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. In some groups, practically it has had not significance at either the species level or above (i.e. where the karyotype is of no or little taxonomic value and has same 2n chromosome number). In some other groups the number and morphology of the chromosomes can be investigated and may have great taxonomic value. Numerical Taxonomy The historical development of taxonomy has produced alpha, beta and gama phases; alpha phase was limited to the production of conventional description of species or groups of species; beta phase was concerned with schemes of natural classi- fication of all levels in the taxonomic hierarchy; and the gama phase was concerned with the problems of evolutionary relationships. Today classification tends to aim at grouping like with like without neces- sarily making any evolutionary of phylogenetic impli- cation. Numerical taxonomy is beginning to make 13

important contributions to these studies. The aim of the numerical taxonomy is to develop methods by means of which different scientists working independently will arrive at somewhat identical estimates, and these methods may lead to the stable classifications which will not need extensive revision as new knowledge be- comes available. This wide use of quantitative measures of relationship will increase the ac- curacy and precision of taxonomy. ,wSokal and Sneathm' (1963) have defined numerical taxonomy as "It is the numerical,evaluctionof the affinity or similarity between taxonomic units and the ordering of these units into taxa on the basis of their affinities" and the outstanding aims of numerical taxonomy are repeatability and objectivity" The idea of a natural classification as a phenetic one based on affinity is not new. It was first introduced by a French botanist Adnanson (1763). He was the first taxonomist who conceived of the use of every feature impartially and with equal weight, he insisted that every part of the plant should be used in making a classification. 14

The viewpoint of Gilmour (1940) is close to that of Adansonian that the ideal classification is one with the greatest predictive value. Such a concept is based on giving every feature equal weight. Gilmour (1961b) in discussing the mathe- matical approach to taxonomy notes that it may help to decide between disputed classifications. The reason for the great usefulness of natural classification is that when the members of a group share many correlated attributes, the"implied in- formation" or "content of information" (Sneath, 1957a) is high; this amounts to Gilmour's saying that a system of classification is the more natural the more propositions there are that can be made regarding its constituent classes. Remaine (1956, P. 4) indicates that the predictive value of taxon- omic groups is only true of natural taxa, not of artificial ones. It is obvious that artificial groups established on a single character are of low predictive value. A natural classification can be used for a great variety of purposes, while an arti- ficial one serves only the limited purpose for which it was constructed. During the past few years these ideas have 15

been further developed and have been summarised by Sneath (1958) and modified by Sokal and Sneath (1963) as follows: 1. The ideal taxonomy is that in which the taxa have the greatest content of information and which is based on as many characters as possible. 2. Every character is of equal weight in creating natural taxa. 3. Over-all similarity (or affinity) between any two entities is a function of the similarity of the many characters in which they are being com- pared. 4. Distinct taxa can be constructed because of diverse character correlations in the groups under study. 5. Taxonomy is, therefore, strictly empirical science. 6. Affinity is estimated independently of phylogenetic considerations (for detail see Sokal and Sneath, 1963). At present the numerical taxonomy is very useful in the middle and lower ranks, in the study of genera, species and perhaps of families. Its aim is to make phenetically stable groups, and tries 16

to overcome some of the faults found in conventional taxonomic groups. Numerical taxonomy is becoming a very rapidly growing field. It has become practical for taxonomists to use numerical taxonomic techniques today. A considerable body of work has already been done with quite satisfactory results. The calculations require electronic computers and these are increasingly available and processing of the data is increasingly fast. The difficulty today may be the collecting and study of the material and the codification of the characters, not the analysis and classification. The proponents of numerical taxonomy insist on the separation of the taxonomic process (based on affinity) from phylogenetic speculations for the following reasons. 1 Phylogenetic speculation is not competable with the stated aims of objectivity and repeatability 2 Fossil record is very fragmentary and the phylogeny of most of the taxa is unknown. 17

MATERIALSAND METHODS On Mount Maiella, central Italian Apennines, were found a range of Cerastium taxa. Twenty-four taxa were collected together with seeds. These taxa fall within the limit of five species. Below are given the names of five species, together with their respective taxa used in the measurements of characters and cytological observations. 1. C. laricifolium Viii. includes taxa A, B, C, D and I.

2. C. thomasii Ten. includes taxa• E, F, G and H. 3. C. rigoi Huter et Porta includes taxa J, K and L. L. C. matrense Kit. includes taxa M, N and 0.

5. C. album Presl includes taxa P, Q, R, S, T, U, V, W and X. Measurements Almost all the material collected from Mount Maiella used in measurement was dry and not very well-mounted on herbarium sheets. Boiling of the material in water was found necessary. In most of the speciemena the petals were broken, torn out, and shrivelled as such the petal character had to be eliminated. 18

The fifteen characters chosen were: a - Sepals length average b - Sepals breadth average c - Scarious tip left, average of large sepals d - Scarious tip left, average of small sepals e - Scarious tip right, average of large sepals f - Scarious tip right, average of small sepals g - Scarious tip height average of sepals h - Bracts length average i - Bracts breadth average j - Bracts scarious tip height average k - Seeds length average 1 - Seeds breadth average m - Pollen grains diameter average n - Leaves length average o - Leaves breadth average. In the measurements of the characters a, b, c, d, e, f, g, h, i, j, k, 1, n and o One ocular division = 734

