Taxonomy—Definitions, Terms and History a - 4-1. Introduction = Variation is the rule of the Nature: The amount of diversity in the living world is staggering. About more than one million species of animals and falf a million species of plants have already been described and estimates on the number of still undescribed living species range from three to ten millions. “An estimate of half a billion for the extinct species is consistent with the known facts. Furthermore each species may exist in numerous different forms, e.g. sexes, classes, seasonal forms, morphs and other phena. It would be impossible to deal with this enormous diversity if it were not ordered and classified. Systematic Zoology endeavors to order this rich diversity of the animal world and to develop methods and principles to make this task possible. 1-11, DEFINITION OF TAXONOMY The term Taxonomy is derived from the Greek words taxis, arrangement, and nomos, law. It was firstproposed, in its French form by de Candolle (1 813) for the theory of plant classification. Later on it was accepted for zoological classification too. Accord- ~ ing to Simpson, “taxonomy fs the theoretical study of classification, Including tts bases, pri neiples, procedures ‘and rules.” Ernst Mayt (1969) has defined it as “the theory and practice of classifying organtsms.”” 1-1:2, ROLE OF TAXONOMY The role of taxonomy in biology.can be briefed as below :— 1. It works out for. us a vivid picture of the existing organic diversity of our earth and is the only science that does 80. 2. It provides much of the information permitting a recon- struction of the phylogeny of life.*. 2 Animal Taxonomy and Museology 3. It reveals numerous interesting evolutionary Phenomena and thus makes them available for casual study by other branches of biology. 4. It supplies, almost exclusively, the informations needed for entire branches of biology. 5. It supplies classifications which are of great heuristic and explanatory value in most branches of biology, e.g. evolu- tionary biochemistry, immunology, ecology, ethology and historical geology. : 6. It is indispensable in the study of economically or medi- cally important organisms. In the hands of its foremost exponents it makes important conceptual contributions for example population thinking, not otherwise so easily accessible to experimental biologists. Thus, it contributes significantly to the broadening of biology and toa better balance within biological science as a whole. - 1-2. Definition of Systematics : The term systematics is derived from latinized Greek word systema, as applied to the systems of classification developed by the early naturalists. According to Simpson (1961) “systematics ts the scientific study of the kinds and diversity of organisms and of any and all relationships among them'’, or more simply, ‘systematics is the sclence of the diversity of organisms”. -The term ‘relationship’ is not used in a narrow Phylogenetic sense, but is broadly conceiv- ed to include all biological relationships among organisms. This explains why such a broad area of common interest has developed between systematics, evolutionary biology, ecology and behavioral biology. 1-3. Definition of Classification : According to Simpson (1961), “zoological classification ts the ordering of animals into groups or sets on the basis of their relation- ships, that is, of assoclations by contiguity, similarity or both’ Thus Classification coincides largely with what is sometimes designated as beta taxonomy, 1-31. STAGES IN CLASSIFICATION The classification of a given taxonomic group goes through Several stages of maturation before it is establisbed. In the first stage, called alpha taxonomy, the emphasis is on the description aTaxonomy—Definitions, Terms and Historp 3 of new species and their preliminary arrangement in comprehen- sive genera, In the next stage, referred to as beta taxonomy, relationships are worked out on the species level and on that of the higher categories. In other words here emphasis is on the development of a sound classification. In the third stage, f.e. the level of gamma taxonomy much attention is paid to, intra-specific variation, to various sorts of evolutionary studies ad to a, casual interpretation of organic diversity. The three levels discussed above overlap and intergrade. 1-3-2. ECOLOGICAL CLASSIFICATION Ecological classification defines sets according to such criteria as the communities in which the organisms, live, for example, soil micro-organisms and forest insects or other swamp plants, fresh water fishes environmental factors such as alpine plants, and cave salamanders. , 1-33. TELEOLOGICAL CLASSIFICATION Teleological classification defines seté according to their usefulness or lack of it, usually withresp¢ct to man. Such sets might be, for example, domesticated animals, with meat animals, draft animals, pets etc. as subsets; eUible, non-edible, and poisonous fishes or herbs classed according to the diseases for which they are considereds peclfics. 