University of Kentucky UKnowledge Plant Sciences Science, Technology, and Medicine 1979 The Aloineae: A Biosystematic Survey Herbert Parkes Riley Shyamal K. Majumdar Click here to let us know how access to this document benefits ou.y Thanks to the University of Kentucky Libraries and the University Press of Kentucky, this book is freely available to current faculty, students, and staff at the University of Kentucky. Find other University of Kentucky Books at uknowledge.uky.edu/upk. For more information, please contact UKnowledge at [email protected]. Recommended Citation Riley, Herbert Parkes and Majumdar, Shyamal K., "The Aloineae: A Biosystematic Survey" (1979). Plant Sciences. 2. https://uknowledge.uky.edu/upk_plant_sciences/2 THE ALOINEAE THE UNIVERSITY PRESS OF KENTUCKY THE ALOINEAE A Biosystematic Survey . '. Herbert Parkes Riley ;' , Shyamal K. Majumdar Library of Congress Cataloging in Publication Data Riley, Herbert Parkes, 1904- The Aloineae, a biosystematic survey. Bibliography: p. Includes indexes. 1. Liliaceae. 2. Plant cytotaxonomy. I. Majumdar, Shyamal K., 1938­ joint author. II. Title. QK495.L72R53 584'.324 77-92927 lI.·laceae. 1. 1938- ISBN: 978-0-8131-5592-0 Copyright© 1979 by The University Press of Kentucky Scholarly publisher for the Commonwealth serving Berea College, Centre College of Kentucky, Eastern Kentucky University, The Filson Club, Georgetown College, Kentucky Historical Society, Kentucky State University, Morehead State University, Murray State University, Northern Kentucky University, Transylvania University, University of Kentucky, University of Louisville, and Western Kentucky University. Editorial and Sales Offices: Lexington, Kentucky 40506 Contents PREFACE vii Chapter One INTRODUCTION 1 Chapter Two GEOGRAPHICAL & ECOLOGICAL FEATURES 7 Chapter Three MATERIALS & METHODS FOR STUDY 25 Chapter Four KARYOTYPES 34 Chapter Five CHROMOSOME NUMBERS 42 Chapter Six POLYPLOIDY 66 Chapter Seven ANEUPLOIDY 77 Chapter Eight DELETIONS, DUPLICATIONS, & INVERSIONS 83 Chapter Nine TRANSLOCATIONS 97 Chapter Ten MISCELLANEOUS STUDIES 114 Chapter Eleven HYBRIDIZATION 135 Chapter Twelve EVOLUTIONARY PROBLEMS 143 LITERATURE CITED 161 INDEX OF FAMILIES, GENERA, & SPECIES 171 GENERAL INDEX 177 This page intentionally left blank Preface During the last fifty years the chromosomes of the Alo­ ineae have been the basis of a number of studies by cytol­ ogists and cytogeneticists in many parts of the world. The early studies were concerned with learning about the chromosomes themselves and their structure and behavior during mitosis and meiosis, two processes that were then just beginning to be understood. Later studies were con­ cerned chiefly with the evolution of the Aloineae, espe­ cially with chromosome factors involved in their evolu­ tion. Much more recently factors other than chromosomal that might have an influence on the evolution or biosyste­ matics of a group of plants have been investigated and are being appreciated; even thirty or forty years ago they were occasionally considered, although not so extensively. The studies on the Aloineae have been written in En­ glish, French, German, Italian, Japanese, Portuguese, and Afrikaans, and have been published in many journals (some rather obscure) in Europe, Africa, Asia, and North and South America. Some of the publications were by us and our associates. The purpose of this book is to assemble in one place all this published material and to try to interpret from the works of the various authors some evo­ lutionary trends and the possible taxonomic significance, if any, of the chromosomal and other factors operating in the development of the group. The extensive literature on Aloineae has been covered in this book as completely as possible. Only an occasion­ al rare foreign publication has been unavailable; one, first published in Portuguese, was of little concern since the original author later republished it in English in a better-known journal. Nearly all Japanese articles in­ clude English or German summaries; the only one that did not was primarily a list of chromosome numbers of various species of plants and therefore largely useful, even though neither of us is competent in even elementary Jap­ anese. The most important paper on the chromosomes of Aloe is the Ph.D. dissertation of F. S. MUller, which was published in Afrikaans. This language is little known in the United States but is easy to read; it is basically a simplified form of Dutch, with practically all the declen­ sions and conjugations eliminated, and has many words that are cognate forms of English or German words and therefore easy to assimilate. The senior author wishes to express his gratitude to many people and several organizations in South Africa and Preface vii the United States who have been helpful in the conduct of his research in the Aloineae and/or in the preparation of this summary. The late M. R. Levyns, formerly a senior lecturer in the Department of Botany in the University of Cape Town, and