aan FER 4 bra xe 784 : Biochemical Systematics and Evolution ANDREW FERGUSON, B.Sc., Ph.D. Lecturer in Zoology The Queen's Univesity of Belfast First published 1980 Blackie Glasgow and London.CHAPTER ONE INTRODUCTION TO SYSTEMATICS, SYSTEMATICS IS REGARDED BY SOME BIOLOGISTS AS SYNONYMOUS with taxonomy. By others, such as Blackwelder (1967), Mayr (19696) and Simpson (1961), it is used in a comprehensive sense to cover the study of the diversity of organisms and their relationships. In this book, systematics is used in the widest sense, including aspects of the study of evolution (the process by which the diversity of living organisms is produced) and taxonomy (the process by which this variation is arranged into a meaningful and useful order). Both evolution and taxonomy involve a study of variation among organisms. Biochemical systematics is the study of biochemical variation, which here will be restricted to a consideration of nucleic acid and protein macromolecules, Within the field of systematics, several other terms are used with various meanings by different authors. Adequate definition of all terms would require an entire book, so consideration will be given here only to those necessary for an understanding of later chapters. The term classification overlaps with taxonomy, but generally itis restricted to that part of taxonomy which involves the arrangement of organisms into groups in an hierarchical system (likewise called a classification) on the basis of their relationships. Populations and species If all the organisms living in a particular area are studied, it is found that, although individuals, may be unique if examined closely enough, they ‘occur in groups of similar individuals with a number of features in common, Such groups of recognizably similar individuals constitute populations. In outbreeding sexual organisms, the individuals in a population interbreed among themselves, exchanging genes frecly, and2 BIOCHEMICAL SYSTEMATICS AND EVOLUTION are said to share a common gene pool. To distinguish it from other uses ‘of the term ‘population’, this is sometimes referred to as a Mendelian population (or gamodeme in botanical useage) and it is with this connotation that the term is used subsequently. ‘The population, not the individuals comprising it, is the natural unit of evolution and is therefore the basic unit used in systematic studies. A species is a group of one or more populations which is given a formal distinguishing name. If the populations, or groups of populations, are sufficiently distinct, they may be given sub-specific names. In goneral, the individuals that comprise a species are capable of inter- breeding to produce viable and fertile offspring but, due to the ‘geographical spread of the constituent populations, they may not have the opportunity to do so. Individuals of one species are, as a rule, reproductively isolated from individuals of all other species. However, ‘occasional o localized inter-breeding (or inter-breeding in captivity) does not negate the specific status of the participants. Obviously if two populations are sympatric (live in the same area) without inter-breeding, they are separate species. If they are allopatric (live in mutually exclusive areas with an unoccupied zone between), evidence of reproductive isolation is not available, and delimitation of species is then generally ‘based on their possession of certain features which are not present in others. Normally these features are aspects of internal or external structure (ie. morphology), and such species are referred to as ‘morphospecies, as distinct from biospecies, which are described on the basis of the biological species concept of reproductive isolation. Since the biological species concept does not apply to asexual, parthenogenetic, or self-fertilizing organisms, here again morphospecies are the rule, The term agamospecies has been applied to all species that reproduce asexually. Several terms have been proposed for species delimited by biochemical characters, but these terms (such as physiological species) are misleading and superfluous. Evolution of diversity—speciation Classifications are necessary because of the immense variability of organisms. This diversity occurs both within populations, and among populations and species. Variation can either be genetically or environmentally induced, or can be the product of a combination of these two forces. Genetic variation is produced in organisms by mutation and itis this variation which is the basis of all evolution. In most species, more offspring are produced than can survive if the species numbers are to remain stable. For example, individual female fish such as the cod Gadus ‘morhua produce upwards of ten million eggs each year, but only two INTRODUCTION TO SYSTEMATICS 3 offspring need survive to replace their parents over their reproductive lifespan. Since individual offspring differ in their genetic make-up, they will differ in their ability to cope with the environment in which they find themselves. Those which survive are those which are genetically best adapted, and thus they in turn will produce the next generation. Natural selection is the name given to this process, whereby the best-adapted individuals survive to reproduce. Random processes may also be important in deciding which individuals survive. The relative importance of natural selection and random events has been the subject of debate among biologists for many decades. Due to mutation, over-production of offspring, natural selection and stochastic events, the genetic composition of a population will change gradually, ic. it will evolve. Over the generations there is a constant refinement of the genetic composition of the population, as the genes selected for are those best adapted to the conditions in which the population lives. However, the environment is constantly changing, and @ stable situation is not usually attained for a long time, since new adaptations are frequently necessary. Over a period of many thousands of years, a species will change in its genetic make-up and consequently in its structural and other features. After a time it may have changed sufficiently for the taxonomist to regard it as a different species. This process, which is a change within a single lineage, is called anagenic or phyletic speciation (figure 1.1). species 8 mutation and lection species A ‘gonatc eitforentiation| Figure 1.1. Digg ilostating he rosso angen seston