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An Approach to the Problem of Taxonomy and Classification in the Study of Sporae Dispersae

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An Approach to the Problem of Taxonomy and Classification in the Study of Sporae Dispersae AN APPROACH TO THE PROBLEM OF TAXONOMY AND CLASSIFICATION IN THE STUDY OF SPORAE DISPERSAE D. C. BHARDWAJ Birbal~i Institute of Palaeobotany, Lucknow ABSTRACT ( 1950 ) system, the American workers prefer The present state of disagreement among spore that of Schopf, Wilson and Bentall's ( 1944), workers with regard to the taxonomy and classi• in Germany, an amplification of Ibrahim's fication of Sporae dispersae has been reviewed. ( 1933) system is mostly in vogue and the The taxonomic categories for Sporae dispersae have Russian workers classify according to Nau• been defined and practicability of organ-genus concept over the form-genus concept for the circum• mova's (1937) system. This tendency to scription of spore taxa has been discussed. stick to one or the other of these systems, none of which excels the others, has uncons• ciously led to thwart the evolution of one such INTRODUCTION system of classification for the Sporae dis• persae which may be universally followed. Of late, in view of the greater applied use of DETAILEDand pollen,studyrecoveredof the dispersedfrom sedimen•spores Sporae dispersae in coal and oil prospecting, tary strata, dates back to Reinsch the need for a universally accepted standard ( 1881) when he described some of them from system of classification for these is being coals of the Carboniferous age. Thereafter, increasingly felt so that assimilation of data for several decades, these dispersed micro• from all over the world can be more easily remains attracted occasional attention of achieved. palaeobotanists and petrologists. However, it is only since the early thirties that the THE PROBLEM British, German, Russian, American and Indian palaeobotanists as well as geologists, All the systems of classification proposed realizing the importance of the Sporae dis• in the past, in spite of their diversities, tend persae in geological dating and stratigraphical to fall into two groups depending upon the correlation of sedimentary strata, set them• differences in their taxonomic treatment of selves to a serious study of these plant micro• spore taxa: remains. This increased research activity 1. Artificial ( morphologic) systems which has resulted into very significant and substan• consider the morphologic characters of tial contributions on the study of Sporae spores and pollen as the basis of specific, dispersae of various ages and from many parts generic and suprageneric grouping of the world. (BENNIE & KIDSTON,1886; IBRAHIM, Prompted by a desire for exactitude and 1933; NAUMOVA,1937; ERDTMAN,1947; clarity in description of the Sporae dispersae, KNOX,1950; PANT,1954and POTONIE& spore workers have always adopted some KREMP,1954). system of taxonomy and classification to 2. Natural (phylogenetic) systems which present their results. A number of these emphasize circumscription of spore systems are a result of original thought but genera on the analogy of higher taxa others are a sequel to synthesis of the then ( Orders-Classes) of the natural system prevalent older conceptions. Each of these of classification of plants ( SCHOPF,1938; efforts on systematics of Sporae dispersae has SCHOPF,WILSON& BENTALL,1944). its merits and also shortcomings. However, The real problem among palynologists in spite of the latter these efforts are highly today is to decide about the suitability of commendable inasmuch as they have been any of the two approaches between which able to single out the numerous difficulties most of the opinions tend to segregate. besetting the evolution of an ideal system. However, to choose between the two is not From a perusal of the current literature on so easy unless we discuss the usefulness of Sporae dispersae it becomes evident that the one over the other in the light of scientific as British spore workers mostly follow Knox's well as practical needs and the limitations 3 4 THE PALAEOBOTANIST imposed by the nature of the material hand• Study of Sporae dispersae reveals these floral• led. breaks readily. It is for the detection of such ASPECTS IN THE STUDY OF SPORAE floral-breaks and for fine stratigraphic cor• DISPERSAE relation of coal seams as well as other strata, that this branch of palaeobotany is proving The study of Sporae dispersae has twofold to be of great practical value. applications, i.e. botanical and geological. From the studies of Mason (1936) and The botanical aspect concerns primarily, with others it is now well known that floral changes the taxonomy and phylogeny of spores and are caused by the evolution or migration of pollen and secondarily with ecology in so far such constituent species which, in view of as these plant microremains are representa• their specialized needs of the environmental tives of the flora of that area. The geologi• factors, behave as the borderline species in cal aspect of stratigraphical correlation and the population. Thus to detect the floral dating of strata depends upon