In the measurements of the character m One ocular division = 3.73t1 19 These were measured as follows: A. Sepal Sepal length was measured from the inner point of attachment on the receptaole to the end of scarious tip, not including hair bases,, Sepal breadth was measured as being the greatest distance at right angles to the length. In measuring breadth, sepal was cut beneath the greatest distance at right angles to the length to facilitate the spreading of the sepal espedially of the scarious tissue fully, by pressing it with needle at right angles to the length. Sepal scarious tip height was measured from the point, at which, in the centre of the sepal, the cells ceased to contain chlorophyll to the extreme point of the sepal, not including hair bases. The scarious tissue breadth of the sepal both larger and smaller both on right, and left hand sides respectively was measured from the points (both right and left on sepal breadth) at which the cells ceased to contain chlorophyll to the extreme points on both sides of the sepal breadth. (see fig.3 Pg•36) B. Bracts Mostly the bract was measured as being the topmost bracteoli subtending a developed bud or flower. 20

The length was taken from the inner point of attach- ment on the stem to the extreme point not including hair bases. The breadth vas taken as being the greatest distance at right angles to the length. The Scarious tip was measured from the point at which, in the centre of the bract, the cells ceased to con- tain chlorophyll, to the extreme point of the bract not including hair bases. Seed Length of (unboiled) seed was measured as being the distance between the funicle and the centre of the opposite cord. Breadth was measured as being the greatest distance at right angles to the length. D. Pollen Grains They were found to be more or less spherical so the diameter of the pollen grains was measured. E. Leaves Usually five pairs of older and larger leaves from the same shoot bearing flowers were taken, first pair being the topmost beneath the bracts and the rest at the successive nodes. The length was measured from the inner point of attachment on the stem to the extreme point, excluding hair bases. 21 The breadth was measured as being the greatest dis- tance at right angles to the length.

All the measurements of 15 characters of some 600 specimens were made under the microscope using X 10 power eye piece having graduated disc in 100 divisions and adjustable objective fixed at x 1.8 in all the characters except in the character pollen grain diameter where x 40 high power objective was used. ,Sowing i. Petridish Seeds (of Cerastium taxa) were sown on Whatman filter paper in petridishes and kept in moist condition using a thin layer of cotton wool in the petridish. Whenever sowing was done in summer, cold treatment was given to hasten germination. Germination occurred between 3 - 6 days. Germination percentage was found to be very high among all the Cerastium taxa under -investigation. ii. Seed-pans Seeds were also sown in seed pans. The soil used in sowing was the mixture of clay, sand and peat in equal proportion and appropriate amount of John Innes fertiliser. The seed pan was filled with

22

the soil, leaving an inch of empty space from the rim of the seed pan. The surface level was then made uniform and very shallow holes with equal dis- tances were made in rows on it. A definite number of seeds was,..; placed one by one in each hole, the seeds were then covered with a thin layer of fine sand by means of a fine mesh sieve. After this operation the seed pans were watered very carefully without causing any disturbance to seeds, the seed pans were then placed in shady and cool place to cut the rate of evaporation and to hasten germination. First sign of germination appeared between Li. - 7 days. After first sign of germination the seed pans were put under the greenhouse light. The duration of light was fixed fourteen hours a day. The seeds have been found to remain viable even after two years. Seeds also germinated when they were sown immediately after the collection from Mount Maiella. It suggests that these seeds reOire no speciol treatment to break dormancy. Cuttings Cuttings of the branches of many taxa grown in the greenhouse as well as in the field,were taken and after removing all the leaves,except the upper two rows from the cuttings,were planted in a propagating 23 frame with bottom heat in rows at equal distances one inch apart. After one week it was observed that almost all the cuttings had taken root in the soil. After two weeks the cuttings were ready to be transplanted. No fungus infection was found either among the cuttings or among the seedlings. Practically all the seedlings and cuttings showed healthy growth were not therefore treated with fungicides. Transplantation After the emergence of first pair of leaves the seedlings of the Cerastium taxa were transplanted in 6" diameter pots. The soil used in transplanting pots contained manure and the top was covered with a thin layer of sand. Seedlings were removed one by one from the petridishes and the seed pans (when used) very carefully, taking care not to damage the seedlings even slightly as the seedlings were very small and delicate to handle. Seedlings were transplanted very carefully and gently one in each pot. After transplantation the pots were thoroughly watered. Culture number and plant number was given to each plant. All the seedlings transplanted from the petri-dishes (or seed pans) bore the same culture 24 number and separate plant number. After watering the transplanted plants the pots were kept in cool and dark place to check transpiration. Two days later the pots were placed under the greenhouse light (fixed at fourteen hours a day) buried in the gravels, leaving the rim of each pot above the gravel surface. The vegetative growth was quite vigorous and normal.(See Figs. 1 & 2). During the early summer-time half of the Cerastium taxa population was retransplanted in the bigger pots as the vegetative growth was very vigor- ous and were transferred to a plunge bed and the pots buried leaving only the rim of the pots above the soil surface. Plants grow well in the field as well as in the greenhouse. Cuttings when ready to be transplanted were transplanted in the pots (method mentioned above). No fungus or any other infection was found in the seedlings after transplantation. A few seedlings died after transplantation probably from mechanical injury; and not by any infection. However, the fully grown plants were infected very heavily by reddish brown aphids. Lambers D. Hille Ris (1946) also reported that

25

Fig. 1. Showing the Cerastium taxa grown in the green- house.