1-4. Nomenclature : : “1¢ 1s the application of distinctive names to'each of the groups recognized in classffication.”” roader than taxonomy and includes it as well Systematics is bi as classification and nomenclature, which are narrower than taxo- nomy and are, ina somewhat different way included in it. This relationship among systematics, taxbnomy, classification and nomenclature can be represented as under -— l<-Taxonomy Systematics<| . |<-Classification<-Nomenclature +5. Important. Taxonomic Terms : Some impertant terms, frequently used in taxonomic litera- ture are as follows :— Allochronic species—Species’ which do not occur at the same time level. Synchronic species—Species which occur at the same time level.< Antmal Taxonomy and Museology Alpha taxonomy—The level of taxonomy concerned with the characterization and naming of species. Beta taxonomy—The level of taxonomy concerned with the arranging of species into a natural system of lower and higher taxa. Gamma taxonomy—The level of taxonomy dealing with various biological aspects of taxa, ranging from the study of intra. specific populations to studies. of specimen and of evolutionary rates and trends, Binominal nomenclature—Tho system of nomenclature, adopt- ed by the International Congress of Zoology, by which the scienti- fic names of an animal is designated by both the generic and specific name. Biota—The fiora and fauna of a region. Cladism—A taxonomic theory by which organisms are order- ed and ranked entirely on the basis of “recency of common des- cent”, that is, on the basis of the most recent branching point of the inferred phylogeny Ecospecies—A group of Populations so related that they are able to exchange genes freely without loss of fertility or vigor in the offspring. Gause's rule—The theory that no two species with identical ecological requirements can co-exist in the same place. Genus—A category for a taxon including one species or a group of species, presumably of common phylogenetic origin, which is separated from related similar units (genera) by a decided gap. Sympatry—The occurence of two or more Populations in the same area ; more precisely the existence of a population in breed- ing condition within the cruising range of individuals of another Population. Category—It designates rank or. level in a hierarchic classi- fication. It isa class, the members of which are all the taxa assigned a given rank. Cline—It isa gradual and nearly continuous change of a character in a scries of contiguous populations ; a character gradient. A term coined by J.S, Huxley (1939), it may be smooth or they may be step clines with rather sudden changes of value,Taxonomy—Definitlons, Terms and History 5 Deme—lIt is a local population of a apecies ; the community of potentially interbreeding individuals at a given locality. Key—It is a tabulation of diagnostic characters of species or genera etc. in dichotomous couplets facilitating rapid identi- fication. Key character—In taxonomy a character of special utility in a key is known as key character. Monophyly—The derivation of taxon through one or more lineages from one immediately ancestral taxon of the same or lower rank, . Phenon—There is no generally accepted technical term for a phenotypically reasonably uniform sample. Camp and Gilly (1943) coined the term phenon (pl. phena) for this, It isa sample of phenotypically similar specimens. Sibling species—Pairs or groups of closely related species which are reproductively isolated but morphologically identical or nearly so. Sob-species—It is an aggregate of phenotypically similar populations of a species, inhabiting a geographic sub-division of the range of a species and differing taxonomically from other populations of the specics. 4 Taxon—It is a taxonomic group of any rank. that is sufficiently distinct to be worthy of being distinguished by name and to be ranked in a definite category. According to Simpson taxon is “a group of real organisms recognised as a formal unit at any level of a hierarchic classification”. Its pleural is taxa. Variety—It is an ambiguous: term of classical taxonomy used for a heterogeneous group of phenomena including non- genetic variations of the phenotype, morphs, domestic breeds and geographic races. 1-6. History of Taxonomy : The history of taxonomy is related with the history of man- kind. So far no authentic history of taxonomy has been written. Only superficial and descriptive attempts ‘have been made from time to time to give an idea of history of taxonomy in the stan- dard histories of zoology and botany, In fact, it remained impossi- ble to give an authentic history of taxonomy as long as roots of taxonomic method and principle in essentialism : in Aristotelian a6 Animal Taxonomy and Museology logic : an Arbitrarily the history of tax six periods :-— 1. First period—Therstudy of local fauna. 2. Second period—Linnacus and his contemporaries. 3. Third perlod—The Empirical approach. 4, Fourth period—Darwin and phylogeny. 5. - Fifth period—Population systematics. 