Biosystematic Studies in the Genus Silphium L.(Compositae): Silphium
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ThU dissertation has been microfilmed exactly as received _ _ - ,. 66-15,144 SWEENEY, Carol Reichert, 1937- BIOSYSTEMATIC STUDIES IN THE GENUS SILPHIUM L. (COMPOSITAE): SILPHIUM COMPOSITUM MICHAUX. The Ohio State University, Fh.D., 1966 Botany University Microfilms, Inc., Ann Arbor, Michigan BIOSYSTEMATIC STUDIES IN THE GENUS SILPHIUM L. (COMPOSITAE): SILPHIUM COMPOSITUM MICHAUX DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Carol Reichert Sweeney, A.B., M.S. The Ohio State University 1966 Approved by Department of Botany and Plant Pathology ACKNOWLEDGMENTS I wish to express my sincere appreciation to my adviser, Dr. T. Richard Fisher, for suggesting this problem and unselfishly aiding and advising in the work. Gratitude is also extended to Drs. Emanuel D. Rudolph, John A. Schmitt and Ronald L. Stuckey who read this paper and offered many helpful criticisms. Special recognition is given to John M. Speer who provided several photo graphs and gave much help in photography, cytological methods and collection of specimens; to Dr. Dale A. Ray who aided in the statistical treatment; and to Mr. Joseph Harvey who did much of the art work. Many thanks are extended to Drs. J.W.A. Burley and Jerry McClure for suggestions and help with the chromatographic work. The curators of the following herbaria are gratefully thanked for their kindness in loaning specimens and photographs: Chicago Natural History (Field Columbian) Museum (F) Florida State University (FSU) Gray Herbarium, Harvard University (G) Michigan State University (MSC) New York Botanical Garden (NY) Philadelphia Academy of Natural Sciences (PH) United States National Herbarium (US) University of Indiana (IND) University of Kansas (KANU) University of North Carolina (UNC) University of Tennessee (TENN) University of West Virginia (WVA) University of Wisconsin (WIS) ii VITA January 9» 1937 • • Born— Meadville, Pennsylvania 1939 ....... A.B., Antioch College, Yellow Springs, Ohio 1960-1962 ........ Kettering Research Laboratory Fellow, Department of Botany and Plant Pathology, The Ohio State University, Columbus, Ohio 1962 .............M.S., The Ohio State University, Columbus, Ohio 1962-1965..... ..... Teaching Assistant, Department of Botany and Plant Pathology, The Ohio State University, Columbus, Ohio 1965-1966.... ..... Research Assistant, Department of Botany and Plant Pathology, The Ohio State University, Columbus, Ohio FIELDS OF STUDY Major Field: Botany Studies in Experimental Plant Taxonomy. Professor T. Richard Fisher Studies in Cytogenetics. Professor E. F. Paddock Studies in Plant Physiology. Professor Carroll A. Swanson iii c o n t e n t s Page ACKNOWLEDGMENTS .................................................ii VITA ............................................................ iii TABLES ......................................................... v ILLUSTRATIONS ................................................ wii INTRO DUG TI ON .................................................. 1 GiJ-i’ji': CijAT 'HAL i*ISTT RY ■ •*•#•••*••**•#*♦•• 5 GENERAI MORPHOLOGY OP q,:E GEL ITS ............................... 10 TAXONOMY......................................................... 13 ECOLOGY OF SILPHIUM COMPOSITUM ............................... MORPHOLOGICAL ANALYSIS OF RESEARCH GARDEU CLONES AITD POPULATIONS COLLECTED IN THE F I E L D .........................^6 BIG CHEMICAL S CoTEMATICS: BACKGROUND .V.: DBASIS ................. 70 METHODS AND RESULTS CP CHROMATOGRAPHIC AN A L Y S I S ............... ?6 CYTOLOGY OF THE SILPHIUM COMFOSITUM RESEARCH SARDS, CLONES . 101 INTERSPECIFIC AMD INTEASFSCIFIC HYBRIDISATION OF SILPHIUM COMPOS IT U M ....................................... 112 THE OCCURRENCE OF NATURAL IMTERSPLCIFIC AMD INTRASPECIFIC HYBRIDISATION ............................................ 123 DISCUSSION AMD CONCLUSIONS ................................... 123 SUMMARY ........................................................ 139 BI3LIC5RAPHY ................................................... l'+l iv m ► ■ ; Table Page 1 Diagnostic characters cf Silphium conpositum subspecies 40 2 numbers of 3. compositum subspecies associated with forest and soil types 42 3 humbers and. collection sites of research garden clones of iilnhiiuo comuositun . • .......... , be 4 Population analysis of £. c cropsiturn ssp. composittwi cO and ssp. renifcme .......... ...... 5 Population, analysis of jj. compositum ssp. conpositum, ssp. venosun, and ssp. ovatifolium ........ 62 6 Collection sites of gog: al~ tiers used in polygono- grapli analysis .................. 63 nI Chromatogram running tires ..... ........... •• 34 3 Characterization cf spots found on two-dimensional paper chromatograus cf 3. compositun ....... 