26

Fig. 2. Showing the Cerastium taxa grown in the field. 27

Caustium spp. particularly C. tomentosum, C. arvense and C. caespitosum can be infected by aphids (Myzus persicae Sulzer). Aphidt. hibernate eggs on Cerastium spp. and on some other genera of caryo- phyllaceae. It seemed very difficult to get rid of them. By the application of 'pesticide' twice a week continued for about six weeks and the green- house also was smoked twice and ultimately the plants got rid of the aphids and the damage done by the aphids was not considerable. All these sowings and transplantations were done to raise the populations of almost all the taxa and when these taxa start flowering the object was to examine meiosis and to make crosses between the taxa into as many directions as possible within the available time, so as to study the relationship of these taxa cytogenetically in relation to taxonomy. The vegetative growth continued abundantly. However, the plants did not produce flowers. To overcome this difficulty of non-flowering various methods were tried to induce flowering as follows. i)The pots were buried in gravels. ii)Duration of light illumination was increased and 28 kept at 14 hours a day. iii)Plant-pots were kept buried in the plunge bed to be faced with high wind velocity and natural sunlight, (whenever available in summer) as high wind velocity and longer period of light are two of the many factors necessary to induce flowering. iv)Plant pots were also left in the field during winter as low temperature during vegetative growth later encourages summer flowering. We managed to keep the plants growing ab- undantly for about more than two and a half years and still they are growing quite normally and there is no sign of flowering so far. If we are going to have a good summer these plants may flower. Cytology Technique Somatic Chromosome Counts Seeds were collected from Mount Maiella and stored in paper envelopes. Seeds were sown whenever required on Whatman filter-paper in steri- lised petri-dishes and kept in moist conditions using a thin layer of cotton-wool underneath the filter-paper. Germination was observed within 3 - 6 days. Sometimes cold treatment was given to hasten germination. 29 It was found that the most suitable part of the plant to use for mitosis was the first root- tip of the germinating seed. As this has a large root-tip and sqUashee very easily. Young root-tips of fully grown plants were also used for mitosis. Squash method is as follows:- Prefixation:- It is very difficult to study and count the chromosomes from the tissue, without the addition of prefixative because of the clumping of the chromosomes. A saturated aqueous solution of paradichlorobenzene (PDB) was found to be the most suitable prefixative. Whereas colchicine and acenaphthene proved to be of little use. Higher concentration of colchicine is poisonous and harmful to the seedlings and lower concentration has little effect. Acenaphthene was also too drastic in its effect and caused the chromosomes to bunch together in light clusters. As tl-e chromosome number of the taxa under investigation was found to be very high, it seemed most essential to put the material when it is in prefixative, in the mechanical shaker for shak- ing at moderate speed, which helped in breaking-of the spindle and setting free the chromosomes in the cell completely, thereby making the counting of the chromosomes easy and accurate. 30

The root-tips were thoroughly washea in distilled water and prefixed in PDB and clamped to the mechanical shaker for two hours. After pre- fixation the root-tips were thoroughly washed in distilled water, following several changes so that no trace of prefixative remained in the root-tips. Fixation The root-tips were fixed in acetic alcohol (mixed in 1 : 3 proportion) and placed in the mech- anical shaker for one hour for thorough fixation. The root-tips were then passed through one change in absolute alcohol for an hour. If not hydrolysed immediately the root-tips were stored in 70% alcohol for a period of 2 - 3 days. Stored material in 70 per cent alcohol for a week or so did not give good results. Hydrolysis Root-tips were thoroughly washed in dis- tilled water for several changes. Transferred to specimen tubes containing normal Hcl. Hydrolysed for 20 - 25 minutes in incubator at a temperature of 58°C - 63°C. Staining Normal Hcl was drained off completely and 31 a little leucofuchsin was added. In 20 - 25 minutes time the root-tips were found to be brilliant red. Most previous workers have used gentian violet as a nuclear stain but it is a very lengthy process and the results are not so good. Hagerup (1941) used Feulgen for C. subtetrandrum. Squashing The stained portion of the root-tip was cut with sharp blade or scalpel and placed in a drop of 45 per cent acetic acid, or in acetocarmine on a new clean slide. It was then teased with the help of an ordinary needle or a spear-headed needle. Cover glass was then mounted over the teased material in such a way as to leave the teased material in the centre of the cover slip. Tapping gently and little pressing of cover slip to spread the tissue uniformly was done. The slide was passed over a spirit lamp for a few seconds for making the chromosomes more clear. It was then ready for observation under the microscope and for photomicrography. 32

Method for making the slide permanent The slides were made permanent following the schedule of Darlington and Lacour (1945) as given below:- a) the slides were inverted in the smearing dish containing 10 per cent acetic acid for the separa- tion of cover slip from the slide. b) both the slide and cover slip were placed in freshly prepared acetic alcohol of 1 : 3 ratio for two minutes; followed by one change in 8.0 per cent alcohol. Finally passed through two changes in absolute alcohol for one to two minutes in each. The cover slip was remounted in a drop of euparol on the same position of the slide. The permanent slide was then placed in oven for twenty- four hours. As the material under investigation was the mountainous species and subspecies of the genus Cerastium mitosis appeared to be very difficult. It took a pretty long time to find out the right tide at which the cell division in root tip was most active. Active cell division occurred mostly between 10.30 a.m. to 11.30 a.m. It was found after several 33 attempts that the time required for perfect hydrolysis was between 2G - 25 minutes at a temperature ranging between 58°C - 63°C. and the root-tips took very brilliant red colour and no crystals were seen under the microscope. 34 TAXONOMIC DESCRIPTION