6. Sixth period—Current trends. The account of these periods of facilitates the understanding of the progress that has been made in the subject. The complexity of taxonomy must be kept in mind when studying its history. All periods are not strictly separated from each other and sometimes various trends are concurrent during a single period. Progress in the taxonomy. of various animal groups and in the study of animals from different regions of the world has been very uneven. Taxonomic studies are most advanced in the most popular groups of animals e.g. birds ; butterflies ; mammals and beetles, while in others it may still be on an elementary level. It is most advanced in north temperate zone and lagging behind in the tropics and ger distant. places. Lét us discuss now each period in some details. . 1-61. FIRST PERIOD—The Study of Local Fauna : Natives of even. the most primitive tribes may be excellent naturalists, with specific names for local trees, mammals, birds, fishes and the more conspicuous or edible invertebrates. A tribe of Papuans in the mountains of New Guinea was found to have 137 specific names for 138 species of birds. Only one species was confused with another. Often the nomenclature of such tribes is clearly binominal, hence it is not the sole invention of Linnaeus. Several early Greek scholars, notably Hippocrates (460—377 B.C.) and Democritus (465—370 B. C.) studied different kinds of animals, but there is no indication of a useful classification in their works. The key figure of this period was Aristotle (384—322 B. C.) who brought together the knowledge of his time and formulated it ints the pogcnaing ofa science. He did not propose a formal ication of animals, but i i Sleseifontion, of ani t he provided the basis for such a d in nominalism were not understood. onomy is divided into followingTaxonomy—Definitions, Terms and History 7 according to their way of living, thetr actions, thelr habits and thetr bodily parts. He referred to such major groups of animals as birds, fishes, insects and whales, distinguishing among insects mandibulate and haustellate types and winged and wingless conditions ; he used specific terms for lesser groups e.g. Coleoptera and Diptera, which persist even today. He also established numerous collective categories, or genera, using as differentiating characters such as blooded or bloodless; two-footed or four-footed; hairy or feathered; with or without an outer shell and so forth. Aris- totlc suggested to naturalists the idea of arranging all animals in a single graded scala naturae according to their degree of ‘perfec- tion’. This led to the endeavor to classify animals into those that are ‘lower’ and those that are ‘higher’, at the grade of organisation. The ‘typological’ or ‘essentialist’ thinking is also proposed by Aristotle. Since all this was a tremandous advance over anything that hdd previously existed so it completely dominated animal taxonomy for the next 2000 years. It is because of this contribu- tion, Aristotle is sometimes called as, ‘Father of biological taxonomy”. John Ray (1627—1705) was first who recognised the difference between genus and the species and who through evaluation of both similarities and dissimilarities in animals, arrived at a more natural higher classification than did those who had gone before him. On the whole, through out this period the botanists were far ahead of the zoologists and were the first to look new methods and new principles. 1-62, SECOND PERIOD — Linnaeus and his Contemporaries ; Linnaeus (1707—1778) exerted such a great influence on the entire subsequent development in taxonomy that he has been called the “Father of modern taxonomy”. In the tenth edition of his Systema naturae (1758), the binominal system of nomenclature was for the first time consistently applied to animals and this work of Linnaeus became the foundation of modern systematic zoology. In addition, his work was characterized by clear cut species diagnoses and by the adoption ofa blerarchy of higher categories : genus, order and class. These methods of Linnaeus were by no means wholly original, but his eminently practicalAnimal Taxonomy and Museo) logy 8 system was quickly adopted, expanded and elaborated because ige and the influence of his followers The L . le his great personal presti, Linnaean philosophy dominated taxonomy for the next centy, and most of the essentials of his methods are still components of _the modern taxonomy. 1-6-3. THIRD PERIOD—The Empirical Approach : This period, between the tenth edition of the Systema naturae and the publication of Darwin’s Origin of species, may also be termed as period of explorations. It was characterised by an intense interest in the fauna of far away places, expeditions and in the. accumulation of vast number of specimens from all over the world. A steady and enormous increase in the number of known animals characterized this period. The local naturalists were being replaced by specialists who studied birds, reptiles, molluscs and similar specific groups or indeed smaller groups The taxonomic work of this period of these specific groups. confirmed the conviction of the early naturalists that the endless variety of organic life is organised into natural groups. Deductive, a priori principles were increasingly rejected and more and more taxonomists delimited taxa empirically, on the basis of the totality of characters, not just a few essential ones. The empirical taxonomists of this perlod did a magnificent job in developing a ‘Natural System’, in the new meaning of this term. The empliricists also ‘undertook the evaluation of gaps between taxa, and they supported hierarchial arrangement of categories on the basis of degree of similarity. 