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University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St. John's Road, Tyler's Green High Wycombe, Bucks, England HP10 8HR 77-16,963 LACKEY s James Andrew, 1948- A SYNOPSIS OF PHASEOLEAE (LEGUMINOSAE, PAPILIONOIDEAE). Iowa State University, Ph.D., 1977 Botany Xerox University Microfilms, Ann Arbor, Michigan 48io6 0 1977 JAMES A^roRBV LACKEY ALL RIGHTS RESERVED A synopsis of Phaseoleae (Leguminosae, Papilionoideae) by James Andrew Lackey A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of The Requirements for the Degree of DOCTOR OF PHILOSOPHY Department; Botany and Plant Pathology Major: Botany (Taxonomy) Approved: Signature was redacted for privacy. In Charge of Major Wor^ Signature was redacted for privacy. For the Major Department Signature was redacted for privacy. For the Gradifate College Iowa State University Ames, Iowa 1977 Copyright © James Andrew Lackey, 1977. All rights reserved. ii TABLE OF CONTENTS Page I. INTRODUCTION 1 II. HERBARIUM STUDIES 2 A. Introduction 2 1. Relationships of the Phaseoleae to other tribes 2 2. A taxonomic history of the Phaseoleae 7 a. de Candolle 7 b. Bentham 1837 9 c. Bentham I865 9 d. Taubert and Harms 12 e. Piper lU f. Recent workers 15 3. Terminology 15 B. Materials and Methods I8 C. A Revised Classification of the Phaseoleae 19 1. Introduction to the classification I9 2. The subtribes 26 a. Cajaninae 26 b. Diocleinae 32 c. Kennedi inae Uo d. Phaseolinae 4l e. Glycininae 58 f. Ophrestiinae 91 g. Erythrininae 95 h. Genera excluded from the Phaseoleae 99 D. Sketches of Selected Floral Dissections 101 III. CHROMOSOME STUDIES I67 A. Introduction 167 B. Materials and Methods 167 C. Results 168 D. Discussion 20i+ 1. General considerations 204 2. Systematic review 206 ill R. Conclusions 212 IV. LEAPLET ANATOMY STUDIES 213 A. Introduction 213 B. Materials and Methods 2lk C. Results—by Anatomical Characters 215 D. Results—by Taxonomic Groupings 230 E. Conclusions 236 V. SEED PROTEIN STUDIES 2k2 A. Introduction, 2k2 B. Materials and Methods 2^2 C. Results and Discussion 21+5 D. Conclusions 258 VI. EPILOGUE 259 A. Glycine and Associated Genera 259 1. Historical review 259 2. Results from this dissertation 262 3. Conclusions 266 B. Phaseoleae 267 VII. LITERATURE CITED 277 VIII. ACKNOWLEDGMENTS 291 1 I. INTRODUCTION This dissertation had more modest beginnings than a synopsis of Phaseoleae. It was originally planned as a study of the genus Glycine and allies, but it soon became obvious that this required a general review of the entire tribe Phaseoleae. To this end, this dissertation includes a survey of the tribe by traditional herbarium taxonomy methods. The conclusions are compared with taxonomic evidence derived from chromosome numbers, leaflet anatomy, seed protein identification on polyacrylamide gels, and canavanine distribution. The epilogue contains a synthesis for the Phaseoleae with special emphasis on Glycine and associated genera. 2 II. HERBARIUM STUDIES A. Introduction 1. Relationships of the Phaseoleae to other tribes The Phaseoleae belong to the family Leguminosae, subfamily Papilionoideae (Fabaceae, Faboideae; Leguminosae, Lotoideae (nom. illeg.); Papilionaceae). Bentham (1865a) divided the Papilionoideae into 11 tribes: Swartzieae, Sophoreae, Galegeae, Dalbergieae, Phaseoleae, Genisteae, Podalyrieae, Trifolieae, Loteae, Vicieae, and Hedysareae. The following are Bentham's conspecta as translated from the Latin by Hutchinson^ (1964): I. Podalyrieae. Shrubs, rarely herbs. Leaves simple or digitately compound. Stamens 10, free. II. Genisteae. Shrubs or herbs. Leaves simple or digitately compound; leaflets entire; racemes terminal or leaf- opposed, or flowers solitary or subfasciculate in the leaf-axils. Stamens 10, monadelphous, rarely diadelphous. III. Trifolieae. Herbs, very rarely fruticose. Leaves pinnately, rarely digitately, 3-foliolate, the nerves of the leaflets often running out into teeth; flowers solitary or racemose, peduncles axillary, rarely crowded in a terminal raceme. Stamens 10, diadelphous or monadelphous. IV. Loteae. Herbs or shrublets. Leaves pinnately 3-more- foliolate, leaflets entire. Flowers capitate or umbellate, rarely solitary, peduncles axillary or crowded at the ends of the branches. Stamens 10, diadelphous or monadelphous; filaments alternately dilated at the apex. 