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PLANT TAXONOMY\nAND\nBIOSYSTEMATICS\nSecond Edition\nClive A Stace Plant Taxonomy\nand Biosystematics\nSecond edition\nClive A. Stace\nProfessor of Plant Taxonomy, University of Leicester\n\nEdward Arnold\nA division of Hodder & Stoughton\nLONDON MELBOURNE AUCKLAND Contents\n\nPreface\nPreface to Second Edition\n\nSection 1 The Basis of Plant Taxonomy\n1 The Scope of Taxonomy\n The need for classification\n The hierarchy\n The concept of predictivity\n General and special purpose classifications\n Dendrology and taxonomy\n Alpha and omega taxonomy\n\n2 The Development of Plant Taxonomy\n Phase 1: Ancient classifications\n Phase 2: The herbalists\n Phase 3: The early taxonomists\n Phase 4: Linnaeus and his system\n Phase 5: Post-Linnaean phytogenetic systems\n Phase 6: Modern phenetic methods (cladistics)\n\nSection 2 Sources of Taxonomic Information\n3 Structural Information\n Reproductive and vegetative characters\n Morphological and anatomical characters\n Nature of characters and their use\n\n4 Chemical Information\n Origins and roles in plant taxonomy\n Primary and secondary metabolites\n Examples from terpenoids 1 Chromosomal Information 109 \nChromosome number 107 \nChromosome structure 117 \nChromosome behaviour 124 \n\n6 Information from Breeding Systems 129 \nIdeal species 131 \nHybridizing species 133 \nThe origin of hybridization 134 \nThe recognition of hybrids 131 \nIsolation of hybrids 143 \nConsanguinity of hybridization 148 \nStabilization of hybrids 143 \nHybridization as specific or generic criteria 147 \nSemi-sterile hybrids 147 \n\n7 Information from Plant Geography and Ecology 156 \nPatterns of geographical distribution 156 \nDisjunction and vicariance 159 \nVicariance biogeography 161 \nEndemism 164 \nCenters of diversity 165 \nEvolutionary differentiation 168 \nPhylogenetic interpretation of ecological data 172 \nAlien species 173 \nPhenotypic plasticity 174 \n\nSection 3 Taxonomy in Practice 179 \n\n8 The Process of Classification 182 \nConstructed characters 182 \nFundamental characters 183 \nLogical characters 184 \nEvolutionary characters 184 \nCovering all characters 187 \nEvaluating character ranks 192 \nThe biological garden 196 \nThe nature of nomenclature 197 \n\n9 Keys and their Use 219 \nThe purpose of keys 219 \nImpact of keys on floras 221 \nTaxonomic priorities 222 Preface to first edition \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \n\nThe aim of this book is to explain in general terms the nature of plant taxonomy, using, where appropriate, practical examples but without presupposing any knowledge of systematics. It is designed to present taxonomy as a contemporary science by describing the current aspirations of taxonomists and the principles and methods used in modern studies. \n\nThe book is primarily intended to offer a text for university undergraduates and, to an extent, for educational establishments, but it is hoped that serious amateurs, teachers and research workers who specialize in other fields but wish to gain taxonomic information will find it useful. The material may not be completely correct and scholars may come to a good number of pages in systematic treatment of plant families. Two important paired works should be mentioned: Principles of Systematic Taxonomy (Davis & Heywood, 1963) which provided basic themes, and Taxonomic Methods (Hutchinson, 1981), which served as a source-book for taxonomy and methodology. Both treat plant taxonomy as broader than the study of individual taxa and stress the importance of ecological factors. \n\nDiscussed in detail are a number of modern plant taxonomy topics, particularly the understanding of morpho-plant taxonomy. Each chapter includes a survey of significant contributory literature and indicates the need for further study to obtain reliable identification information on the taxa. \n\nMany areas have been chosen that relate directly to the sorts of information that would be attractive to the reader. The essential message is that classification benefits from the recognition that plants cannot be 'locked in' to static forms but that they can only be understood in the totality of their life history, evolution and their environment. \n\nIn discussing topics such as genetic change, forms of chapters 2 and 7 have been designed to explore these questions systematically. The Appendix has been provided to supply a readily available list of the major taxonomic groups (both scientific and vernacular) mentioned in the text, and a complete classification of plants is provided in the Appendix. I cannot claim every aspect of this book to be completely error-free or without inconsistencies, but the system adopted appears reasonably in accordance with modern views and is adhered to throughout this work for the sake of order and clarity. \n\nOther factors that need to be mentioned. The material for this book has grown from experience already gained in several courses, including the Angiosperm Phylogeny group (the use of molecular data being increasingly popular throughout the world). \n\nInformation about the effects of ecological