A Systematic Study of the Genus Lasiacis (Gramineae, Panicoideae) Gerrit Davidse Iowa State University
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Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1972 A systematic study of the genus Lasiacis (Gramineae, Panicoideae) Gerrit Davidse Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons Recommended Citation Davidse, Gerrit, "A systematic study of the genus Lasiacis (Gramineae, Panicoideae) " (1972). Retrospective Theses and Dissertations. 4731. https://lib.dr.iastate.edu/rtd/4731 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]. INFORMATION TO USERS This dissertation was produced from a microfilm copy of the original document. 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University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 A Xerox Education Company 73-9438 DAVIDSE, Gerrit, 1942- A SYSIBIATIC STUDY OF TŒ GENUS LASIACIS (GRAMDŒAE, PANICOIDEAE). Iowa State University, Ph.D., 1972 Botany- University Microfilms, A XEROX Company, Ann Arbor, Michigan THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. A systematic study of the genus Lasiacis (Gramineae, Panicoideae) by Gerrit Davidse 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. In Charge of Major Work Signature was redacted for privacy. For the Major Department Signature was redacted for privacy. For the Graduate College Iowa State University Ames, Iowa 1972 PLEASE NOTE: Some pages may have indistinct print. Filmed as received. University Microfilms, A Xerox Education Company ix TABLE OF CONTENTS Page INTRODUCTION 1 METHODS AND MATERIALS 4 GROSS MORPHOLOGY 13 LEAF ANATOMY 25 BREEDING SYSTEM 43 CHROMOSOME NUMBERS 47 HYBRIDIZATION 54 FRUIT DISPERSAL 85 TAXONOMIC TREATMENT 97 LITERATURE CITED 224 ACKNOWLEDGMENTS 230 1 INTRODUCTION Lasiacis is a small neotropical genus of about 16 species of grasses distributed from the southern tip of Florida and central Mexico south through the West Indies, Central America, and in South America to southern Peru and northern Argentina. Many species are rather striking in appear ance because of their large, erect, woody culms. However, an almost equal number of species have procumbent, creeping, herbaceous culms. As striking are the shiny, black mature spikelets. All species are of little or no economic importance. Lasiacis was first delimited as a section of Panicum (Grisebach, 1864). Hitchcock subsequently elevated the section to a genus (Hitchcock and Chase, 1910). This was followed by a treatment of all species north of Colombia (Hitchcock, 1920). This revision has continued to form a solid basis for later work in Lasiacis, as has the bulk of Hitchcock's work for all subsequent American agrostological research. Since Hitchcock's 1920 revision, published later in very similar form in North American Flora (Hitchcock, 1931), most systematic work with Lasiacis has been limited to regional floristic treatments, new species descriptions, and species trans fers. The most important contributions were made by Hitchcock (1930, 1936), Parodi (1943), and Swallen (1934, 1936, 1943, 1953, 1955a, 1955b, 1957). Systems of grass classification at the tribe and subfamily level relied largely on gross morphological features of the spikelets and inflorescences during the early part of this century. In recent years classifications have in addition been built on anatomical, cytological. 2 physiological, and a wider array of gross morphological characters. Sig nificant in the use of such a wide array of characters in constructing a system of grass classification was the pioneer effort of Avdulov (1931). He used size and base number of chromosomes, leaf anatomy, size and posi tion of first seedling leaves, type of starch grains, and the type of nucleoli in non-dividing cells to propose a new system of classification. Important among the new systems of classification that have used these characters are those of Prat (1960), Parodi (1961), and Stebbins and Crampton (1961). Although differing in detail, these authors agree in recognizing six subfamilies which are, using the Stebbins and Crampton (1961) nomenclature; Bambusoideae, Oryzoideae, Arundinoideae, Festucoideae, Eragrostoideae, and Panicoideae. These new systems are phylogenetic and probably reflect genetic relationships much better than previous systems. The broad outlines of this new system seems rather stable at the present time. There are many genera for which the new characters are not or only partially known, and these genera are placed in tribes and subfamilies strictly on the basis of spikelet and inflorescence structure. Lasiacis is such a genus. Lasiacis has never been monographed in its entirety. No biosystematic studies have been made, as in most tropical plants; such studies will prob ably give a better understanding of the relationships between species. My interest in Lasiacis first developed while collecting, identifying, and cytologically examining Costa Rican grasses with Dr. Richard W. Pohl of Iowa State University. As a result of these studies, I felt that existing taxonomic treatments for Lasiacis were incomplete. 3 My reasons for a detailed study of the genus were; 1. to delimit and provide a taxonomic treatment for all species of Lasiacis 2. to suggest possible evolutionary relationships among the species 3. to study the evolutionary forces that might have been important in the development of Lasiacis 4. to place the genus in its natural tribe and subfamily. 4 METHODS AND MATERIALS Herbarium Studies Data gathered from herbarium specimens included morphological measure ments and observations, species distribution, ecological observations reported by collectors, as well as vernacular names that have been applied to the plants. Since Lasiacis has been rather extensively collected, I borrowed specimens only from some major herbaria, marked by an asterisk in the following list. Collections in other herbaria were studied ^ situ or specialized collections were borrowed. The herbaria are cited according to Lanjouw and Stafleu (1964), except for Universidad Central de Venezuela at Maracay, assigned MÏ by Venezuelan botanists (Bunting, personal correspon- dence). A Arnold Arboretum; Cambridge, Massachusetts AWH Natuurwetenschappelijk Museum; Antwerpen, Belgium C Botanical Museum and Herbarium; Copenhagen, Denmark EAP Escuela Agrlcola Panamericana; El Zamorano, Honduras F* Field Museum of Natural History; Chicago, Illinois GH* Gray Herbarium of Harvard University; Cambridge, Massachusetts IJ Science Museum, Institute of Jamaica; Kingston, Jamaica ISC* - Iowa State University; Ames, Iowa L - Rijksherbarium; Leiden, Netherlands MEXU* - Herbario Nacional del Institute de Biologia, Universidad Nacional de Mexico; Mexico City, Mexico M - Botanische Staatssanmlung; Munchen, Germany MO* - Missouri Botanical Garden; St. Louis, Missouri 5 MÏ - Universidad Central de Venezuela; Maracay, Venezuela NY* - New York Botanical Garden; New York, New York P - Muséum National d'Histoire Naturelle, Laboratoire de Phanirogamie; Paris, France TRIN - University of the West Indies; St. Augustine, Trinidad UC* - University of California; Berkeley, California US* - United States National Museum; Washington, D.C. VEN* - Institute Botanico; Caracas, Venezuela Field Studies I have been able to study Lasiacis in the field in the northern half of its distributional range. The following trips have been