A range of Cerastium taxa found growing on Mount Maiella, Central Italy, appear to be an in- tergrading series ranging from distinct C. laricifolium taxa, (C. thomasii taxa), through intermediate taxa to distinct C. album taxa. These plants were collected together with seeds and divided into 24 taxa (referred to as A to X. The taxa P to X are included in group A and the taxa A to 0 represent the group B) by eye and on the evidence of the measurements of 15 variables (characters) used in the present investigation. Cytological studies show that these taxa represent high degrees of polyploidy with 2n chromo- some number ranging from 2n = 36 to 2n = 126. C. laricifolium Vill. consists of taxa A, B, C, D and I with 2n = 36. These are found in crevices often fairly sheltered in the cliffs at the edge of the altipiano. C. thomasii Ten. consists of taxa E, F, G and H. These are found in exposed parts of the altipiano and at the edges of the cliff tops. 2n chromosome number ranges from 2n = 36 to 2n = 54. 35

C. album Presl includes taxon T with 2n = 36, Taxon u, 2n = 63 and taxa P, Q, R, S, V, W and X, 2n = 72. These are found in large groups growing from just below the shallow cliffs at the edge of the altipiano down to the lower slopes of the mountain, an altitudinal range of some 7000 ft. They are never found on the altipiano proper and only in sheltered habitats at the higher altitudes. C. rigoi Huter et Porta includes taxa J, K and L with 2n chromosome number ranging from 2n = 36 to 2n = 126. These are found at the base of the altipiano cliffs. C. matrense Kit. includes Taxa M, N and 0. These are found below the cliffs of the alti- piano and extend a little way down into the valley. 2n chromosome number ranges from 2n = 72 to 2n = 108. A brief description of Cerastium L. and each of the 24 perennial taxa (by eye) of the group B and A based on the measurements of 15 variables (characters) used, is given below. The characters used in the measurements of the scariousness of the sepals are shown in Fig. 3, page 36.

36

SCO,r 04.

wi dth S moll scar kl tcp Y ,'51rtt

%-ci. %Ile. SC 0..r 10 US 1-4y le. SCcLY u,s ti? trp test

Svr%oI Sccoloti tip u,8_t

WT-cam sko the c hoTo..c.ters used. the yr, eQsktr errients

of tile SCcUr ousrtess o.f the sepal.

Fig. 3 37 Cerastium L. The description of the genus as per Clapham, Tutin and Warburg (Flora of the British Isles, IInd Ed. 1962, p.233) is as follows:- "Annual to perennial herbs or dwarf shrubs, usually hairy, with opposite, entire, sessile leaves. Flowers usually in cymose inflorescences, sometimes solitary; sepals free, with membranous margins; petals white, bifid up to half-way or emarginate, sometimes 0; stamens usually 8 or 10, sometimes 5 or fewer; nec- taries present; ovary one celled; styles usually 5, opposite to sepals, sometimes 4, 3 or 6. Fruit an oblong capsule exceeding the sepals, usually ± curved, opening by twice as many short teeth as styles. Seeds numerous, spherical or reniform, rough. Per- haps c.100 spp., cosmopolitan, but principally in the north temperate regions of the "Old World".

Taxon A. ± Caespitose•, perennial, with flowering shoots, up to 6 cms. and shorter sterile ones, all shoots hairy. Leaves 4.34 - 5.78 x 1.37 - 1.86 mm., linear-lanceolate to lanceolate; bracts 2.92 - 4.35 x 1.42 - 2.19 mm., with a scarious tip 0.29 - 0.91 mm. wide, hairy, lanceolate. Sepals 4.75 - 6.13 x 1.82 - 2.54 mm., large scarious tip left 0.51 - 0.87 mm., small scarious tip left 0.21 - 0.54 mm.,

38 large scarious tip right 0.51 - 0.80 mm., small scarious tip right 0.16 - 0.54m.scarious tip 0.27 - 0.94 mm; wide, hairy. Seds 1.02 - 1.28 x 0.76 - 0.97 mm. more or less spherical to ellipsoidal. Pollen grains 33.9 - 40.2µ in diameter, circular. Chromosome number: 2n = 36.

Taxon B. ± Caespitose; perennial, with flowering shoots up to 9 cms. and shorter sterile ones, all shoots hairy. Leaves 5.22 - 7.15 x 1.20 - 1.68mm, hairy, linear-lanceolate; bracts 3.30 - 4.24 x 1.53 - 2.15 mm., with a scarious tip 0.43 - 0.98 mm. wide, hairy, lanceolate. Sepals 5.18 - 6.64 x 1.97 2.49m1l.arge scarious tip left 0.58 - 0.84 mm., small scarious tip left 0.21 - 0.58 mm., large scarious tip right 0.51 - 0.80 mm., small scarious tip right 0.18 - 0.60 mm., scarious tip 0.45 - 1.08 mm. wide, hairy. Seeds 0.91 - 1.24 x 0.69 - 0.96 mm. more or less spherical to ellipsoidal. Pollen grains 33.5 - 44.7µ in diameter, circular. Chromosome number: 2n = 36.

Taxon C. Caespitose, perennial, with flowering shoots up to 7 cms. and shorter sterile ones, all shoots hairy. Leaves 4.02 - 6.00 x 1.13 - 1.88=1.;

39

Fig. 4. Showing the plant and the mitotic metaphase of the taxon A. 2n chromosome number = 36. Plant magnification -= x.9. Chromosome magnification = x 1525.

