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CYTOTAXONOMY in the GENUS CERASTIUM L. a Thesis Presented

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CYTOTAXONOMY in the GENUS CERASTIUM L. a Thesis Presented CYTOTAXONOMY IN THE GENUS CERASTIUM L. A thesis presented in part fulfilment of the requirements for the degree of Doctor of Philosophy in the Faculty of Science in the University of London by NASIR AHMED BAIL M.Sc. Department of Botany, Imperial College of Science & Technology, London, S.W.7. January, 1965 ABSTRACT The genus Cerastium has always presented difficulties to taxonomists. The present work represents an attempt to use cytological, statistical and experimental methods to investigate one of the most confused parts of this genus. On Mount Maiella, Italy, were found a range of taxa which appeared to be an inter- grading series ranging from C.laricifolium taxa to C.album taxa. These were collected together with seeds and grown under similar conditions at the Imperial College Field Station. It was found that with the exception of general greater luxuriance, all the taxa showed no significant differences from their growth in their original habitat. All seeds showed high degree of fertility ( usually c. 90c%). These taxa have well defined ecological niches in nature and it can be concluded from these experiments that these taxa form a range of forms, genetically adapted to particular ecological conditions and not exhibiting any significant phenotypic variability. Measurements of 15 variables of some 600 specimens were. made. The computation of 'canonical variates' was carried out on sirius computer. This biometric treatment could estimate the objectivity of the subjectively determined taxa of the group A (containing 9 taxa) and group B (containing 15 taxa). For the group A the total variability was 82520.79. The first five components accounted for 97.7% of total variability. For the group B the total variability was 181474.48. The first six components accounted for 99.86%of the total variability. This indicates that variables carry differential weighting. This biometric treatment separated the group A in three sub-groups containing three taxa each. These sub-groups fall within the limits of the species C.album. Group B waa divided into four sub-groups, each of which falls within the limit of separate species (C.laricifolium, C. rigoi, C.thomasii and C.matrense). The taxonomic descriptiOn of the taxa was given. Somatic chromosome numbers of 21 taxa were determined. Cytologically the taxa showed many degrees of polyploidy 2n number ranging from 36 to 126. The basic number for the genus was suggested as 9 and not 9, 17 and 19. The importance of polyploidy wak discussed. CONTENTS Page ABSTRACT 2 I INTRODUCTION 6 a)Cytotaxonomy 11 b) Numerical taxonomy 12 II MATERIAL AND METHODS 17 Measurements 17 Sowing 22 Cuttings 23 Transplantation 2L1. Cytology Technique 29 III TAXONOMIC DESCRIPTION 34 IV CYTOLOGY 76 Review of literature 76 Cytological observations 89 Page V BIOMETRICS 924 Nomenclature 138a VI DISCUSSION 139 Cytological discussion 155 Basic chromosome number in the geAus Cerastium 155 Polyploidy 159 VII ACKNOWLEDGEMENTS 167 VIII BIBLIOGRAPHY 169 IX APPENDIX 178 5 CYTOTAXONOMY IN THE GENUS CERASTIUM L. 6 Introduction The history of taxonomy reveals that there have always been certain groups whose taxo- nomy has presented more difficulty to the taxonomists than others. These groups have been called "Critical groups" and much controversy has arisen regarding their treatment. These groups may be defined as groups where the variation of morphological charac- ters overlap to such an extent that there are no complete correlations giving a definite bimodality of the distribution of variation. There has been confusion about the descriptions of such groups especially where there has been a practice of naming intraspecific groups on the basis of the dif- ferences in one minor character only, and the accuracy of such classification is difficult to test or assess. These difficulties have suggested the need to adopt appropriate biometric and statistical methods which can give a means of assessing the objective validity of any grouping and which can easily be tested or verified by other workers in this field. The genus Cerastium has always presented difficulties to the taxonomists as the included species show a great deal of variation. In some 7 annual species this variation is mainly phenotypic due to eniironment (Whitehead, 1956). There is also a high degree of genetic variability and con- siderable overlapping of the range of characters (Whitehead, 1955). Most work on this genus has consisted of a series of "reshufflings" often based on dubious characters. At the same time many collections have been identified from descriptions from floras without comparison from correctly named herbarium material. Even if the taxa can clearly be distinguished their nomenclature presents a difficult problem. The present investigation is an attempt to deal with some of the confused parts of this genus by cytological, statistical and experi- 1 mental methods. There follfts a brief chrono- logical, account of the systematic treatment of the genus