Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1978 Section Chamaefistula, series Corymbosae (Leguminosae, Cassia) of the New World Alex Lasseigne Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons Recommended Citation Lasseigne, Alex, "Section Chamaefistula, series Corymbosae (Leguminosae, Cassia) of the New World " (1978). Retrospective Theses and Dissertations. 6463. https://lib.dr.iastate.edu/rtd/6463 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. 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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 7913236 LASSEIGNE» ALEX section ChAMAEFlSTULÂ, SERIES CCRy^BOgAE (LEGUHINGÛAE, CASSIA) OF THE NEw .OPLC. lO^A STATE university, °-i,0.r 19?c Universiw Microfilms Intematcnal 300 N ZEES «OAD.ANN ARBOR, MI «SIOS Section Chamaefistula. series Corymbosae (Leguminosae, Cassia) of the New World by Alex Lasseigne 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. Signature was redacted for privacy. For the Major Department Signature was redacted for privacy. Iowa State University Ames, Iowa 1978 11 TABLE OF CONTENTS Page I. INTRODUCTION 1 II. TAXONOMIC HISTORY 4 III. MORPHOLOGICAL AND ANATOMICAL STUDIES 15 A. Leaves 16 1. Previous studies 16 a. Phyllotaxy 16 "b. Epidermis 16 c. Vascular system 20 d. Crystals 21 e. Secretory organs 22 2. Materials and methods 22 3. Results and discussion 25 a. Epidermis 25 b. Venation 41 c. Crystals 5^ B. Seeds 55 1. Previous studies 55 a. Gross morphology 56 b. Internal morphology $8 c. Other 59 2. Materials and methods 60 3. Results and discussion 6l C. Seedlings 76 1, Previous studies 76 a. Morphology 76 "b. Nyctlnasty 79 ill 2. Materials and methods 81 3. Results and discussion 81 D. Flowers 94 1. Previous studies 94 a. Calyx 94 b. Enantiostyly 95 c. Anthers 96 d. Tapetum 97 2. Materials and methods 98 3. Results and discussion 98 E. SEM 117 1. Previous studies 118 2. Materials and methods 120 3. Results and discussion IV. CHROMATOGRAPHY STUDY 146 A. literature Review 146 1. History of chromatography as applied to taxonomy l46 2. Secondary substances l47 3. Previous studies in the Leguminosae 149 4. Previous studies in Cassia 15O B. Materials and Methods 152 C. Results and Discussion 154 V. CHROMOSOME NUMBERS 199 A. Previous Literature 199 B. Materials and. Methods 199 C. Results and Discussion 200 iv VI. CONCLUSIONS BASED ON STUDIES 214 VII. SERIES CORÏMBOSAE 216 A. General Information 216 B. Infrageneric Concepts 223 C. Distribution and Ecology 225 D. Artificial Key to the Taxa 22? E. Enumeration of Taxa 234 1. Cassia aurantia 234 2. Cassia 'bica'DSTJlaris 237 3. Cassia "bracteosa 260 4. Cassia candolleana 263 5. Cassia corymbosa 268 6a. Cassia floribunda var. floribunda 275 éb. Cassia floribunda var. scandens 298 7. Cassia hilariana 301 8. Cassia indecora 303 9. Cassia morongii 317 10. Cassia obLongifolia 325 11a. Cassia -pendula var. rendula 329 lib. Cassia pendula var. hracteosa 355 11c. Cassia -pendula var. erlocarpa 358 lid. Cassia pendula var. hemirostrata 36I 12a. Cassia santanderensis var. santanderensis 365 12b. Cassia santanderensis var. canarensls 372 12c, Cassia santanderensis var. .iahnli 376 13. Cassia subulata 380 14, Cassia tomentosa 383 F. Excluded Taxa 394 1, Cassia chinensis 394 2. Cassia blcapsularls var, chinensis 394 V 3. Cassia <d.andullflora 395 h. Cassia laevigata 395 5. Cassia T?entagonia 396 G. Taxa of Uncertain Disposition 396 1. Cassia bjcapsularis var, guadri.Tuga 396 2. Cassia "bonariensis 396 3. Cassia crassiseDala 397 4. Cassia frondosa 397 5. Cassia indecora var. advena 398 6. Cassia inflata 398 7. Cassia reinwardtil 399 VIII. LITERATURE CITED 400 IX. ACKNOWLEDGEMENTS 411 X. AEEENDIX ^15 1 I. INTRODUCTION Cassia is one of the largest genera of legumes (600+ species), and. it has been considered among the 25 largest of dicotyledons (Irwin and Turner I960). The genus consists of three subgenera—Cassia (Fistula of early workers), Senna, and Lasiorhegma—the latter two further resolving into nine sections. Subgenus Senna includes five sections; one of these, Chamaefistula. contains four series, of which Corvmbosae is the topic of the present study. Series Corvmbosae includes the Cassia bjcaipsularis complex (those plants with obovate terminal leaflets having rounded apices), considered to be one of the major imponderables in the genus (isely [lowa State University] and Irwin [The New York Botanical Garden]—personal communication). These are shrute and small trees (sometimes vine-like), naturally dispersed from Argentina to Mexico and the West Indies, in the northern part of the range Including both indigenous and introduced kinds. Conspicuous aspect-variance is exhibited in habit, flower size, length of pedicels, and rostration of the lowest anthers; in number, shape, color, and pubescence of leaflets; and in pod size and pubescence. Introduced plants from warm regions about the world show a similar, but less extreme, range of variation. Most of the material is Identified in hertaria as Cassia bicapsularis L. Bentham (1871) was the last worker to study the Cassia bicapsularis complex as a whole, treating it as a single species. He assigned nearly 30 names in synonymy to the group, and almost an eq.ual number have been 2 added to the complex since then. Subsequent comparative analyses of some of the taxa have been primarily concerned with the interpretation of cultivated material in various parts of the world (de Wit 1955» Brenan 196?} Isely 1975)» The initial objective of the present investigation was an attempt to introduce some order into the Cassia blcapsularis complex. It was soon evident, however, that the complex was not an entirely natural group, and that an understanding of it required a survey of all of the New World^ taxa of the seemin^y definable and natural series, Gorymbosae. Obviously also, interpretation of character trends within the series requires a comparison with comparable traits among other segments of the genus. A considerable portion of the research efforts herein reported was, therefore, directed toward the latter end. The overall problem has taken two phases» (l) a determination of characters—aiiatomical, biochemical, chromosomal, and morphological— common to taxa within series Gorymbosae. and the comparison of them with comparatiLe traits in other sections and series of the genus; (2) an assessment of the specific and varietal taxa of the series, and revisionary action where appropriate. Judgements have been based on examination of more than 2700 herbarium specimens and experience gained ^ Series Gorymbosae contains three African species (Bentham I87I). These are excluded from the present study, however, since their affinities do not seem to lie within the bounds of the series. As Bentham put it: "... the species of the two [areas—New Ïorld/Old World] are separated by characters which may perhaps indicate a separate origin, and should form perhaps separate groups." Thus, whenever the Gorymbosae are heretofore indicated, the New World taxa are intended. 3 in two seasons of field work in tropical America. The Corymbosae are herein defined to include 20 taxa—14 species and 6 varieties. There are two basic philosophies in writing a dissertation: (l) include all research, whether or not a certain part might be considered enlightening to the overall study; (2) include only parts which might be considered suitable for publication. In light of the usual procedure at this university, I have chosen the first philosophy, much to the dismay of the members of my thesis committee.