Fig. 5. Showing the plant and the mitotic metaphase of the taxon B. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525.

L1

Fig. 6. Showing the plant and the mitotic metaphase of the taxon C. 2n chromosome number = 36. Plant maginfication = x.9. Chromosome magnification = x1525. 42 hairy, linear-lanceolate to lanceolate; bracts 2.46 - 3.83 x 1.56 - 2.09 mm., with a scarious tip 0.16 - 0.87 mm. wide, hairy, lanceolate. Sepals 4.40 - 5.94 x 1.82 - 2.89 mm., large scarious tip left 0.43 - 1.02 mm., small scarious tip left 0.14 - 0.62 mm., large scarious tip right 0.43 - 1.02 mm., small scarious tip right 0.14 - 0.55 mm., scarious tip 0.- 36 - 0.94 mm. wide, hairy. Seeds 0.83 1.21 x 0.65 - 0.94 mm., more or less spherical to ellipsoidal. Pollen grains 31.7 - 42.54 in diameter, circular. Chromosome number: 2n = 36.

Taxon D. ± Caespitose; perennial, with flowering shoots up to 11 cms. and shorter sterile shoots, all shoots hairy. Leaves 5.11 - 6.39 x 1.16 - 1.76 mm., hairy, linear-lanceolate; bracts 3.43 - 4.41 x 1.24 - 1.78 mm., with scarious tip 0.56 - 0.98 mm. wide hairy, lanceolate. Sepals 5.03 - 6.65 x 1.60 - 2.01 mm., large scarious tip left 0.40 - 0.65 mm., small scaribus tip left 0.14 - 0.38 mm., large scarious tip right 0.40 - 0.65 mm., small scarious tip right 0.15 - 0.37 mm., scarious tip 0.64 - 0.77 mm. wide, hairy. Seeds 1.03 - 1625 x 0.73 - 0.91 mm., more or less spherical to ellipsoidal. Pollen grains 32.08 - 39.1711 in diameter, circular. Chromosome number: 2n = 36.

43

Fig. 7. Showing the plant and the mitotic metaphase of the taxon D. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525. 44 Taxon E. Loosely matted to caespitose; perennial, with flowering shoots up to 10 cms., hairy. Leaves 6.21 - 8.68 x 1.71 - 2.45 am., very slightly tomen- tose, linear-lanceolate; bracts 3.06 - 4.74 x 1.38 - 2.48 mm., with a scarious tip 0.61 - 0.99 mm. wide, hairy, lanceolate. Sepals 5.4 - 6.87 x 1.89 - 2.52 mm., large scarious tip left 0.55 - 0.88 mm., small scarious tip left 0.15 - 0.51 mm., large scarious tip right 0.51•- 0.88 mm., small scarious tip right 0.07 - 0.49 mm., scarious tip 0.73 - 1.18 mm. wide, hairy. Seeds 1.0 - 1.2 x 0.78 - 0.96 mm., more or less spherical to ellipsoidal. Pollen grains 36.2 - 46.6µ in diameter, circular. Chromosome number: 2n = 36.

Taxon F. Mostly matted to caespitose; perennial, with flowering shoots up to 11 ems. and slightly shorter sterile ones, hairs'. Leaves 5.50 - 8.24 x 2.02 - 3.14 mm., hairy, very slightly tomentose, linear-lanceolate; bracts 3.2 - 4.67 x 1.43 - 2.19 mm., with a scarious tip 0.55 - 1.13 mm. wide, hairy, lanceolate. Sepals 5.85 - 7.16 x 2.17 - 2.62 mm., large scarious tip left 0.58 - 0.92 mm., small scarious tip left 0.15 - 0.47 mm., large

4.5

Fig. 8. Showing the plant and the mitotic metaphase of the taxon.E. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525.

46

Fig. 9. Showing the plant and the mitotic metaphase of the taxon F. 2n chromosome number'= 54. Plant magnification = x.9. Chromosome magnification = x1525. 47 scarious tip right 0.58 - 0.95 mm., small scarious tip right 0.15 - 0.46 mm., scarious tip 0.66 - 1.06 mm. wide, hairy. Serds 0.87 - 1.29 x 0.66 - 0.98 mm., more or less spherical to ellipsoidal. Pollen grains 33.6 - 47.7µ in diameter, circular. Chromosome number: 2n = 54.

Taxon G. Caespitose to loosely tufted; perennial, with shoots up to 12 cms., hairy. Leaves 5.53 - 8.27 x 1.49 - 2.54 mm., hairy, lanceolate; bracts 2.77 - 4.7 x 1.35 - 1.97 mm., with a scarious tip 0.58 - 0.95 mm. wide, hairy, lanceolate. Sepals 5.22 - 6.68 x 2.13 - 2.64 mm., large scarious tip left 0.51 - 0.88 mm., small scarious tip left 0.15 0.58 mm., large scarious tip right 0.51 - 0.88 mm., small scarious tip right 0.15 - 0.51 mm., scarious tip 0.6 - 0.93 mmo wide, hairy. Seeds 0.8 - 1.23 x 0.69 - 0.9 mm., somewhat sherical to ellipsoidal. Pollen grains 35.8 - 46.64 in diameter, circular. Chromosome number: 2n = 36.

Taxon H. Cnespitose to loosely tufted; perennial, up to 6 cms. with sterile shoots shorter than flower- ing shoots, hairy. Leaves 5.39 - 9.72 x 2.08 - 3.56 mm., hairy, lanceolate; bracts 2.92 - 4.29 x

148

Fig. 10. Showing the plant and the mitotic metaphase of the taxon G. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525.