Cerastiurl ';hick shows how far the various authors intended to reveal the natural relationships in this genus. Linnaeus Linnaeus's system was an artificial one based, exclusively on floral characters, the stamens being given a preponderance of attention. His twenty four classes were based on the number or 8 some other obvious characteristic of the stamens. In his system, Cerastium was placed in the class of ten stamens, the Decandria, and in the order of five styles, the Pentagynia. This artificial system prevented him from realising that species which had different number of styles and stamens sometimes came under the same genus. He placed Co cerastoides (L.) Britton in his order trigynia because it usually has three styles, and assigned it to the genus Stellaria. Linnaeus described fourteen species of Cerastium in his "Species Plantarum" (1753), which he arranged in two groups according to the shape of the capsule. De Candolle's classification was not an artificial one like Linnaeus's. He made an attempt to formulate a natural system, by which he meant one which would reveal the Divine Plan, accepting the doctrine of Special Creatian. De Candolle believed strongly that morphology alone provided a key to taxonomy. His classification of the genus Cerastium is therefore based entirely upon morphological cha- racters, although when De Candolle published his Prodromus Systematis Naturalis Regni Vegetabilis in 1824, he gave a much more extensive treatment 9 describing in all sixty-nine species and divided the gent's into sections according to the capsule teeth. Species with dentibus circinnatis were included in the section Strephodon and the species with dentibus margine revolutis were placed in the section Orthodon. He further divided the section Orthodon into two subsections, one with rather small flowers having petals and sepals of equal size, and the other with large flowers having petals larger than the sepals. Endlicher (1836-40) in his genera Plantarum divided the genus into five sections. His first section, Dichodon, contained plants with three styles. His other two sections were the Strephodon and Orthodon of De Candolle, and two others, Schizodon and Monchia. Fenzel in his classification divided the genus Cerastium into four subgenera; 1. Dichodon, 2, St:,ephedon, 3. Schizod-n and 4. Orthodon, He further divided the subgenus Strephodon into T,pAeyetala (Petals glabrous at base) and Ciliatopetala (hair at the base of Petals). The subgenus Orthodon 0-0 was dividedCugpoin (annual species) and Perennia (perennial species), The section. Fugacia was further divided into Ciliatopetala and Leiopetala. 0 The classification of Boissier (1867) was more or less similar to Fenzel's except that C. dichotomum was pTaced in the section Orthodon and not in a separate group. Nyman (1878) in his classification simply divided the genus into four sections, of which Strephodon Orthodon and Dichodon, were already well established, the fourth he call Cryptodon. Englar and Pranti in their work 'Die Naturlichen Pflanzenfamilien" published the volume dealing with the family Caryqphyllaceae in 1889. Pax described the family. Dealing with the genus Cerastium Pax divided the genus first into Dichodon and Eucerastium. T-Tr then subdivided Eucnrastium into Strephodon, Orthodon and Cryptodon. Moschl (1936-38) working in the European. Orthodon species in the section Leiop_ttala of Fenzel concluded that these species Duld be naturally placed into Ovoglcndulosa• -,wn•to r (speies wi-Th the terminal cells of the glandular hair ovoid) and Olavatmlandulosa (species with the terminal cells of the glandular hair club shaped). He thought that possibly the whole genus perastilim could be divided in this way. 11 Cytotaxonomy Cytology is a study of nuclear phenomena, and pLrticularly of chromosomes. It has become of great importance to taxonomy. Cytotaxonomy has become a useful tool for modern evolutionary classification of plants, and often makes possible the investigatim of relation- ship and barriers between taxa at or above the species level. This approach studies the chromo- somes in all their aspects, in natural populations as well as in experimental offspring, but the most C=MCD and basic phase is concerned mainly with the study of the number and morphology of chromosomes. The cytological studies of plants began more than half a century ago, and earlier result revealed that considerable variation existed in the number and morphology of the chromosomes between any two species and within the same species. With improvements in the cytological technique, examina- tion of chromosome complements has become now almost a routine operation especially in the investigation cf critical taxonomic groups. The chromosome set is seen at its best at the mitotic metaphase which is known as the Karyctype. The most easily observable 12 Karyotypic differences are those involving chromo- some number and the phenomenon of polyploidy. Apart from the chromosome number the karyotypes can be useful in groups where the chromosomes are large enough for good morphological observations. The taxonomic value of cytological data varies from group to group.
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