This dissertation has been 65—3903 microfilmed exactly as received PINKAVA, Donald John, 1933- BXDSYSTEMATIC STUDY OF GENUS BERLANDIERA DC. (COMPOS1TAE). The Ohio State University, Ph.D., 1964 Botany University Microfilms, Inc., Ann Arbor, Michigan BIOSYSTEMATIC STUDY OF OENUS BERLANDIERA DC. (COMPOSITAE) DISSERTATION Presented in Partial Fulfillment of tha Requirement* for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By DONALD JOHN PINKAVA, B.Sc.t M.Sc. The Ohio State University 1964 Approved by Adviser Department of Botany and Plant Pathology PLEASE NOTE Figure pages are not original copy. They tend to "curl". Filmed in the best possible way. University Microfilms, Inc. ACKNOWLEDGMENTS I wish to express my sincere gratitude to Dr* T. Richard Fisher, my adviser, who not only suggested thia problem, but moat effectively guided it to ita completion* I am thankful alao to Drs* Clara G. Weiahaupt, Emanuel 0. Rudolph, Dale A. Ray and Carroll A* Swanson for reading thia dissertation and for their helpful criticisms and suggestions* Special recognition is extended to John M* Speer* Unless otherwise cited, the photography is to be credited to his talents so unselfishly shared* I am deeply indebted to Dr* Ray for statistical assistance; to the curators of the herbaria for loaned specimens and/or photographs; and to my colleagues for their many useful suggestions* Financial assistance was provided by The Ohio State Research Foundation and the National Science Foundation* ii TABLE OF CONTENTS Pag* ACKNOWLEDGMENTS....................................... 11 LIST OF T A B I E S ......................................... It LIST OF ILLUSTRATIONS................................. Yl INTRODUCTION........................................... 1 GENUS D E S C R I P T I O N ..................................... 6 HISTORY OF G E N U S ....................................... 12 ECONOMIC I M P O R T A N C E ................................... 16 ARTIFICIAL KEY TO HERLANDIERA T A X A ..................... 17 DESCRIPTION AND DISCUSSION OF T A X A ..................... 19 PROBLEMS OF NOMENCLATURE............................... 66 EXPERIMENTAL METHODS AND MATERIALS..................... 68 CYTOGENETICS ........................................... 75 CROSSABILITY........................................... 10A GERMINATION AND SURVIVAL............................... 113 ANALYSIS OF PARENTAL EXPERIMENTAL P L A N T S .............. 117 KEY TO AND ANALYSIS OF HYBRID EXPERIMENTAL PLANTS . 129 FIELD COLLECTION ANALYSIS (EAST) ....................... l46 FIELD COLIECTION ANALYSIS (WEST)....................... 157 DISTRIBUTION M A P S ..................................... 168 SUMMARY ............................................... 172 BIBLIOGRAPHY ........................................... 17^ AUTOBIOGRAPHY......................................... 178 111 LIST OF TABLES 1 Summary of chromosome count* reported in lltormturo .............................. 2 Chromosome data of current s t u d y .......... • 3 Pollen atainability data from fisld oollsctlons.................... .. 4 Genetic data supporting tstraploidy hypothesis .......... .................... 5 Chi-square tsst of gsnstic data supporting tstraploidy hypothssis ••*••••••• 6 Sslfing achene-set data ......... .......... 7 Summary of achsns-sst data from artificial crosses .............. ............ 8 Germination and ons-ysar survival records of artificial crosses .............. •••• 9 Germination data of field-collected fruits of eastern Berlandiera taxa .............. 10 Morphological analysis of parental experi­ mental plants .......... ................ 11 Morphological analysis of artificial hybrid plants ••••»•• ...................... 12 List of herbaria specimens used in hybrid index analysis of eastern taxa complex • • . 13 Hybrid index analysis of eastern taxa complex .................................. 14 Analysis of variance between populations of eastern taxa complex 15 Analysis of variance between species-hybrid groups of eastern taxa complex •••••• iv LIST OF TABLES (Continued) Table Page 16 Liat of herbaria specimens used in hybrid index analysis of western taxa complex . • . 158 17 Hybrid index analysis of western taxa complex • • 160 v LIST OF ILLUSTRATIONS Figure Page 1 Berlandiera texana DC. (Holotype illustration)............................ 7 2 Berlandieralyrata Benth. var. lyrata (Holotype .......................... 21 3 Berlandiera lyrata var. macrophylla 6ray (Holotype, Sheet 1) . .......... 32 Berlandiera lyrata var. macrophylla Oray (Holotype, Sheet 2) . .......... 32 5 Berlandiera aubacaulia Nutt. (Holotype) .... 37 6 Berlandiera x humilia (Small) comb. nov. Tfeolotype of B. humilia Small)............ kz 7 Berlandiera x humilia (Small) comb. nov. (Ieotype of B. humilia Small) ....... kz 8 Berlyidiera pumila (Michx.) Nutt. (Holotype) . ........................ kk 9 Berlandiera pumila (Michx.) Nutt. (Holotype of Polymnia caroliniana Poiret) .......... kk 10 Berlandiera pumila (Michx.) Nutt. (Holotype and paratypea of B. tomentosa var. deal­ ba t a T. & 0.) . 7 ........................ k6 11 Berlandiera pumila (Michx.) Nutt. (Paratypea of B. tomentosa var. dealbata T. & G.) . 12 Berlandiera texana DC. (Iaotype) ............ 5^ 13 Berlandiera texana DC. (Holotype of B. longifolia Nutt.) ...................... 51* l*f Berlandiera x betonicifolia (Hooker) comb. nov. (Syntype of silphium betonicifolium Hooker) . 7 7 . ... *1 vi LIST OF ILLUSTRATIONS (Continued) 15 Berlandiera x betonicifolia (Hooker) comb, nov. (Syntype of Silohium betonicifolium Hooker) 16 Microsporogenesis I ...................... 17 Microsporogenesis II ...................... 18 Microsporogenesis III .................... 19 Microaporogenesia I V ............... • . 20 Microaporogeneaie V ...................... 21 Microsporogenesis VI .............. .... 22 Heteromorphic Pollen ...................... 25 Corolla Colora ............................ 2*+ Leaf Shapea .............................. 25 Pubescence Types I ........................ 26 Pubescence Types II .......... ...... 27 13. lyrata var. lyrata 1 0 1 0 ................ 28 B. texana 1002 ...•• .................. 29 B. x betonicifolia 655 .................... 30 B. x betonicifolia 657 .................... 31 — * Puaila 525 ............................ 32 <B. subacaulls 101*+........................ 33 lyrata var. lyrata 35-*+................ B. lyrata var. lyrata x subacaulls 5-5 . • • LIST OF ILLUSTRATIONS (Continued) Figure Page 35 B. subacaulis 11-2 .... .................. 131 36 B. texana 660-1 .................. ..... , 13 2 37 B. lyrata var. lyrata x texana 40-2 ........ 132 38 B. lyrata var. lyrata x oumila 255-1 ..... 132 39 B. texana x oumila 197-3 ..................... 134 40 B. texana x subacaulis 2-1 ................... 134 41 B. oumila x subacaulis 170-2 ................ 134 42 B. lyrata var. lyrata x B. x betonicifolia i i 8 - i .................... ....... 135 43 B. texana x B. x betonicifolia 176-1 ........ 135 44 B. x betonicifolia x B. pumila 242-1 ........ 135 45 B. x betonicifolia x B. subacaulis 144-1 . 136 1*6 offspring of B. lyrata var. lyrata x F2 oumila 31-1 ............................... 136 offspring of B. subacaulis x texana 469-1 . 156 1*7 F2 48 Histograms of hybrid index values of eastern and western taxa complexes .............. 163 49 Distribution map of eastern t a x a ........ .. 168 50 Distribution map of western taxa ............ 170 viii INTRODUCTION The genus Berlandiera was described by Augustin-Pyrasue De Candolle in 1828 in honor of Jean Louis (or Luis) Berlandier who had served as his eaissary to the then relatively unknown Texas region and as a scientist of the Mexican Boundary Coaaission (1827-1829)* Although born into a poor faaily near Fort-de-l'£cluse, France, Berlandier's initiative and keen interest in natural sciences enabled hia, as a young aan in his early twenties, to be the first to collect extensively and explore the Texas boundary region* According to Qeiser (1937) and McKelvey (1933)* the travels of Berlandier extended froa Mexico City to the Trinity River via Laredo and San Fernando de Bejar along the Old Bejar Road* The expedition's objective was Nacogdoches, but Berlandier and others of the party contracted aalaria* The scientific staff never crossed the Trinity River but were forced back to Bejar by June, 1828* During this tiae Berlandier collected a series of speciaens that later were naaed in his honor, Berlandiera texana DC* With Berlandier ill and his speciaens daaaged again and again by the incleaent weather, the elder De Candolle's hopes of a large and valuable collection froa a virgin land faded* Subsequent sojourns were aade later in 1828 and again the following year but none included the type locality of B* texana* After the Commission had completed its task, Berlandier remained in Mexico, married and settled at Matamoras. He made still other excursions, particularly one to Goliad and Bejar in l$3k. He continued his investigations until his death in 1851 when he drowned in an attempt to cross the San Fernando River south of his home. As to his scientific accomplishments, McKelvey (1955) lists his "Memories de la Comleion de Limites. Historia Natura por El General Teran y L. Berlandier" (1832) which contains descriptions of eleven newly proposed species and four new genera; his handwritten Expedition (in U.S. National Museum); co-authorship with Raphael Chovel in 1850 of Diario de viage de la comision de limites aue puso el gobierno de la republica.