1-6-4, FOURTH PERIOD—Darwin and Phylogeny : Charles Darwin (1809-1882) in 1831 joined as a naturalist on Voyage of the Beagle. During this voyage, he encountered various phenomena of distribution, variation, structure and adaptation which caused him to think of evolutionary interpretation. By this the problem of Natural system was solved. “Natural” groups exist because the members of such a group had descended from a common ancestor. This evolutionary theory supplied the expla- nation for the fact that variation in nature is not continuous but consists of ‘groups within groups”. The interest of this period was pre-eminently phylogenetic.Taxonomy—Definitions, Terms and History 9 The chief effect of the acceptance of the theory of evolution on taxonomy has consequently beena greater pre-occupation with phylogeny. Darwin realized that two processes occur during phylogeny. These are: (i) Branching, and (ii) Subsequent divergence. Accordingly Darwin stressed that the separation of taxa must be based on branching (propinquity of descent), but that in the ranking of these taxa into various categories due consideration must be given “‘to the different degrees of modifications which they have undergone”. Darwin, like empiricists, rejected both the priori weightings of taxonomic characters as practiced by Linnaeus and Cuvier and as well as the disapproval of all weightings. He, instead, proposed a number of empirical rules on how to discover taxonomically useful characters, that is, how to undertake a posteriori weighting. They include the constant presence of the character in related forms, “‘especially those having different habits of life,” and particularly a constant association of several characters. Ernst Haeckel (1834—1919) introduced the method of repre- senting phylogeny by means of trees or branching diagrams. This method was useful and stimulating and provided the taxonomist with a graphic means for expressing supposed relationship. The search for facts to improve the designs of phylogenetic trees dominated biology during the second half of the 19th century and led to a boom in the fields of comparative systematics, com- parative morphology and comparative embryology. Particularly in taxonomy, it spurred the search for ‘missing links” for example Amphioxus, Peripatus and Archeopteryx etc. and ‘primitive ances- tors.” These efforts led toa far-reaching understanding of the animal forms and to the establishment of a Natural system that is still considered essentially valid. 1-65. FIFTH PERIOD—Popalation Systematics : This period is characterized by the study of evolution within species instead the study of evolution of the higher categories, The typological ‘concept of species, which was already shaky in the preceding period, was totally abandoned and replaced by a dyna- mic, polytypic concept. Species were no longer considered as something fixed and uniform, but rather as polytypic, consisting10 Antmal Taxonomy and Museology of many sub-species and local population, each differing from the others and each showing considerable variability within itself. Two re important :— : “ t s between sub-speciés and species were 1, that the difference: cies the total of very numerous small variations, and 2, that much of the local and geographical variations were closely correlated with the environment. Mistakenly the second point was interpreted by some pro- Lamarckians as indicating a direct effect of the environment on species. However, the early Mendelians e.g. De Vries and Bateson emphasized the role of large mutations and thought that they pro- duced new species by a.single step. They minimized the role of the environment, even as a selective agent. The geneticists began to appreciate extreme importance of small and very small genetic changes and the concept of mutation was extended to include these. Fisher’s (1930) demonstration that even a very amall selective advantage of a new gene or gene combination would cause in due time a genetic transformation of populations wes an important contribution. Goldschmidt, Sumner and Dobzhansky combined taxonomy and genetics together and proposed a new prosperous branch of science the ‘Population genetics". This thinking of geneticists forced taxonomists to give up their Lamarckian thinking and made them realize that the small variations which they had known so long were actually small mutations. J. S. Huxley (1940) labeled this change in thinking from typo- logical to population as New Systematics. . According to this the purely morphological definition of species has been replaced by a biological definition which takes ecological, geographical, geneti- cal and other factors into consideration. The population repre- sented by an adequate sample, the series of museum specimens, has become the basic taxonomic unit. Following are other important events of this period : 1. In 1889 first International Zoological Congress adopted the Balanchard code. 2. In 1900 the Mendel’s work was rediscovered. 