia using LE sequence data...........57 Figure 2.5 Bayesian inference 50% majority consensus tree for the representative North American species of Oxytropis sect. Arctobia using matK sequence data ......58 Figure 2.6 Bayesian inference 50% majority consensus trees for members of Fabaceae, with an emphasis on North American species of Oxytropis section Arctobia from the combined Bayesian analysis of matK and TRPT ........................................................... 62 Figure 2.7 Morphological characteristics of Oxytropis arctobia............................... 73 Figure 2.8 Distribution map of Oxytropis arctobia in the Canadian Archipelago, the Yukon Territory and British Columbia .................................................................... 74 Figure 2.9 Morphological characteristics of Oxytropis bryophila.............................. 80 Figure 2.10 Distribution map of Oxytropis bryophila in Alaska and neigh­ boring Russian and Canadian territories ..........................................................................81 vii Figure 2.11 Morphological characteristics of Oxytropis gorodkovii ...... .............96 Figure 2.12 Distribution map of Oxytropis gorodkovii in Alaska as well as neigh­ boring Russian and Canadian territories ..........................................................................97 Figure 2.13 Morphological characteristics of Oxytropis huddelsonii......................102 Figure 2.14 Distribution map of Oxytropis huddelsonii in A laska .......................... 103 Figure 2.15 Morphological characteristics of of Oxytropis kokrinensis .................. 108 Figure 2.16 Distribution map of Oxytropis kokrinensis in Alaska ........................... 109 Figure 2.17 Morphological characteristics of of Oxytropis mertensiana .................113 Figure 2.18 Distribution map of Oxytropis mertensiana in A laska ......................... 114 Figure 2.19 Morphological characteristics of Oxytropis podocarpa ......................121 Figure 2.20 Distribution map of Oxytropis podocarpa in North America .............. 122 Figure 2.21 Morphological characteristics of Oxytropis scammaniana ...................129 Figure 2.22 Distribution map of Oxytropis scammaniana in Alaska as well as the Yukon Territory and British Columbia ...................... 130 LIST OF TABLES Page Table 1.1 Specimens examined along with voucher information and locality data ... 17 Table 1.2 Summary statistics of the multinomial logistic regression with A1C model selection of seed coat micromorphology ..............................................................................27