249

Fig. 11. Showing the plant of the taxon H. magnification = x.9. 50 1.33 - 2.34 mm., with a scarious tip 0.31 - 0.92 mm. wide, hairy, lanceolate. Sepals 5.4 - 7.36 x 2.2 - 2.86 mm., large scarious tip left 0.66 - 0.87 mm., small scarious tip left 0.15 - 0.47 mm., large scar- ious tip right 0.66 - 0.8 mm., small scarious tip right 0.1 - 0.44 mm., scarious tip 0.66 - 1.0 mm. wide, hairy. Seeds 0.86 - 1.27 x 0.61 - 0.95 mm., somewhat spherical to ellipsoidal. Pollen. grains 33.5 - 47.74 in diameter, circular. Chromosome number: 2n = ?

Taxon I. t Caespitose, perennial, with flowering shoots up to 8 cms. and shorter sterile ones, all shoots hairy. Leaves 5.83 - 1.04 x 1.47 - 2.83 mm., linear-lanceolate to lanceolate, hairy; bracts 3.43 - 4.16 x 1.55 - 2.19 mm., with a scarious tip 0.53 - 0.97 mm. wide, hairy, lanceolate. Sepals 5.1 - 6.8 x 2.0 - 3.1 mm., large scarious tip left 0.66 - 0.99 mm., small scarious tip left 0.37 - 0.62 mm., large scarious tip right 0.66 - 0.95 mm., small scarious tip right 0.31 - 0.6 mm., scarious tip 0.42 - 0.98 mm. wide, hairy. Seeds 0.99 - 1.25 x 0.79 x 0.96 mm., more or less spherical to ellip- soidal. Pollen grains 33.9 - 45.14 in diameter, circular. Chromosome number: 2n = 36.

51

Fig. 12. Shoeing the plant and the mitotic metaphase of the taxon I. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525. 52

Taxon J. Caespitose to loosely tufted; perennial, up to 11 cms. with sterile shoots shorter than flower- ing shoots, hairy. Leaves 4.96 - 8.59 x 1.83 - 2.86 mm., lanceolate, very slightly tomentose; bracts 3.0 - 4.96 x 1.71 - 2.6 mm., with a scarious tip 0.63 - 0.99 mm. wide, hairy, lanceolate. Sepals 4.83 - 6.35 x 2.48 - 3.18 mm., large scarious tip left 0.73 - 1.04 mm., small scarious tip left 0.22 - 0.75 mm., large scarious tip right 0.73 - 0.99 mm., small scarious tip right 0.15 - 0.66 mm., scarious tip 0.66 - 0.98 mm. wide, hairy. Seeds 0.96 - 1.28 x 0.74 - 1.0 mm., more or less spherical to ellip- soidal. Pollen grains 34.3 - 46.24 in diameter, circular. Chromosome number: 2n =

Taxon K. Caespitose to loosely tufted; perennial, up to 7 cms., sterile shoots shorter than flowering shoots, hairy. Leaves 6455 - 9.62 x 2.15 - 3.32 mm., more or less tomentose; bracts 3.32 - 4.96 x 1.64 - 2.94 mm., with a scarious tip 0.44 - 0.99 mm. wide, hairy, lanceolate. Sepals 5.41 - 6.86 x 2.54 3.29 mm., large scarious tip left 0.77 - 1.09 mm., small scarious tip left 0.29 - 0.69 mm., large scarious tip right 0.73 - 1.09 mm., small scarious

53

Fig. 13. Showing the plant of the taxon J. magnification = x.9.

54

Fig. 14. Showing the plant and the mitotic metaphase of the taxon K. 2n chromosome number = 36. Plant magnification x x.9. Chromosome magnification = x1525. 55 tip right 0.29 - 0.68 mm,, scarious tip 0.69 - 1.05 mm. wide, hairy. Seeds 0.74 - 1.28 x 0.63 - 0.96 mm., more or less spherical to ellipsoidal. Pollen grains 30.5 - 42.8µ in diameter, circular. Chromosome number: 2n= 36.

Taxon L. ± Caespitose; perennial, with flowering shoots up to 13 cms. and with shorter sterile ones, all shoots hairy. Leaves 8.32 - 12.64 x 1.62 - 3.18 mm., linear-acicular to lanceolate, hairy; bracts 2.92 - 4.08 x 1.82 - 2.7 mm., with a scarious tip 0.66 - 0.95 mm. wide, hairy, lanceolate. Sepals 4.64 - 6.29 x 2.42 - 3.26 mm., large scarious tip left 0.73 - 1.02 mm., small scarious tip left 0.37 - 0.58 mm., large scarious tip right 0.73 - 0.99 mm., small scarious tip right 0.31 - 0.58 mm., scarious tip 0.61 - 0.9 mm. wide, hairy. Seeds 1.02 - 1.35 x 0.84 - 1.18 mm., more or less spherical to ellipsoidal. Pollen grains 38.0 - 46.6µ in diameter, circular. Chromosome number: 2n = 126.

Taxon M. ± Caespitose; perennial, with flowering shoots up to 14 cms. and shorter sterile shoots, hairy. Leaves 8.5 - 15.9 x 1.73 - 2.66 mm., linear-

56

Fig. 15. Showing the plant and the mitotic metaphase of the taxon L. 2n chromosome number = 126. Plant magnification = x.9. Chromosome magnification = x1525.

