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A Study of Morphological Variation Within Pseudoroegneria Spicata

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A Study of Morphological Variation Within Pseudoroegneria Spicata AN ABSTRACT OF THE THESIS OF Jack Revnold Carison for the degree of Master of Science in Crop Science presented on March 2O 1986 TITLE: . !st St f p soicata (Pursh) A. L&ve (Poaceae: TriticeaeJ Abstract Approved: Signature redacted for privacy. Robert J. Metzger Chromosome counts were determined for 152 accessions of Pseudo- roegneria spicata (Pursh) A. L6ve and, combined with existing count data, used to plot the distribution of diploid and tetraploid popula- tions. Morphological variation of 55 characters was examined in five groups totaling 205 operational taxonomic units (OTU's), using cluster, principal factor, and discriminant analyses. The five groups included diploid and autotetraploid spicata, an allotetraploid pre- viously included in spicata, a control group including four Old World Pseudoroegneria species, and a small control sample of Elymus lanceolatus (Scrib. and Smith) Gould. The analyses were not able to separate diploid from autoploid spicata nor identify any clear sub- groupings within the dip].ojds. However, the alloploid was separated from spicata and aligned with Elymus lanceolatus based on glume and spike characters. This study recommends the alloploid be included in lanceolatus as a new subspecies, Elymus lanceolatus ssp. wawawai. The chromosome count data indicate it is distributed in the canyons and tributaries of the lower Salmon and Snake Rivers of northern Idaho, northeastern Oregon, and southeastern Washington. The new subspecies keys to Elymus lanceolatus based on glume characters and is separated from subspecies .ianceolatus and albicans by its cespitose growth habit. A S11LPY OF MORPHOLOGICAL VARIATION WITHIN PSELVOROEGNERIA SPICATA (PURSH) A. LOVE (POACEAE: TRITWEAE) By Jack Reynold Carison A THESIS Submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Completed March 20, 1986 Commencement June, 1986 APPROVED: 1' Signature redacted for privacy. Professor of Crop Science in charge of major Signature redacted for privacy. - Read of Department of Crop Science Signature redacted for privacy. Dean of Graduate hoo 1 Date thesis is presented Typed by Marguerita Balint-Carden for JackR.Carison ACKNOWLEDGEMENTS I would like to extend my appreciation to my Major Professor and good friend, Dr. Robert J. Metzger, for his excellent guidance throughout the course of my graduate study. Together with Dr. Douglas R. Dewey, their expertise in cytogenetics provided an outstanding foundation for my work on this thesis problem. I am especially indebted to Dr. Dewey for his willingness to serve on my graduate committee, despite the difficulty in doing so from his position at Logan, Utah. I feel very fortunate to have received training from hint. I thank Dr. Kenton Chambers, Dr. Warren Kronstad, and Dr. Harry Mack for accepting the membership of my graduate committee. A special thanks to Dr. Chambers for his guidance on biosystentatics and his out- standing teaching in this area. Appreciation also is extended to Dr. Mary Barkworth and Dr. Steven Broich for assistance with numerical taxonomy questions. I thank the USDA Soil Conservation Service for the opportunity and support to complete this graduate study. Special thanks to the late Archie Fuchs and my current supervisor, Clarence Maesner, for their patience and helpfulness. I want to express my gratitude to Marguerita Balint-Carden for her excellent assistance with word pro- cessing in the development of this manuscript. I want to acknowledge the friendship and collaboration of Dr. Luiz Gonzaga E. Vieira, my former fellow graduate student in plant gene- tics. Luiz is a wizard in the laboratory, likes to argue passionately about scientific issues (among others), and is very open and helpful to others. He created a day-to-day learning environment that was a tremendous benefit to me. I shall miss this interaction. Finally, I feel very fortunate in the love, encouragement, and support from my wife, Vicki, and children, Chris and Sara, while completing this degree. TABLE OF CONTENTS Pace INTRODUCTION 1 MATERIALS AND METHODS 5 RESULTS 10 DISCUSSION 21 CONCLUSIONS 24 BIBLIOGRAPHY 46 APPENDICES 49 LIST OF FIGURES Yiqure Pane 1 Distribution of Pseudoroegneria spicata in 27 western North America, including the alloploid form. 2 Condensed dendogram obtained from average linkage 28 cluster analysis of 205 OTU's of Pseudoroegneria spicata and Elymus. 