3. In 1904 International Commission on Zoological Nomen- clature (ICZN) was formed. 4. Gone Powers were granted:to International ‘ological Nomenclature, andTaxonomy—Definitions, Terms and History iu 5. In 1944 “Systematics and the origin of species” book was presented by E. Mayr. According to this book taxono- mists must study the processes of speciation rather than just its products. 1-66. SIXTH PERIOD—Current Trends : It is characterised by the following three important develop- ments :— 1. Renewed examination of the whole theory of taxonomy (Simpson, 1961 ; Gunther, 1962; Mayr, 1965). 2. The use of electronic computers and associated endeavors to revive a nominalistic approach of taxonomy (Sokal and Sneath, 1963). 3. The vigorous introduction of biochemical techniques and a growing realization among molecular biologists of the importance of understanding the phylogeny of organisms as a basis for the understanding of evolution of macro- molecules. Comparative ethology i.e. science of character formation or science of animal behaviour likewise has stimulated a deeper interest in taxonomy. 1-7. How Taxonomist Works : Practicing taxonomy usually means constructing classifications of particular group of organisms and also identifying specimens in accordance with a classification already made. Back of this practice and of still deeper importance are the theories, principles, rules, and procedures that are, strictly speaking, the science of taxonomy. Formulating and testing those theories, principles and so on are, of course, the most basic things a taxonomist does. The next question is, by what taxonomic principles and procedures an actual classification is to be produced. When, as he certainly should, a taxonomist is going to practice classification, the techno- logical application of his science, just how does he go about it ? In general, he does the following things, more or less in this sequence but usually with a good deal of overlapping, jumping and back-tracking :— 1. He selects or obtains the organisms (by some specific techniques mentioned in chapters 10 and 11) to be classified. He may study them in the field and leave them there alive, but usually they are collected as museum specimens, preferably but not neces- sarily by the taxonomist who later classifies them. For large| 12 Animal Taxonomy and Museoligy groups, at least, he will almost always Heed to use older Collec. tions, perhaps scattered in different institutions. . 2. He observes and records data about the Organisms, ang also assembles from the. literature as nearly as possible all the data already recorded about the particular groups he is ‘Studying. This activity overlaps the first, because some data, minimally those of locality and (for fossil) horizon, can only be observed in the field, and other, or in exceptional cases, all the data may be obtained in the field. Fi 3. He sorts the organisms into taxonomic units, demes, sub. species or species as may be appropriate. He then analyses the data on polymorphic forms and all kinds of variations. The analy. sis is statistical in Principle and now is generally also so in details of practice, inferring Population characteristics from the sample in hand. This activity overlaps the two Preceding and also the two following activities. 4. He makes comparisons among the characteristics of the varying units, with special attention to the kinds resemblances, differences and sequences shown. T! too, are largely statistical in Princfple. 5. He interprets the relationships revealed by comparisons in terms of basic taxonomic concepts, especially homology, para- llelism, convergence, Primitiveness and specialization. 6. On the interpretations made in the last Procedure, he bases inferences as to the evolutionary pattern among the popula- tions studied. He may draw an actual diagram, preferably an inferred phylogenetic tree or a dendrogram intended to be topo- logically as nearly as possible equivalent to a Pologenetic tree. He may use some other visualization or may proceed in wholly verbal form, In any event, he reaches conclusions as to affinities such as are represented by a tree. His conclusions will also involve some considerations, as of degree of divergence, that can not readily be shown by a treo diagram. 7. He translates his conclusions on affinities, So on into hierarchic terms, assembling and dividi; groups of organisms into taxa of various ranks, 8. According to rules and usages, publications names applicable to the recog there are no applicable published names h divergence and ing the various he selects from various nized taxa, and when © coins new ones,