57

Fig. 16. Showing the plant and the mitotic metaphase of the taxon M. 2n chromosome number = 108. Plant magnification = x:9. Chromosome magnification = x1525. 58 acicular to lanceolate, hairy; bracts 3.32 - 5.48 x 1.99 - 2.82 mm., with a scarious tip 0.64 - 1.17 mm. wide, hairy, lanceolate. Sepals 5.97 - 7.39 x 2.45 - 2.98 mm., large scarious tip left 0.73 - 1.02 mm., small scarious tip left 0.29 - 0.58 mm., large scarious tip right 0.73 - 0.95 mm., small scarious tip right 0.29 - 0.55 mm., scarious tip 0.69 - 1.28 mm. wide, hairy. Seeds 0.99 - 1.39 x 0.76 - 0.96 mm., more or less spherical to ellip- soidal. Pollen grains 35.4 - 46.94 in diameter, circular. Chromosome number: 2n = 108.

Taxon N. I Caespitose; perennial, with flowering shoot up to 9 cms. and shorter sterile shoots; all shoots hairy. Leaves 6.94 - 11.05 x 2.39 - 3.82 mm., linear-lanceolate) hairy; bracts 2.79 - 4.34 x 1.5 - 2.26 mm., with a scarious tip 0.46 - 0.95 mm. wide, hairy, lanceolate. Sepals 5.12 - 7.4 x 2.23 - 3.06 mm., large scarious tip left 0.66 - 0.95 mm., small scarious tip left 0.22 - 0.62 mm,, large scar- ious tip right 0.66 - 0.95 mm,, small scarious tip right 0.22 - 0.58 mm., scarious tip 0.38 - 1.08 mm. wide, hairy. Seeds 1.02 - 1.27 x 0.8 - 1.0 mm#, more or less spherical to ellipsoidal. Pollen grains 35.0 - 45.84 in diameter, circular. Chromosome number: 2n = 72.

59

Fig. 17. Showing the plant and the mitotic' metaphase of the taxon N. 2n chromosome number = 72. Plant magnification = x.9. Chromosome magnification = x1525. 60 Taxon 0. ± Caespitose to loosely tufted; perennial, with flowering shoots up to 9 cms. and longer than those of the sterile shoots, hairy. Leaves 6.32 - 9.7 x 1.73 - 2.47 mm., linear-lanceolate, slightly tomentose; bracts 3.2 - 4.92 x 1.42 - 2.4 mm., with a scarious tip 0.55 - 1.41 mm. wide, hairy, lanceolate. Sepals 6.25 - 8.0 x 2.55 - 2.95 mm., large scarious tip left 0.73 - 0.92 mm., small scarious tip left 0.17 - 0.58 mm., large scarious tip right 0.73 0.89 mm., small scarious tip right 0.17 - 0.51 mm., scarious tip 0.63 - 0.89 mm. wide, hairy. Seeds 1.07 - 1.3 x 0.8 - 0.96 mm., more or less spherical to ellipsoidal. Pollen grains 33.9 - 46.9µ in dia- meter, circular. Chromosome number: 2n = ?

Taxon P. Caespitose to matforming; perennial, with tomentose stems up to 20 oms. Leaves 8.0 - 13.75 x 2.42 - 4.17 mm., lanceolate, white, lanate; bracts 4.08 - 2.07 x 2.07 - 3.04 mm., with a scarious tip 0.8 - 1.83 mm. wide, lanceolate, tomentose. Sepals 5.87 - 8.13 x 2.85 - 4.1 mm., large scarious tip left 0.77 - 1.3 mm., small scarious tip left 0.4 - 0.8 mm., large scarious tip right 0.73 - 1.24 mm., small scarious tip right 0.4 - 0.77 mm., scarious

61

Fig. 18. Showing the plant of the taxon 0. magnification = x.9.

62

Fig. 19. Showing the plant and the mitotic metaphase of the taxon P. 2n chromosome number = 72. Plant magnification = x.9. Chromosome magnification = x1525. 63 tip 0.6 - 1.36 mm. wide, lanceolate, tomentose. Seeds 1.06 - 1.46 x 0.8 - 1.33 mm., more or less spherical to ellipsoidal. Pollen grains 40.2 - 45.1µ in diameter, circular. Chromosome number: 2n = 72.

Taxon Q. Caespitose to matforming; perennial, with tomentose stems up to 20 cms. Leaves 7.85 - 14.0 x 2.66 - 5.0 mm., lanceolate, innate; bracts 3.72 - 6.42 x 2.4 - 4.12 mm., with a scarious tip 0.92 - 1.75 mm. wide, lanceolate, tomentose. Sepals 5.75 - 8.25 x 2.89 - 4.05 mm., large scarious tip left 0.95 - 1.28 mm., small scarious tip left 0.1ih - 0.99 mm., large scarious tip right 0.91 - 1.28 mm., small scarious tip right 0.4 - 0.92 mm., scarious tip 0.93 - 1.37 mm. wide, lanceolate, tomentose. Seeds 1.09 - 1.53 x 0.74 - 1.1 mm., more or less spherical to ellipsoidal. Pollen grains 35.4 - 43.6µ in diameter, circular. Chromosome number: 2n = 72.

Taxon R. Caespitose to matforming; perenninl,up to 16 cms. with tomentose stems. Leaves 8.25 - 1.66 x 2.52 - 6.19 mm., lanCeolate, lanate; bracts 3.64 - 5.91 x 2.08 - 4.1 mm., with a scarious tip

64

Fig. 20. Showing the plant and the mitotic metaphase of the taxon Q. 2n chromosode number = 72. Plant magnification = x.9. Chromosome magnification = x1525.