'All subsequent translations in this dissertation are my own. 3 V. Galegeae. Herbs, not climbing, erect shrubs, rarely trees or tall climbing shrubs. Leaves pinnately 5-more rarely 3-1-foliolate, leaflets often entire; petiole not tendril- iform. Flowers solitary, racemose, or paniculate. Stamens 10, diadelphous, or if monadelphous vexillary stamen often free at the base. Legume 2-valved, or if rarely indehiscent small 1-2-seeded or membranous-inflated. VI. Hedysareae. Habit of Loteae, Galegeae, or Phaseoleae. Legume jointed. VII. Vicieae. Herbs, leaves abruptly pinnate, petiole (rhachis) ending in a bristle or tendril; leaflets often denticulate at the apex. Stamens and legume as in Phaseoleae. VIII. Phaseoleae. Climbing herbs or rarely erect or shrubby, very rarely trees. Leaves pinnately, very rarely subdigitately, 3-foliolate, rarely 1-5-7-foliolate, leaflets entire or lobed, mostly stipellate; flowers racemose or fasciculate; peduncles often axillary. Stamens diadelphous or submon- adelphous. Legume 2-valved. IX. Dalbergieae. Trees or tall shrubs or high climbers. Leaves pinnately 5-more-foliolate, very rarely 3-1-foliolate. Inflorescence various, often paniculate or fasciculate. Stamens monadelphous or diadelphous. Legume exserted, indehiscent, membranous, coriaceous, woody or drupaceous. X. Sophoreae. Trees or tall shrubs or tall climbers, very rarely small shrubs or subherbaceous. Leaves pinnately 5- or more- foliolate, or large and 1-foliolate, rarely 3-foliolate. Stamens 10, free. XI. Swartzieae. Trees or tall shrubs. Leaves pinnately 5-more-, rarely 3-1-, foliolate. Calyx always (rarely in Sophoreae) entire before flowering, closed. Stamens numerous, rarely sub-10, free. Since Bentham, workers have attempted to organize papilionoid tribes into larger groups and define evolutionary lines or trends in the subfamily. Dormer (19^6) proposed a dichotomy in papilionoid evolution, one branch composed of "pulvinate types", and the other u of "epulvinate types", corresponding to the presence or absence of a foliar pulvinus. He listed shared characters for each group: Features common in the pulvinate types, rare or unknown in epulvinate 1. Stipels 2. Ridge bundles in the petioles and rachises (Watari 193^) 3. Anomalous secondary thickening 4. Secretory reservoirs 5. Longitudinal walls in the epidermal hairs, making them • multiseriate 6. Subsidiaty cells to the stomata of the leaves 7. Multilacunar nodes (Sinnott 19l4; Watari 193%) 8. Opposite position of the first two leaves of the seedling Features common in epulvinate types, extremely rare in pulvinate 1. Closed vascular systems Alignment of the Bentham tribes by Dormer's criteria is summarized in Table 2-1. The tribal groups, with the exception of the manifestly unnatural Galegeae and Hedysareae, largely maintain their unity. Turner and Fearing (1959)» based on chromosome numbers, arrived at. groupings similar to Dormer's; the pulvinate types have chromosome numbers based on eleven or reductions therefrom, but the epulvinate types have a base number of eight, or reductions from it. I have observed that the pulvinate types tend to be woody with bracteoles below the calyx, whereas epulvinate types tend to be herbaceous and lack bracteoles. Many workers now recognize more than the 11 tribes of Bentham