3 Scatter plot of factor scores for spike/spikelet 29 and awn characters in principal factor analysis of 205 Pseudoroegneria spicata OTU's. 4 Scatter plot of factor scores for spike/spikelet 30 and awn characters in principal factor analysis of 165 Pseudoroegneria spicata OTU's. 5 Scatter plot of factor scores for spike/spikelet 31 characters and leaf/cuim lengths in principal factor analysis of 165 Pseudoroegneria spicata OTUS. 6 Scatter plot of factor scores for spike/spikelet 32 characters and leaf/cuim widths in principal factor analysis of 165 Pseudoroegneria spicata OTU's. 7 Scatter plot of factor scores for leaf/cuim widths 33 and awn characters in principal factor analysis of 165 Pseudoroegneria spicata OTU's. 8 Scatter plot of canonical variable scores for five 34 groups of Pseudoroegneria and Elymus. 9 Scatter plot of canonical variable scores for three 35 groups of Pseudoroegneria spicata, including the a].loploid form. LIST OF TABLES Table Paae 1 Species of the genus Pseudoroegneria (Nevski) 36 L6ve. 2 Characters used to score OTU's of five groups of 38 Pseudoroegneria and Elymus. 3 Morphological characters selected for use in 40 cluster analysis of Pseudoroegneria and Elymus OTU' s. 4 Group frequency in five major clusters formed by 41 analysis of 17 morphological characters of 205 OTU's of Pseudoroegneria and Elymus. 5 Variance explained by factors identified in 42 principal factor analysis of three combinations of morphological characteristics measured on 205 OTU's of Pseudoroegneria and Elymus. 6 Rotated estimated factor loadings in principle 43 factor analysis of 165 OTU's of Pseudoroegneria. 7 Variables selected by stepwise discriminant 44 analysis and classification functions for groups of Pseudoroegneria and Elymus OTU's. 8 Jackknifed classification of OTU's into five and 45 three groups of Pseudoroegneria and Elymus by discriminant analysis. LIST OF APPENDIX TABLES Table Pane 1 Collection information for 205 OTU's of 50 Pseudoroegneria and Elymus. 2 Environmental conditions at four uniform 56 garden nursery sites which were sources of 192 Pseudoroegneria and Elymus OTU's. 3 Source of known diploid, triploid, auto- 57 tetraploid, and allotetraploid forms of Pseudorogenria spicata. 4 Means and standard deviations of 55 morphological 65 characters for five groups of Pseudoroegneria and Elymus. A STUDY OF MORPHOLOGICAL VARIATIONWITHIN PSEUDOROEGNERIASPICATA (PURSHJ A. LOVE (POACEAE: TRITICEAE) INTRODUCTION The genus Pseudoroegneria (Nevski) L5ve, was recently con- structed (L6ve 1982) to accommodate all S genome species of the Triti- ceae tribe of grasses. Several members of this new genus came from section Pseudoroegneria in the genus Elytrigia, established by Nevski (1934), hence the new generic name. The new genus includes one North American species, still commonly known as Agropyron spicatum (Pursh) Scrib. and Smith, after the treatment by Hitchcock (1951). The name Pseudoroegneria spicata (Pursh) Lve, will be used for this species, which is the focus of this study, as proposed in the new nomenclature for North American Triticeae by Barkworth et al. (1983) and Barkworth and Dewey (1985). This new treatment of North American Triticeae is based on the genomic system of classification advocated by Dewey (1982, 1983) and L6ve (1982, 1984). That is, different genomes and combinations of genomes determine generic limits and pro- vide a clearer understanding of biological relationships, a major objective of taxonomy. Bluebunch wheatgrass, a widely used common name of P. spicata, is a major cool-season range grass, native to much of western North America. It is characterized by Hitchcock (1951) as cespitose with erect cuims, awned and distant spikelets, a continuous rachis, diver- gent awns, more than 7 spikelets per spike, spikes 8 to 15 cm long, and spikele'cs usually shorter than the internode. The awnless form, considered by Hitchcock (1951) as the separate species Agropyron inerme (Scrib. and Smith) Rydb., differs from P. spicata only in the awnless spikelets and is not considered to have much taxonomic significance (Daubenmire 1939, Dewey 1982). However, it is considered a subspecies in some treatments (LBve 1984). Bluebunch wheatgrass is a major component of many plant coirimuni- ties, ranging from arid desert sagebrush(Artemisia)habitats to mesic, subhumj.d pine/fir(Pinus/Pseudotsuga)woodlands.It is often associated or sympatric with other Triticeae such as thickspike wheatgrass,Elymus lanceolatus(Scrib. and Smith) Gould; squirrel- tail,E. elymoides(Raf.) Sweezy; slender wheatgrass,E.trachy- caulus(Link.) Gould ex Shinners; western wheatgrass,Pascopyrum smithii(Rydb.) L6ve; basin wildrye,Leymus cinereus(Scrib. and Merr.,) Love; and beardless wildrye,L. triticoides(Buckley) Pilger.Natural hybrids betweenP. spicataandE. lanceolatus, and betweenspicataandE. elymoides,are conunon and are evi- dence for introgression of specific characters.For instance, spi- catapopulations found in mesic woodland habitats often are mildly rhizomatous, a characteristic perhaps contributed bylanceolatus (Dewey 1982). E. lanceolatus spp. albicans(Scrib. and Smith)
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