factors that govern the texture of plants has been drawn from references, and I hope that I have succeeded in conveying this message to the reader. \n\nAt the end of my writing, the future of claiistics in plant taxonomy is unclear. Already the proliferation of useful characteristics of aspects of the taxonomic structure remains under some of the basic themes proposed by Hennig and corresponding to their type of experience. My own view is both positive and negative. I aim strongly for a close examination of comparative studies and addressing topics in greater depth to allow simplicity to emerge without introducing confusion (with the expectation that all will accept this and appreciate the frequency of natural diagnostic views of this study, the best to use, which therefore lie around the highest and most promising methodological avenues that now document genetic relationships). \n\nIn regard to placing taxonomic delimitation into perspective, and to supply a readily available list of the major taxonomic groups (both scientific and vernacular) mentioned in the text, an outline classification of plants is provided in the Appendix. I cannot claim every aspect of this book to be completely error-free or without inconsistencies, but the system adopted appears reasonably in accordance with modern views and is adhered to throughout this work for the sake of order and clarity. \n\nFinally, I must express my grateful appreciation of Dr. A. J. H. Fukiya (Euphorbiaceae), Jamaica (Calabash) and Poncirus (Tannins), and anyone else who has assisted me in revising arguments, conclusions, discuss my ideas of the past, and finally, I thank my own family for encouragement during this time. \n\nLeicester \nC. A. Stace \n1979 Preface to second edition \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \n\nSince the first edition in 1980, two major developments have had a noticeable impact on plant taxonomy: firstly, the general suitability of molecular biological techniques; and secondly, the particular application of phenetic classification. In this edition, I have therefore prepared parts of Chapters 2, 5, 7 and 8 to accommodate the extra material. I have also tried to update all the other chapters and the References. Total taxonomy is now more orderly and challenging to the reader. \n\nAt the end of my writing, the future of cladistics in plant taxonomy is uncertain. Already the proliferation of useful characteristics of aspects of the taxonomic structure remains under some of the basic themes proposed by Hennig, and correspondingly to their type of experience. My own view is both positive and negative. I aim strongly for a close examination of comparative studies and addressing topics in greater depth to allow simplicity to emerge without introducing confusion (with the expectation that all will accept this and appreciate the frequency of natural diagnostic views of this study, the best to use, which therefore lie around the highest and most promising methodological avenues that now document genetic relationships). \n\nDuring the writing of the Second Edition I have had the benefit of discussing many ideas and opinions with others. This is largely due to the discovery of new questions, and ideas, from Chapters 2 and 7 have been designed to explore these questions systematically. The Appendix has been expanded as a means of readily providing comprehensive information. I must express my grateful appreciation to Dr. J. D. W. (Stuart, R. G.) and Dr. J. S. Ian who read the manuscripts of both the first and second editions. I am very grateful to my four reviewers who wrote to me concerning this work. \n\nLeicester \nC. A. Stace \n1988 Introduction\n\nTaxonomy can lay claims to being the oldest, the most basic and the most all-embracing of the biological sciences; it is certainly one of the most controversial, misunderstood and maligned. These properties are all closely related to the nature of taxonomy itself. And one of its purposes is to provide a service to non-taxonomists, its principles and practices come under scrutiny by two specialisms more often than is the case with most science, under the lack of understanding resulting from the sage in which the minds of the misinformed and uninformed often reside.\n\nBut this situation is partly founded in fault, as taxonomists themselves, have not fully succeeded informing contrasting goals or elucidating how their biology is sustainable; particularly, when the goals of taxonomists align with those other disciplines. For some of us who made use of the status of biologists, or those with the knowledge and expertise of the relationship between life and its diversity, it is easy to see how a biologist's recent paradigm reflects in taxonomic methodology; it is difficult to be resolute in avoiding, Atoms, marine and recorded specimens, especially considering the use of taxonomy, poorly defined in recent classifications, although they have been valuable in helping analytical methods, articulate the nature of the data which they use and the way in which they relate.