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Thesis-1960D-M498c.Pdf (3.787Mb) CY'l'OT.AXONOMIC STUDY OF THE Dichanthium annulatum COMPLEX By KHARAITI LAL MEHRA,, Bachelor of Science Delhi University Delhi, India 1952 Master of Science Delhi University Delhi, India 1954 Submitted to the faculty of the Graduate School of the Oklahoma State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY May, 1960 OKLAHOMA STATE UNIVERSITY LIBRARY SEP 1 1960 CYTOTAXONOMIC STUDY OF THE Dichanthium _!!nnulatum COMPLEX Thesis Approved: ;7nean of the Graduate School 4528'10 ii DEDICATION To the memory of the Late Dro Robert Paul Celarier whose researches have contributed so nru.ch to our knowledge of the difficult genera of the Andropogoneae iii ACKNOWLEJ:GEMF.NTS I would like to take this opportunity to express my sincere ~rat= iiude to Professors J. R. Harlan and the late R. P. Celarier for ·their il;lteree-t in my scientific growth, encouragement, and constructive C!'itiei~m throughout the course of this study. I con~ider it a ~·eat wivil~ge to have 1,mrked under their guidance on the -world ns largest i c-pllection of the Dichanthium annulatum complex. I ~cknowledge my indebtedness to Drs. J. S. Brooks, L. H. Bruneau, a,.1d J. A. Whatley, Jr. for their assistance and guidance in this in- ~stiga;tion. I wish to express Iliy thanks to Dr. B. B. Hyde of Oklahoma un1ver- sity, N·orman, Oklahoma for his valuable suggestions and constructive criticism in preparing the final manuscript. I -would also like to take this opportunity to express my grat= itude to Professor and Mrs. Edgar Anderson and their daughter Miss Phoebe Anderson of Missouri Botanical Gardens, St. Louis, Missouri for their keen interest in the development of my understanding of the semigraphical_methods and for their hospitality during my residence with them in the spring of 1958. l also appreciate the help and cooperation of Mi:-. W. L. Richardson anc;! Mr. Ro M. .A.bring D I must acknowledge my indebtedness to the Agronomy Department and ,, the Agricultural Experiment Station, Oklahoma State University for iv the research assistantship and the facilities, without which this in= vestigation could not have been completed. My sincere appreciation is also due to the Department of Botany and Plant Pathology for making the laboratory facilities available to me throughout the present studyo I express my special thanks to Dr. Jo S. Bakshi, Mr'. Ao Ho Gill, Mr. S. Ao Faruq19 rlfr o Do So Borgaonkar, and Mr. Ao Po Singh for their help during this studyo I am also thankful to Mr·s. Shirley Tracy for completing the pains= taking work of typing this dissertationa I am also thankful to my parents, Dro Po No Bhaduri, Dro S. L. Tandon,and Mr. K:rishen Lal for their interest in my education, without which my study in the United States would have been impossibleo V TABLE OF CONTm!TS Page IN'l'RODUCT ION a 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 REVIEW OF LITERATURE. 5 Taxonomy 5 Historical. 5 Deseriptive. 6 Geographical Distribution. • 11 Cytology • 12 Cytogeography. '. 12 Meiotic behavior. • 13 Cytotaxonomy. • 14 MATERIALS AND MEI'HODS. • 15 Accessions • • 15 Herbarium s_pecimens • 0 15 Cytological studies • • 16 Morphological studies • 17 Hybridization between the Tropical and the Mediterranean type • 0 17 RESULTS . 0 • 19 Cytology . 19 . Cytogeography • • 19 Diploids • 0 • 21 Tetraploids • • 21 Hexaploids 30 External Morphology. • 3.3 Diploids • • 36 Tetraploids • • 36 Hexaploids 0 40 Cytogeography and Morphological Variation 0 4.3 Relationships of the Tropical and Mediterranean Types • • 45 F1 hybrids • 0 45 F2 population. 0 46 Cytology. 46 Morphology 0 47 Origin of Jl. papillosum Stapf 0 54 Fl hybrids • . 54 F2 hybrids. 0 54 DISCUSSIO:tf, • • 60 SUMMARY 0 72 vi Page SELECTED BIBLIOGRAPY. 75 APPEND.ID'S • 81 vii LIST OF TABLES Table Page I • .Artificial and NatUTal Hybrids between the Tropical and Mediterranean Types of the 12· annulatum Complex. 18 II. Class Intervals of the Cytological Features in the 12· annulatum Complex 24 III. Comparison of the Range of Morphological Variation in the Diploid, Tetraploid, and Hexaploid Accessions of the]. annulatum Complex. 35 IV. Class Intervals of the Morphological Characters in the 12• annulatum Complex 38 V. Comparison of the Morphological Characters in the Parents, F1· Hybrids, and F2 Populations of the ~. annulatum Complex. 49 viii LIST OF FIGURES Figure Page 1. Geographieal Distribution of Ninety=three Collections of the ]2. annulatum Complex. 20 2. Frequeney Histograms Showing Cytological Variation in the 12· annulatum Complex 25 3. Pictorialized Scatter Diagram Showing the Cytological Variation in Forty Tetraploid Accessions' of the 12· annulatum Complex. 26 4. Histogram Illustrating the Frequency Distribution of Cytological Index Values in Forty Tetraploid Accessions of the 12· annulatum Complex. 