her husband, J. E. P. Levyns, provided in­ spiration and valuable suggestions and spent much time showing him botanical features of the landscape of the western Cape Province. Professor J. D. J. Hofmeyr, now retired from the Department of Genetics of the University of Pretoria, provided him space and research facilities during his trip to South Africa over twenty years ago. Other South Africans who were helpful primarily in supply­ ing seeds or live plants of the A10ineae were A. Berg of the University of Pretoria, Harry Hall of Kirstenbosch, and Hans Herre of the University of Ste11enbosch. Several organizations were particularly important in furthering this study. First and foremost was the Ful­ bright committee that provided the senior author the op­ portunity to visit South Africa in 1955-56 and to collect material, observe the plants in their natural habitats, and make contacts that enabled him to continue the work later. More recently, in 1963, an invitation to join in the celebration of the Golden Jubilee of the National Bo­ tanic Garden of South Africa and to spend a month on a conducted tour of the whole country enabled him to con­ tinue the observations he had made previously and to spend considerable time studying the collection in that fabu­ lously beautiful garden. To Professor H. Brian Rycroft, the director, he is very grateful. Indirectly the govern­ ments of the Union of South Africa in 1955 and its succes­ sor, the Republic of South Africa, in 1963 helped by pro­ viding him the necessary visas and permits to enter and study in their fascinating country. Grants from several organizations provided funds that really made this study possible. They include the Haggin Scholarship of the University of Kentucky and a National Science Foundation summer fellowship to the junior author. Grants from the Program of Genetic Biology and the Program of Systematic Biology of the National Science Foundation, from the Joseph Henry Fund of the National Academy of Sci­ ences, and from the University of Kentucky Research Foun­ dation were given to the senior author. Leaves of absence to visit and study in other coun­ tries are rarely granted as a matter of right but usually as a privilege. Therefore, the senior author is espe­ cially appreciative of three men who were responsible for his two leaves of absence to go to South Africa: Herman L. Donovan and Frank G. Dickey, both former presidents of the University of Kentucky, and M. M. White, dean emeritus of the College of Arts and Sciences. For many years the authors and their co-workers have been studying various aspects of this group of plants. We especially mention Debdas Mukerjee, who contributed much to the study of the Aloineae. Several authors, scientific journals, and publishers have given us permission to reproduce or redraw various illustrations from their published works. Specifically we would like to mention Peter E. Brandham of the Jodre11 Laboratory of the Royal Botanic Gardens at Kew, Gordon D. viii Preface Rowley of the Agricultural Botany Department of the Uni- versity of Reading, the Journal of South African Botany, the Canadian Journal of Genetics and Cytology, the Journal of Heredity, Chromosoma, The Nucleus (India), Advancing Frontiers of Plant Science (India), Grana, 30th Annual Proceedings of the Electron Microscopy Society of America, Pyton, The American Biology Teacher, Taxon, Plant Syste- matics and Evolution, the Ronald Press Company, and John Wiley and Sons, Inc. These permissions are gratefully ac­ knowledged. Herbert Parkes Riley Shyamal K. Majumdar Preface ix 1.1 1.3 Chapter One INTRODUCTION The tribe Aloineae of the family Liliaceae is fundamental­ ly a South African group, but some of the genera included in it are also found elsewhere. It is rather small and is variable, since not all authors include the same genera within it. As treated here the tribe will include Aloe, Gasteria, Haworthia, Astroloba (Apicra), Poellnitzia, Chamaealoe, Chortolirion, Lomatophyllum, Leptaloe, and Guillauminia. The Aloineae comprise a group of generally succulent plants characterized by having a loculicidal capsule and anthers with introrse dehiscence. The leaves are general­ ly thick or fleshy and often toothed on the margins; they often have a bitter sap that sometimes is used medicinal­ ly; and they contain a soft, green, water-storing tissue. The rootstock is a rhizome and is not bulbous, as it is in most of the Liliaceae. The flowers are generally in a ra­ ceme or panicle, and the segments of the perianth are con­ nivent or united into a tube at the base. Aloe Linn. (Sp. Pl., 1753, p. 319) is a large genus, is the most widely distributed geographically, and is probably the best known. The plants may be herbaceous and Opposite page: 1.1 about 23 cm tall (as in or arborescent Aloe di- Aloe myriacantha) chotoma; 1.2 Aloe speciosa; and about 15 m tall (as in A. eminens) and may be stemless 1.3 Aloe davyana; 1.4 Aloe or caulescent; if stems are present they are simple or marlothii.