Cytotaxonomy of the Genus Echinochloa in Louisiana. James Howard Brooks Louisiana State University and Agricultural & Mechanical College
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Louisiana State University LSU Digital Commons LSU Historical Dissertations and Theses Graduate School 1969 Cytotaxonomy of the Genus Echinochloa in Louisiana. James Howard Brooks Louisiana State University and Agricultural & Mechanical College Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses Recommended Citation Brooks, James Howard, "Cytotaxonomy of the Genus Echinochloa in Louisiana." (1969). LSU Historical Dissertations and Theses. 1640. https://digitalcommons.lsu.edu/gradschool_disstheses/1640 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 70-9040 BROOKS, James Howard, 1932- CYTOTAXONOMY OF THE GENUS ECHINOCHLOA i IN LOUISIANA. j The Louisiana State University and Agricultural and Mechanical College, Ph.D., 1969 Agronomy University Microfilms, Inc., Ann Arbor, Michigan CYTOTAXONOMY OF THE GENUS ECHINOCHLOA IN LOUISIANA A Dissertation Submitted to the Graduate Faculty of the Louisiana State University Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Botany and Plant Pathology by James Howard Brooks B.S., Stephen F. Austin State College, 1957 M .S., Stephen F. Austin State College, 1964 ACKNOWLEDGEMENTS The author wishes to express his sincere appreciation to Dr. Clair A. Brown for serving as chairman of his committee, for aid in selecting the problem, guidance throughout the study, and helpful suggestions for improving the manuscript and companionship during some of the collecting trips. Thanks are due to the other members of his committee, Dr. St. John P. Chilton, Dr. Irwin Forbes, Dr. M. T. Henderson, Dr. B. Exner, and Dr. J. B. Baker, for their encouragement and helpful criticism in the preparation of this dissertation. The author is grateful to the Department of Botany and Plant Pathology for cooperation and assistance in respect to utilization of equipment, materials and greenhouse space. The author also wishes to thank the curators and staff of all the herbaria who supplied their herbarium specimens and to thank other persons who supplied plants and seed for this investigation. Thanks are due also to Mrs. Denise Sylvester for securing several interlibrary loans. He is grateful to Mrs. Jackie Lockwood for typing the final draft of this manuscript and her invaluable assistance during the past three years. Finally, he wishes to express his deep appreciation to his wife, Vera, for encouragement at all times and limitless patience in typing this dissertation. TABLE OF CONTENTS Page ACKNOWLEDGEMENTS...................................................................................... ii TABLE OF CONTENTS ...................................................................................... iii LIST OF TABLES . v LIST OF FIG U RES................................................................................................ vi ABSTRACT. ................................................................................................... vii INTRODUCTION................................................................................................... 1 HISTORICAL ACCOUNT...................................................................................... 2 MATERIALS AND M ETHODS............................................................................ 8 CYTOLOGY.............................................................................................................. 11 Chromos ome s ............................................................................................... 11 Method of Reproduction........................................................................ 17 Correlation of Pollen Size and Chromosome Number .... 18 ANATOMY................................................................................................................ 22 Leaf Epidermis............................................... ' ......................................... 22 Embryo-Caryopsis ................................................................................. 24 ANALYSIS OF HERBARIUM SPECIM ENS...................................... 30 Discussion of Taxonomic Characters ..................... 34 TAXONOMIC TREATMENT............................................................................. 38 SUMMARY....................... 69 iii Page LITERATURE CITED.................................................' ........................................ 71 APPENDIX.............................................................................................. 75 VITA....................................................... 84 iv LIST OF TABLES TABLE Page I. Concepts of Echinochloa taxa correlated with Weigand (1921).................................................................................... 6 II. Mean diameter of pollen grains in different taxa of Echinochloa .................................................................................... 21 III. The difference in caryopsis length and embryo le n g th ....................................................................................................... 29 IV. Mean values of some quantitative characters of Echinochloa taxa................................................................................. 32 V. Ranges of some quantitative characters of Echinochloa ta x a ............................................................................... 33 v LIST OF FIGURES FIGURE Page 1-7. Chromosomes of Echinochloa taxa at diakinesis. 15 8. Photomicrograph of a meiotic anaphase I of E. colonum (n = 2 7 ) ...................................................................... 16 9. Photomicrograph of an endosperm nucleus showing approximately 81 chromosomes (3n) and 54 chromosomes (3 n ) ...................................................................... 19 10-19. Abaxial epidermis of leaves and sheaths of grasses of various Echinochloa ta x a ................................................... 26 20. Caryopsis of various taxa of the genus Echinochloa . showing a longitudinal cross-sectional view .................... 27 21. Caryopsis of various taxa of the genus Echinochloa, showing the dorsal view ............................................................. 28 22. Polygons of Echinochloa ta x a ............................................... 31 23. Spikelets of various taxa of the genus Echinochloa. showing the nature of the spikelet and relative size. Each spikelet is shown in two views, dorsal and ventral....................................................................................... 37 24. Maps showing the distribution of E. crusgalli. E. colonum and E. colonum form JL in Louisiana. 6F 25. Maps showing the distribution of E_. muricata. E.. muricata var. microstachya and E. occidentalis in Louisiana. .............................................. .... 67 26. Maps showing the distribution of E. zelayensis. E. walteri and four forms in Louisiana. ................... 68 vi ABSTRACT The present investigation was undertaken to determine the taxonomic relationship of various taxa of the genus Echinochloa in Louisiana. The investigation included studies of anatomical, cyto- logical and morphological characters of each taxon within the genus. The characters used for delimitation of the tasa were, anther, caryopsis, fertile floret, embryo, palea of lower floret, sheath, spikelet, sterile lemma, panicle, pustulate based setae on the branches of panicle, length of panicle branches, length and width of blades, ligule, and presence or absence of stamens in the lower floret. Based on the above characters, twelve taxa of the genus Echinochloa were recognized in Louisiana, namely, E. colonum (n = 27), E. colonum form JL (n = 27), E. crusgalli (n = 27), E. muricata (n = 18), E. muricata var. microstachya (n = 18), E_. occidentalis (n = 27), E. walteri (n = 18), E. walteri form 1_ (n = 18), E. walteri form_2 (n = 18), E. walteri form 3^ (n = 18), 13. walteri form A (n = 18) and 12. zelayensis (n = 18). A key is provided for the above taxa, based on material from specimens taken from L o u isiana. v ii INTRODUCTION Echinochloa. the sixth largest grass genus in the tribe Paniceae (Hitchcock, 1951), is one of the less economically important genera. Most of the taxa are grazed by stock but are not usually utilized as forage due to the sparse growth habit. Some of the cultivated varie ties of E. colonum and E. crusgalli are used as a source of food in tropical Asia and tropical Africa. In Louisiana, E. colonum and E. crusgalli are weeds in the rice fields as well as other agronomic crops. Echinochloa is one of the important sources of food in the coastal marsh for the water fowl. The taxa of Echinochloa are widely distributed throughout the tropical and temperate regions of the world. There is no agreement as to the exact number of polymorphic species represented in this genus. A total of twenty-one species has been described in North America alone. Yabuno (1962) reported about twenty species world w id e . The most recent treatment of the genus Echinochloa in North America was by Hitchcock (1951). It was evident from the use of this treatment that an extensive investigation of the genus Echinochloa was necessary. The present study, utilizing an anatomical,