27 5. Frequency Histograms Showing Morphological Variation in the ]2 • .annulatum Complex. 34 6. Pictorialized Scatter Diagram Illustrating the Morphological Variation in the 12· annulatum Complex. 37 7. Pi.ctorialized Seatter Diagram Showing Introgression between the ]2. annulatum Complex and the ]. intermedia Complex. 41 8. Histogram· Illustrating the Frequency Distribution of Morpholog'ical Index Values in Fifty=three Accessions of the ]2. annulatum Complex. 42 9. Cytogeography· and Morphological Variation in the 12· annulatum Complex • 44 10. Frequency Histograms Showing Morphological Variation in the F2 Plants Obtained by Selfing the 4n F1 Hybrid between the Tropical and the Mediterranean Type. 48 11. Morphological Analysis of the F2 Pop.1lation Obtained by Selfing the 4n FJ. Hybrid between the Tropical and the\Mediterranean type of the 12• annulatum Complex. 50 12. Frequency Histograms Showing Morphological Variation in the F2 Plants Obtained by Selfing the 6n F]. Hybrid between the Tropical and the Mediterranean Type 55 ix Figure Page 13. Morphological Analysis of the F2 Pop.1lation Obtained by Selfing the 6n F1 Hybrid between the Tropical and Mediterranean Types of the ]2. annulatum Complex. 56 1~ The Origin and Evolution of the D. annulatum Complex. 64 X LIST OF PLATES Plate Page I. Meiosis in ]2. annulatum X 1180. 22 II. Meiosis in ]2. annulatum X 1210. 31 xi INTRODUCTION Most of the forage·cirops grown in the United States are introduc= tions from Europe, AsiaJ and Africao Only a relatively small number of native grasses and practically no native legumes are used as cultivated forage plantso In many cases these introductions have been far super= ior to native species in productivity, persistence under use, aggres= siveness, and in some cases, forage quality (Celarier and Harlan, 195S)o The introduction of :materials has not been very systematic until reci:mt years, and some of the better adapted species, or ecotypes with= in a species, perhaps have not yet been evaluatedo Furthermore, only a few of these introduced species have been collected throughout their geographical range of distributionj even though at least some of them have been known for some time to be of considerable agronomic importanceo It has been recognized that several members of the grass tribe !ndropogonea~ have forms with considerable potential as pasture and range forage (Celarier and Harlan, 1955; Harlan et aL, 1958) o Of the genera that have been studied, Dichanthiumj Bothriochloa, and Capilli= pedil.Hl± are among those which have forms with certain desirable agronomic characterso Although, the group as a whole has much potential, no single collection has a.11 of the desirable ·agronomic qualitieso It -Will be necessary to develop plant breeding methods to utilize selected mem= bers ~f these grasses in the development of altogether new combinations of forms with desirable agronomic qualitieso It has been suggest,ed that if the vigor, productivity, and high quality of some collections i ·l 2 of ~~CJ}fil!'J;,Jti.wm could be combined with the winter hardiness. of some col= lections of -~°-t_t4'19£1}1oa~ strains of great, value to the Great Plains might be obtained (Harlan et al"J 1958L A systematic :survey of the morphological and cytological variation prevalent -within different species of these genera would be helpful in selecting the desirable col= lections to be used in the breeding p:rogramo Althoughjl these genera are of consideTable impo:rt,ie.m.ie in the Old World and seem to have much prom= ise for the New World, they are poorly understood cytologfoally and taxono:micallyo Only an analyBis of Jl9.t.n£19~Jll.,9.~ A~9...h~~-mi,vn, Keng has heen more or less completed (Celar.i.erj) 19571 Celarier and HarlanJ 1958) and a similar suirvey ls needed for the other species of these generao The objecctive of this study was to grow togetherj in an experimen= tal garden~ s,everal int,rodu«::Jed a©ccessfons of the lt:!J:J~~11:!;;hi1±~ !i~l~~;ld.lll;! complex collected throughout it:s geographfoal range of dlstribution, and to condui.::t both a taxonom.foal and cytologfoal BUll:"vey of theroo The carrying out of such a.n obJ ecctive should provide information from whfoh to dra:w conclus:i.ons rconJ~erning the pattern of va.riati.on · prevalent in this complex, and also about Hrs origin and evolutionary mechanicso Plr'eliminai"y cytological and taxonorrd.i:l studies (Cela.rier and HsrlanJ 1956, Celarier et ala~ 1958, Mehra and CelarierJJ 19:58; Harlan et aL 1 1958) have pointed out the presence of' a polyploid series within the l2cl~Jl~ll-J,P.J,.'U,.ID /:lt}W,;,11.).J:.,Ji!A1!11 complex with somat,fo ichromosome numbers of 20j 40)1 and 600 Four mi:irphological types {Tropical, Mediterx0 anean, Senegal, and South
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