8s 9 Characterization spots found on two-dimensional thin-layer chromatograms of ii. compositun . • 87 10 Chromatographic profiles of 3. corpooitun sub species from paper cbr'rr.atograr.s ......... 38 11 Chromatographic profiles of 3. conpositum sub species from thin-layer chrcmatograns ...... 39 12 Paired affirity values cf paper c’.rcaatograns of 3, conpositum arranged in subspecific groups . • 90 13 Paired affinity values cf tiir-layer chromatograms of 3. corrposituc arranged ir. subspecific groups . 91 14 Pesalts of Dancen’s hultigle Pr.r.ge Test for Signifi cant Differences between <,h*: sear, paired affinity values of the intrasubspecific comparisons for paper and thin-layer chrcuetcgrans ........ 93 v TABLES (continued) Table Page 15 Results of the analysis of variance of the paired affinity values cf intrasubspecific and inter- subspecific comparisons fro:.: thin-layer chroma bo grams ...» , , , 95 16 Results of the analysis of variance of the paired affinity values of intrasubspecific and inter subspecific comparisons from paper chromatograms • 96 17 Results of the analysis of variance of the paired affinity values of intraspecific and interspecific comparisons from paper chromatograms cf clone standards of £. conpositum and ntlicr oilphium species in "end arrangement" groups A,. 3, and C • 98 18 hean paired affinity values from thin-layer chromatograms of 7,'2159 tested for reproducibility of the m e t h o d s ............................ 99 19 Cytological behavior of meiotic chromosomes in 3. compositnm ssp, compositun c l o n e s........... » 103 20 Cytological behavior of neiotic chromosomes in C. compositum esp, rer.iforme clones 104 21 Cytological behavior of meiotic chromosomes in 3. compositun ssp, venocmn c l o n e s .................105 22 Cytological behavior of meiotic chromosomes in S. compositum ssp, ovatifolium clones *«•,•• 106 23 Results of interspecific and intraspecific hybridization of S. compositum . • . 116 24 Crossability of S. compositun with Silphium taxa found in the same habitat, other sympatric taxa and allopatric taxa •«••*•*•• ............ 127 vi ILLUSTRATIONS Figure Page 1 Silphium compositum Kichx* (Copy of holotype photograph; I T~............................ 15 2 Silphium orae Smell (Holotype)................. 17 5 Achenes of compositum ssp. compositun (#2162) • 18 k Achenes of S. compositum ssp. conpositum (#1309) * l3 5 Distribution map of S. compositum ssp. compositum and ssp. compositun-reni forme inter grades . 1$ 6 S. compositum ssp. compositum (#131^* field collection) ................................ ... 21 7 S. compositun ssp. conpositum (#1301, field CGllec tion) 21 3 compositum ssp. compositum-reniforne intergrade (#1327» field collection) . .................. 23 9 3. compositum ssp. compositum-reniforme intergrade (#1317| field collection) • 23 10 Distribution map of S, compositum subspecies . ?M 11 Silphium. conpositum ssp. reniforme (liaf. e£ IJutt.) Sweeney & Fisher comb. nov. (Lectotype) .... 26 12 Achenes of S. compositum ssp. reniforme (#2165) • 26 13 Distribution map of compositum ssp. reniforme . 28 1^ S. compositum ssp. reniforme (#2 1 6 6 , garden collection) ............................. 30 15 Silphium compositum ssp. venosum (Snail) Sweeney & Fisher c omb. nov. (Holotype) T ............ 30 16 Silphium lapsuum Small (Holotype) ........ 31 17 Achenes of S. compositum ssp. venosum (#1238) . • 31 vii ILLUSTRATIONS (continued) Page Distribution map of S. compositun ssp. venosum and ssp. ovatifolium ..•••• 33 19 S, compositun ssp. venosum (,^123^, garden collection) ................................... 35 20 Silphium conpositum ssp. ovatifolium (T. Sc G.) Gweeney Zt Fisher comb. nov. (holotype, Sheet 1) • 35 21 3. compositum ssp. ovatifolium (T. Sc 3.) Sweeney 8c Fisher comb. nov. (holotype, Sheet 2) ...... 37 22 Achenes of S, compositun S3]:.ovatlfoliun (,/-12^5) • 37 23 S. compositun ssp. ovatifolium (#123^. garden collection) ........................................... 3° 2k S* compositum ssp. ovatifolium (£l2*+5, field collection) ....... 38 23 Flowering periodicity in oilphium compositum subspecies ...o................ kk 26 Pictorialized scatter diagram of research garden clones of 3, compositum ssp. compositum and ssp. reniforme and 3. te rebin tliinaceum ........... 30 27 Pictorialized scatter diagram o' research garden clones of S. compositum ssp. compositum, ssp. venosun and ssp. ovatifolium......................... 32 28 Graph of mean hybrid number against mean hybrid index for natural population samples of compositum ssp. compositum and ssp. reniforme • 36 29 Graph cf mean hybrid number against mean hybrid index for natural population samples of S. compositum ssp. compositurn, ssp. venosum and