AN ABSTRACT OF THE THESIS OF Barbara L. Wilson for the degree of Doctor of Philosophy in Botany and Plant Pathology, presented on 25 February 1999. Title: Fescue Taxonomy in the Pacific Coast States. Redacted for Privacy Abstract approved: Aaron Liston Accurate taxonomy is essential to preservation and management of native grasses. Four problems in fescue taxonomy were investigated: theidentity of the native fescue that dominates western Oregon and Washington prairies, the relationship of native Festuca roemeri to closely related grasses, the status of F. howellii and F. washingtonica, and fescue identification. The cespitose fine-leaved fescue that dominates prairies in western Oregon and Washington had previously been identified as F. rubra. Two fescues were found to dominate these habitats, one introduced F. rubra var. commutata, and the other native F. roemeri. The two differed in isozymes and subtle morphological traits. Isozymes, flow cytometry of DNA, and leaf anatomy clarified relationships among F. idahoensis, F. roemeri. and the northwest Californiafescue that shares characters with both of them. Isozyme band patterns divided them into two groups; F. roemeri and a mixture of populations of F. idahoensis and the northwest California fescue. All three fescues were tetraploid. All three fescues were distinguished using leaf anatomy. Festuca roemeri and the northwest California fescue shared leaves elliptic to obovate in cross section and usually having three sclerenchyma bands, at margins and midrib; F. idahoensis had leaves round to hexagonal in cross section, with five or more sclerenchyma bands, Festuca idahoensis and the northwest California fescue had numerous long adaxial leaf hairs; F. roemeri had few and short adaxial hairs. Leaf shape was phenotypically plastic in F. roemeri and the northwest California fescue, but constant in F. idahoensis. Leaf sclerenchyma development was greater in the field than greenhouse for all three fescues. The northwest California fescue was recognized asF. roemeri var. klamathensis B. L. Wilson fined. Morphometric analysis of F. howellii and F. washingtonicarevealed that the two are distinct taxa. Festuca washingtonicaappeared to be a rare species endemic to lightly grazed and ungrazed areas of eastern Washington.The F. howellii holotype was conspecific with F. elmeri, not F. viridula as previouslythought. Because F. howellii is an earlier synonym, F.elineri must now be called F. howellii. Identification keys were provided for identifying fertile orsterile fescues and ryegrasses of the Pacific coast states. Copyright by Barbara L. Wilson February 25, 1999 All Rights Reserved Fescue Taxonomy in the Pacific Coast States by Barbara L. Wilson A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented February 25, 1999 Commencement June 1999 Doctor of Philosophy thesis of Barbara L. Wilson presented on February 25, 1999 APPROVED: Redacted for Privacy Major Professor, representing Botany and Plant Pathology Redacted for Privacy Chair of Department of Botany and Plant Pathology Redacted for Privacy Dean of School I understand that my thesis will become part of the permanentcollection of Oregon State University libraries. My signature below authorizes release of mythesis to any reader upon request. Redacted for Privacy Barbara L. Wilson, Author CONTRIBUTION OF AUTHORS Kathryn Beck and Florence Capiow collected Festuca washingtonica and made observations on its growth form and habitat. Susan Aiken wrote the F. ivashingtonica description, which I edited slightly as new specimens became available. She also provided insights into fescue taxonomy. Juan N. Zavala, Beth Parsons, and Krishnawan Panashatham performed flow cytometry on the fescue leaves. ii TABLE OF CONTENTS Page 1. Introduction 1 The Genus Festuca and the Fescue Clade 4 The Fescue Species 6 The Case of Festuca roemeri 9 2. A "New" Native Fescue of Western Oregon Prairies 14 Abstract 15 Introduction 16 Methods 17 Results 17 Discussion 28 Conclusion 30 Acknowledgements 31 3. Isozyme Variation and Taxonomy of Festuca idahoensisand F. roemeri 32 Address 33 Abstract 33 Introduction 34 Methods 35 Results 42 Discussion 52 Acknowledgements 63 111 TABLE OF CONTENTS (Continued) Page 4.Chromosome Number of Festuca idahoensis and F. roemeri Determined by Laser Flow Cytometry 65 Introduction 66 Methods 66 Results and Discussion 68 Acknowlegements 68 5. Phenotypic Plasticity in Leaf Anatomy in Festuca idahoensis and F. roemeri (Poaceae) 71 Address 72 Abstract 72 Introduction 73 Methods 74 Results 79 Discussion 86 Acknowledgements 87 6. A New Variety of Festuca roemeri (Pavlick) E. B. Alexeev 89 Introduction 90 Methods 91 Results 91 Discussion 93 iv TABLE OF CONTENTS (Continued) Page 7. What are Festuca howellii and F. washingtonica 9 98 Addresses 99 Abstract 99 Introduction 100 Methods 102 Results 104 Discussion 116 Conclusion 127 Acknowledgements 127 8. Key to the Fescues (s.1.) and Ryegrasses