the applied changes in a fossil flora it is imperative that use of the botanical knowledge so gained, the fossils of the borderline species should be especially that which concerns the changes distinguished from the other species. The in the distribution of Sporae dispersae or same holds good in the study of Spora; dis• in other words their parent floras, during persae and implies a taxonomic treatment geological time. which aims at distinguishing the spores be• Speaking of the botanical aspects of taxo• longing to each species and genus of the nomy and phylogeny, Sporae dispersae have parent flora. Hence it can now be concluded limited value in view of their dispersed nature, that the primary need in the scientific study because nothing definite can be stated about of Sporae dispersae is to define the taxonomic the identity of their parent plants. As a principles which may enable circumscription. matter of fact, whatever we know today about of the small, spore taxa as nearly equivalent the phylogeny of some of the dispersed spores to natural plant species and genera as possible. has been a contribution from botany or palaeobotany rather than vice versa and it is TAXONOMIC CATEGORIES FOR SPORAE doubtful if an independent study of Sporae DISPERSAE dispersae would ever contribute anything Spore-Species more tangible than probabilities. However, for ecological studies, Sporae dispersae have A scientific definition of a plant speCIes proved to be of real help especially in Tertiary could be that it is a group of morphologi• ecology where Pollen-analysis, in conjunction cally and physiologically similar individuals with the study of plant megafossils, has which are biologically isolated from those always enhanced accuracy of data and there• of the other closely related species. Among by its interpretation. This success of pollen Sporae dispersae, in view of their isolated studies of Tertiary strata is in a large mea• state, a fossil spore-species can be defined sure due to the possibility of correctly identi• only as a group of morphologically ( i.e. mor• fying the parent genera or even species of the phographically) similar individuals. The dispersed spores and pollen. other features such as physiological similarity The application of the study of Sporae dis• and biological isolation are beyond any persae to stratigraphical (geological) prob• means to establish in fossils. Thus, in effect, lems especially of the pre-Tertiary strata the only important basis of defining spore• depends on the detection of floral changes species is the morphography of spore fea• which have occurred in an area during the tures as these are actually evidences of course of time due to migration and evolution evolutionary morphology. of the species constituting its vegetation. Theoretically, a species cannot be strictly Floral changes are normally imperceptibly genetically homogeneous population because gradual, but occasionally they are accentuat• no individual can contain all the hereditary ed due to biotic or environmental factors and, factors which exist in allelic pairs and series. accompanied with migration of species from But we know that in nature, barring excep• and to the area, cause significant change in tions, mostly one set of either of the pair of the composition of succeeding floras, thus allelic characters tend to be represented in a interpolating floral-breaks in the vegetational species. The more usual characters must, history of the area. These floral-breaks are therefore, be accepted as the characters to used for delimiting stratigraphic horizons. define a spore-species. BHARDWAJ - TAXOXOJI,IY AND CLASSIFICATION OF SPORAE DISPERSAE 5 Variation within a Spore-species - The well. The size is a valuable variable in characters tend to vary, within a species some miospores of Lycopsida but ofless consequence varying more but others less. From my for their megaspores, of indeterminate utility work ( BHARDWAJ, 1952) on the spores of the in spores of Sphenopsida and of problematic living genus Anthoceros ( L.) Prosk., the work use in the taxonomy of gymnospermous of Knox ( 1950) on the spores of Lycopo• pollen grains. dium, Phytloglossum, Selaginetla and Isoetes, In other features, e.g. height and width Stokey & Atkinson (1956) on Osmunda• of emergences as well as that of rays, the ceae and Wagner ( 1952) on Diellia, I have range of quantitative variation in a species been able to infer that in Cryptogams the is within definable limits. For delimitation morphographic characters of mature spores, of species within a genus it is important within a natural species, vary quantitatively to work out the range of variation and but the range of variation is within definable the statistical mean of each variable, to limits. Likewise, as far as known to me, enable comparison with that of the other this generalization holds good in the case of comparable species. Unless as pore worker pollen grains of Phaneogams as well. has a rational conception of these varia• It is worth pointing out here that while tions, he is very likely to split or lump writing these generalized observations, I am together natural species.
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