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A Revision of the Phacelia Crenulatae Group (Hydrophyllaceae) for North America

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A Revision of the Phacelia Crenulatae Group (Hydrophyllaceae) for North America Brigham Young University BYU ScholarsArchive Theses and Dissertations 1972-05-01 A revision of the Phacelia crenulatae group (hydrophyllaceae) for North America N. Duane Atwood Brigham Young University - Provo Follow this and additional works at: https://scholarsarchive.byu.edu/etd BYU ScholarsArchive Citation Atwood, N. Duane, "A revision of the Phacelia crenulatae group (hydrophyllaceae) for North America" (1972). Theses and Dissertations. 8015. https://scholarsarchive.byu.edu/etd/8015 This Dissertation is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. A REVISION OF THE PHACELIAa:tENULATAE CEOUP (HYDROPHYLLACEAE)FOR NORTHAMERICA A Dissertation Presented to the Department of Botany and Range Science Brigham Young University In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy by Nephi Duane Atwood May 1972 ACKNOWLEDGEMENTS The author is indebted to Dr. Stanley L. Welsh for his valuable suggestions and also to other members of the committee, Dr. Joseph R. Murdock and Dr. Joseph R. Murphy. Special thanks is due James v. Allen for the Scanning Elec- tron micrographs of the seeds. The author is also indebt- ed to his wife for her cooperation and patience and also to Kay Thorne and Carolyn Christensen for the illDstrations. Appreciation is also extended to the various herbaria for loaning type material and other pertinent specimens. Fi- nancial assistance was provided in part by Cost of Educa- tion Funds through an NDEAfellowship and partly by the Department of Botany and Range Science, Brigham Young Uni- versity. iii TABLE OF CONTENTS Page ACKNOWLEDGEMENTS••••••••••••••••••••••••••••• iii LIST OF ILLUSTRATIONS•••••••••••••••••••••••• V LIST OF MAPS•••••• • • • ••• • •••••• • • ••••• • •• • ••• ix IN'IRODUCTION••••••••••••••••••••••••••••••••• 1 Materials and Methods••••••••••••••••••• 1 History of the Crenulatae group ••••••••• 4 General Morphology •••••••••••••••••••••• 5 Phylogeny••••••••••••••••••••••••••••••• 11 Distribution and Ecology •••••••••••••••• 29 Cytology •• •••••••••••••••••••••••••••••• 34 TAXONOMICTREATMENT • •• • •• • ••••••• • ••••• • • • ••• 36 BIBLIOGRAPHY • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 278 APPENDIX I•• •••• • • • •• • •• •. • • •••• •. • ••• • •••••• 286 APPENDIXII ••••••• • •••••• • •••••••• • •• • ••••••• 289 LIST OF ILLUS'lRATIONS Page Figure 1. A phylogenetic arrangement of the complexes in the Crenulatae group.•·• ••••••••••••••••• 17 A phylogenetic arrangement of the species in the neomexicana complex••••••••••••••••• 20 A phylogenetic arrangement of the species in the welshii complex••••••••••••••••••••• 22 A phylogenetic arrangement of the species in the palmeri complex••••••••••••••••••••• 25 s. A phylogenetic arrangement of the speeies in the crenulata complex••••••••••••••••••• 27 A phylogenetic arrangement of the species in the congesta complex•••••••••••••••••••• 31 A phylogenetic arrangement of the species in the scariosa complex•••••••••••••••••••• 33 a. Phacelia alba Rydberg•••••••••••••••••••••• 46 Dorsal and ventral view of the seeds of P. Rydberg••••••••••••••••••••••••••••: •• 50 10. Phacelia ambira Jones var. minutiflora (Voss in Munz Atwood•••••••••••••••••••••• 59 11. Dorsal and ventral view of the seeds of P. ambira Jones var. minutiflora (Voss in - MunzAtwood••••••••••••••••••••••••••••••• 63 12. Phacelia anelsonii Macbride•••••••••••••••• 67 Dorsal and ventral view of the seeds of P. anelsonii Macbride••••••••••••••••••••••: ••. 71 14. Phacelia arizonica Gray•••••••••••••••••••• 75 15. Dorsal and ventral view of the seeds of P. arizonica Gray••••••••••••••••••••••••••: •• 79 V 16. Phacelia bakeri (Brand) Macbride•••••••••• 84 17. Dorsal and ventral view of the seeds off• bakeri (Brand) Macbride••••••••••••••••••• 88 18. Phacelia bombycina Wooton and Standley.... 93 19. Dorsal and ventral view of the seeds off.• bombycina Wooton and Standley••••••••••••• 97 20. Phacelia coerulea Greene••••••~••••••••••• 101 21. Dorsal and ventral view of the seeds off.• coerulea Greene••••••••••••••••••••••••••• 105 22. Phacelia congesta Hooker•••••••••••••••••• 108 23. Dorsal and ventral view of the seeds off• congesta Hooker••••••••••••••••••••••••••• 112 24. Phacelia constancei Atwood•••••••••••••••• 117 25. Phacelia corrugata A. Nelson•••••••••••••• 122 26. Dorsal and ventral view of the seeds off• corrugata A. Nelson••••••••••••••••••••••• 126 27. Dorsal and ventral view of the seeds of P. coulteri Greenman•••••••••••••••••••••••-=-• 132 28. Dorsal and ventral view of the seeds off.. crenulata Torr. ex s. Wats•••••••••••••••• 137 Phacelia denticulata Osterhout •••••••••••• 141 30. Dorsal and ventral view of the seeds off.