of the Pacific Coast States 128 Introduction 129 Notes on nomenclatural instability 136 Convensions used in the keys 138 Key # 1Plants with inflorescences, collected at or after flowering 140 Key # 2Sterile plants and plants collected before flowering 147 Key # 3The F. rubra complex 154 Key # 4Cultivated fescues of the F. ovina complex 155 Fescue leaf cross sections 158 Distribution and habitat of fescues and ryegrasses in the Pacific States 166 Acknowledgements 186 V TABLE OF CONTENTS (Continued) Page 9. Summary 187 Recommendations for further work 188 Bibliography 190 Appendices 198 Appendix 1. Sources for Illustrated Leaf Cross Sections 199 Appendix 2. Specimens Examined for Chapter 7 702 vi LIST OF FIGURES Figure Page 1.1 Spike lets of type specimens relevant to the nomenclature of F. roemeri 12 2.1 Cross sections of fescue leaves. 23 2.2 Distribution of Festuca roemeri in Oregon. 27 3.1 Malate dehydrogenase band patterns in Festuca. 43 3.2 Phenogram based on isozymes in four fescue taxa. 48 3.3 Phenogram based on isozymes in four fescue taxa. 49 3.4 Diagrams of the common band patterns for dimeric enzymes PGI, TPI, and AAT in the Festuca idahoensis/F. roemeri complex 50 3.5 Abundance of isozyme band patterns with different frequencies 53 4.1 DNA content of cells of Festuca taxa 70 5.1 Measurements and counts taken from leaf cross sections 78 5.2 Leaf cross section of field collected specimens (top row) and greenhouse plants (bottom row) 88 6.1 Leaf cross section of field collected specimens (top row) and plants from a common greenhouse environment (bottom row) 95 6.2 Typical malate dehydrogenase band patterns in Festuca 95 6.3 Distribution of F. roemeri var. klamathensis (left) and F. roemeri var. roemeri (right) 97 7.1 Leaf cross sections of F. elmeri, F. howellii, and F. viridula 108 7.2 Distribution of Festuca washingtonica in the state of Washington 111 7.3 Spikelets of Festuca washingtonica 112 7.4 Leaf cross sections of Festuca washingtonica and F. rubra 113 vii LIST OF FIGURES (Continued) Figure Page 7.5 Principal Components Analysis of fescue morphology 120 7.6 Distribution of Festuca howellii (left) and F. viridula (right) 126 8.1 Leaf anatomy of fine-leaved fescues 139 8.2 Leaf cross sections of Festuca brachyphylla ssp. coloradensis 159 8.3 Leaf cross sections of Festuca californica 159 8.4 Leaf cross sections of Festuca campestris 159 8.5 Leaf cross sections of Festuca filiformis (A), F. glauca (B-C), F.huonii (D)160 8.6 Leaf cross sections of Festuca howellii 160 8.7 Leaf cross sections of Festuca idahoensis 161 8.8 Leaf cross sections of Festuca minutiflora (A) and F. occidentalis (B-D). 161 8.9 Leaf cross sections of Festuca roemeri var. klamathensis 162 8.10 Leaf cross sections of Festuca roemeri var. roemeri 162 8.11 Leaf cross sections of F. rubra 163 8.12 Leaf cross sections of F. saximontana ssp. purpusiana 164 8.13 Leaf cross sections of F. trachyphylla 164 8.14 Leaf cross sections of F. trachyphylla and F. valesiaca 164 8.15 Leaf cross sections of F. viridula 165 8.16 Leaf cross sections of F. washingtonica 165 8.17 counties of the Pacific coast states 167 8.18 Distribution of Festuca arundinacea 169 viii LIST OF FIGURES (Continued) Figure Page 8.19 Distribution of Festuca brackvphylla ssp.brachyphylla 169 8.20 Distribution of Festuca brachphvlla ssp. coloradensis 170 8.21 Distribution of Festuca californica ssp. californica 170 8.22 Distribution of Festuca californica ssp. parishii 171 8.23 Distribution of Festuca catnpestris 171 8.24 Distribution of Festuca liliformis 172 8.25 Distribution of Festuca howellii 172 8.26 Distribution of Festuca idahoensis 173 8.27 Distribution of Festuca minutiflora 173 8.28 Distribution of Festuca occidentalis 174 8.29 Distribution of Festuca pratensis 174 8.30 Distribution of Festuca roemeri var. klamathensis 175 8.31 Distribution of Festuca roemeri var. roemeri 175 8.32 Distribution of Festuca rubra 176 8.33 Distribution of Festuca rubra ssp. arenicola 176 8.34 Distribution of Festuca rubra ssp. commutate 177 8.35 Distribution of F. rubra ssp. pruinosa 177 8.36 Distribution of Festuca rubra ssp. vallicola 178 8.37 Distribution of Festuca saximotztana var. purpusiana 178 8.38 Distribution of Festuca subulata 179 ix LIST OF FIGURES (Continued) Figure Page 8.39 Distribution of Festuca subuliflora 179 8.40 Distribution of Festuca trachyphylla 180 8.41 Distribution of Festuca viridula 180 8.42 Distribution of Festuca washingtonica 181 8.43 Distribution of Leucopoa kingii 181 8.44 Distribution of Lolium multiflorum 182 8.45 Distribution of Lolium perenne 182 8.46 Distribution of Lolium rigidum 183 8.47 Distribution of Lolium temulentum 183 8.48 Distribution of Vulpia bromoides 184 8.49 Distribution of Vulpia microstachys 184 8.50 Distribution of Vulpia myuros 185 8.51 Distribution of Vulpia octoflora 185 x LIST OF TABLES Table Page 2.1 Festuca roemeri specimens examined during thisstudy 18 2.2 Festuca rubra var.