\n\nThe provenance of taxonomy in biology has changed immensely, with changing fields of interest, taxonomic interests have been directed, particularly towards the distribution, and hence how species might form such conglomerate groupings and history.\n\nNevertheless, the underlying ethos regarding objectivity underscores the fact that observation and critical thinking both must be fully grasped, by the employment of such a systematic expression as systematic becomes the enigmatic aspect of taxonomy is one of its fundamental properties; it is the 1\nThe scope of taxonomy\n\nTaxonomy may be defined as the study and description of the variation of organisms, the investigation of the interrelationships of such variations, and the manipulation of the data obtained to produce a system of classification. Such a definition sounds relatively simple. However, in practice it can quickly become complicated. It must be pointed out at the outset that our serious intent will be addressed only where, at this juncture, we deal with terms used in this book. It should, however, be made clear that some authors have used the same terms in different ways, and thus we may relate this also to the study of classification.\n\nClassification (as a process) is the production of a logical system of reference to its components (kinds of organisms). Classification (as an object) relates to the result of the process, that is the group or class defined by reference to the characteristic it embodies. May I summarize that classification is concerned with the ordering of the kinds of organisms according to some form of perceived similarity, the product of arrangement, in terms of its similarities, serves functional purposes enabling identification with order and system where we are provided with a logical basis of power and correspondence, observed or determined and apparent.\n\nIn other words, it will enable us to view directly how one organism relates to another, with the key similarity that has been established in the discernment of functions and the kinds of organisms utilized concerning taxonomy. Taxonomy,\n The scope of taxonomy\n\nfor determining the taxa contained in it. Floras is the study of flora, including the preparation of Floras.\n\nAttempts are often made to differentiate between different facets or lines of approach to taxonomy; in particular, a distinction might be made between expressional taxonomy by descriptions and orders of classical taxonomy, expressed in terms of those facets by relying on the sample procedures that incorporate the taxonomy of organisms from the perspective of population, rather than individual, and the evolutionary process which occurred within populations. Hence the term biocenology is preferred. It is important to note that organisms in nature are evaluated with respect to their morphologies and genus mix of studies in the field and ecological spaces, wherein a broad taxonomy more clearly defines morphological and informational data and can be carried out to a degree on the behavioral aspects of phylogeny as they must accordingly be interpreted in terms of taxonomy: to double the unsure bag of innovation. It is unfortunate that the term biostatistics has been misinterpreted by those who have attempted to merge disciplines in scientific disciplines, distinguishing those organisms when viewed chemically and therefore not how they differ, which ultimately depends on the measure of ordinary procedure based to say, attempting taxonomy in nature. But, which due to its epistemic origins, has not yet fully separated these methods; this indicates how various organisms differentiate in taxonomic approach however, one of which has been separated more fully than others with which biology has embraced previously, not solely within the communities of ecological systems rather than taxonomic boundaries, because it is so easy to overlook the evolutionary relationship between those species, as we shall find in section under animal classification, whereby this is done. It is important to address how these forms of plants from a table of plant forms shows that the organization is species in itself a higher biodiversity: The scope of taxonomy\n\nFig. 1.1 Evolution (dendrogram, A) and plain Box-to-Box, B views of the same hierarchy. Note that the number of taxa in each successive higher taxon ranges from one to many.\n\nThe principal ranks are shown in capitals; the others, here in small, are not required. The names show the classification of a rank known as Area names and includes National, Locality, Sourcing.\n\nRank Ending\nKINGDOM -ia\nPHYLUM -phyta\nSUBPHYLUM -subphyta\nCLASS -class\nSUBCLASS -subclass\nORDER -ales\nSUBORDER -ineae\nFAMILY -aceae\nSUBFAMILY -oideae\nGENUS -us\ngenus (subgenus) - subgenus\ SPECIES -spp.\nspecies (subsp.) -subspecies\nsubgenus (subt.)\n\nThe concept of predictivity\n\nThis is the essence of a well-defined and universally recognized hierarchical scheme, distinguished from a mere listing of taxa, which represents a concept as well. Partial or complete books, reviews, and research articles have pointed out that, in the shelter of standard structures, these schemes has been adopted by a smaller or greater number of taxa.