• denticulata Osterhout••••••••••••••••••••• 145 31. Phacelia glandulosa Nutt. var. glandulosa 155 32. Dorsal and ventral view of the seeds of P. glandulosa Nutt. a. var. argillacea b. - glandulosa •••••••••••••••••••••••••••••••• 159 33. Phacelia howelliana Atwood•••••••••••••••• 162 34. Dorsal and ventral view of the seeds of P. howelliana Atwood•••••••••••••••••••••••:• 166 35. Phacelia integrifolia Torr•••••••••••••••• 171 36. Dorsal and ventral view of the seeds of P. integrifolia Torr•••••••••••••••••••••••:• 173 vi 37. Phacelia intermedia Wooton•••••••••••••••• 180 38. Dorsal and ventral view of the seeds off• intermedia Wooton••••••••••••••••••••••••• 184 39. Phacelia palmeri Torr. ex s. Wats ••••••••• 195 40. Dorsal and ventral view of the seeds of!!• palmeri Torr. ex s. Wats•••••••••••••••••• 199 41. Phacelia pedicellata Gray••••••••••••••••• 203 42. Dorsal and ventral view of the seeds,of f• pedicellata Gray•••••••••••••••••••••••••• 207 43. Phacelia popei Torrey and Gray•••••••••••• 211 44. Dorsal and ventral view of the seeds off• popei Torrey and Gray••••••••••••••••••••• 215 45. Phacelia rafaelensis Atwood••••••••••••••• 219 46. Dorsal and ventral view of the seeds of P. rafaelensis Atwood••••••••••••••••••••••-=-• 223 47. Phacelia robusta (Macbride) Johnston •••••• 227 48. Dorsal and ventral view of the seeds of P. robusta (Macbride) Johnston•••••••••••••7• 231 49. Dorsal and ventral view of the seeds off• rupestris Greene•••••••••••••••••••••••••• 237 so. Phacelia scariosa Brandegee••••••••••••••• 240 51. Dors~l and ventral view of the seeds off• scariosa Brandegee•••••••••••••••••••••••• 244 52. Phacelia serrata Voss••••••••••••••••••••• 248 53. Dorsal and ventral view of the seeds off• serrata Voss•••••••••••••••••••••••••••••• 252 54. Phacelia splendens Eastwood••••••••••••••• 256 55. Dorsal and ventral view of the seeds of P. splendens Eastwood••••••••••••••••••••••:• 260 56. Phacelia utahensis Voss••••••••••••••••••• 263 57. Phacelia vossii Atwood•••••••••••••••••••• 269 vii 58. Phacelia welshii Atwood••••••••••••••••••• 275 viii LIST OF MAPS Map Number Page 1. Distribution of Phacelia ~-........... 48 2. Distribution of Phacelia amabilis •••••••• 54 3. Distribution of Phacelia ambigua ••••••••• 61 4. Distribution of Phacelia anelsonii ••••••• 69 s. Distribution of Phacelia arizonica ••••••• 77 6. Distribution of Phacelia bakeri •••••••••• 85 1. Distribution of Phacelia bombycina ••••••• 95 8. Distribution of Phacelia coerulea •••••••• 103 9. Distribution of Phacelia conqesta •••••••• 110 10. Distribution of Phacelia constancei •••••• 119 11. Distribution of Phacelia corruqata ••••••• 124 12. Distribution of Phacelia coulteri •••••••• 130 13. Distribution of Phacelia crenulata ••••••• 135 14. Distribution of Phacelia denticulata ••••• 143 15. Distribution of Phacelia formosula ••••••• 149 16. Distribution of Phacelia glandulosa •••••• 157 17. Distribution of Phacelia howelliana •••••• 164 18. Distribution of Phacelia integrifolia •••• 177 19. Distribution of Phacelia intermedia •••••• 182 20. Distribution of Phacelia neomexicana ••••• 188 21. Distribution of Phacelia pallida ••••••••• 191 22. Distribution of Phacelia palmeri ••••••••• 197 23. Distribution of Phacelia pedicellata ••••• 205 24. Distribution of Phacelia popei •••••••••.• 213 ix 25. Distribution of Phacelia rafaelensis ••••• 221 26. Distribution of Phacelia robusta ••••••••• 229 27. Distribution of Phacelia rupestris ••••••• 235 28. Distribution of Phacelia scariosa •••••••• 242 29. Distribution of Phacelia serrata ••••••••• 250 30. Distribution of Phacelia splendens ••••••• 258 31. Distribution of Phacelia utahensis ••••••• 265 32. Distribution of Phacelia vossii •••••••••• 271 33. Distribution of Phacelia welshii ••••••••• 277 X INTRODUCTION The Crenulatae group of Phacelia belongs to the sub- genus Phacelia section Phacelia and may be distinguished from other members of the section by the four seeded capsule and excavated ventral surface of the seeds. Many of the species are viscid and ill-scented desert plants confined mostly to western North America and Mexico. The remaining species occur in South America, the midwest, and the west-central part of the United States. The lack of phenologic, edaphic, morphologic, and distributional data plus the description of additional species since the monograph by Voss (1937) have made it necessary for a thorough study of the group. Very little was Jmown about the species occuring in Texas, New Mexico, Arizona, Utah, and California or species in Mexico and South America. Previous revisions were based entirely on herbarium specimens and possibly because of this numerous
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