65

Fig. 21. Showing the plant and splitting of the chromosomes. in two equal halves of the taxon R. 2n chromosome number'= Plant magnification = x.9. Chromosc4e magnificatiOn = x1525. 66

0.88 - 1.46 mm. wide, lanceolate, tomentose. Sepals 5.78 - 8.22 x 2.58 - 3.69 mm., large scarious tip left 0.88 - 1.19 mm., small scarious tip left 0.45 - 0.88 mm., large scarious tip right 0.85 - 1.19 mm., small scarious tip right 0.44 - 0.88 mm., scarious tip 0.85 - 1.37 mm. wide, lanceolate, tomentose. Seeds 1.1 - 1.5 x 0.74 - 1.1 mm., more or less spheri- cal to ellipsoidal. Pollen grains 35.4 - 48.1µ in diameter, circular. Chromosome number: 2n = 72.

Taxon S. Caespitose to matforming; perennial, with tomentose stems up to 16 cms. Leaves 7.85 - 15.0 x 2.47 - 4.85 mm., lanceolate, lanate; bracts 4.06 - 5.84 x 2.21 - 4.38 mm., with a scarious tip 1.09 - 1.97 mm. wide, lanceolate, tomentose. Sepals 6.03 - 7.67 x 3.33 - 4.2 mm., large scarious tip left 1.04 - 1.34 mm., small scarious tip left 0.58 - 1.02 mm., large scarious tip right 1.02 - 1.35 mm., small scarious tip right 0.55 - 0.97 mm., scarious tip 1.02 - 1.4 mm. wide, lanceolate, tomentose. Seeds 1.17 - 1.5 x 0.76 - 1.17 mm., more or less spherical to ellipsoidal. Pollen grains 34.3 - 44.7µ in diameter. Chromosome number: 2n = 72.

67

Fig. 22. Showing the plant and the mitotic metaphase of the taxon S. 2n chromosome number = 72. Plant magnification = x.9. Chromosome magnification = x1525. 68 Taxon T. Caespitose to matforming; perennial, with tomentose stems up to 15 cms. Leaves 7.96 - 13.78 x 3.25 - 6.24 mm., lanceolate, lanate; bracts 4.0 - 5.72 x 2.48 - 3.5 mm., with a scarious tip 0.89 - 1.83 mm. wide, lanceolate, tomentose. Sepals 6.1 - 7.75 x 2.89 - 4.0 mm., large scarious tip left 0.99 - 1.28 mm., small scarious tip left 0.51 - 0.88 mm., large scarious tip right 0.95 - 1.31 mm., small sca- rious tip right 0.53 - 0.9 mm., scarious tip 0.77 - 1.55 mm. wide, lanceolate, tomentose. Seeds 1.17 x 8.0 - 1.24 mm., more or less spherical to ellipsoidal. Pollen grains 34.6 - 43.64 in 9iameter, circular. Chromosome number: 2n = 36.

Taxon U. Caespitose to matforming; perennit 19 with tomentose stems up to 14 cms. Leaves 7.43 - 13.0 x 2.94 - 5.1 mm., lanceolate, lanate; bracts 3.84 - 5.98 x 2.53 - 4.0 mm., with a scarious tip 0.92 - 1.72 mm. wide, lanceolate, tomentose. Sepals 6.0 - 7.6 x 3.43 - 4.57 mm., large scarious tip left 0.99 -

1.4 mm., small scarious tip left 0.69 - 0.91 mm., large scarious tip right 0.98 - 1.42 mm., small scarious tip right 0.66 - 0.88 mm., scarious tip 1.0 - 1.38 mm. wide, lanceolate, tomentose. Seeds

69

Fig. 23. Showing the plant and the mitotic metaphase of the taxon T. 2n chromosome number = 36. Plant magnification = x.9. Chromosome magnification = x1525.

70

Fig. 24. Showing the plant and the mitotic plate of the taxon U. 2n chromosome number = 63. Plant magnification = x.94 Chromosome magnification = x1525. 71 0.95 - 1.4 x 0.73 - 1.09 mm., more or less spherical to ellipsoidal. Pollen grains 36.5 - 45.8µ in diameter, circular. Chromosome number: 2n = 63.

Taxon V. Caespitose to matforming; perennial, with tomentose stems up to 15 cms. Leaves 7.45 - 13.9 x 2.95 - 5.63 mm., lanceolate, lanate; bracts 3.65 - 5.79 x 2.0 - 3.25 mm., with a scarious tip 0.82 - 1.57 mm. wide, lanceolate, tomentose. Sepals 5.6 - 7.87 x 2.69 - 3.43 mm., large scarious tip left 0.8 - 1.21 mm., small scarious tip left 0.44 - 0.77 mm., large scarious tip right 0.8 - 1.17 mm., small sca- rious tip right 0.29 - 0.75, scarious tip 0.66 - 1.36 mm. wide, lanceolate, tomentose. Seeds 1.0 - 1.6 x 0.8 - 1.22 mm., more or less spherical to ellipsoidal. Pollen grains 30.9 - 45.1p. in diameter, circular. Chromosome number: 2n = 72.

Taxon W. Caespitose to matforming; perennial, with tomentose stems up to 25 cms. Leaves 1.14 - 1.57 x 2.88 - 4.91 mm., lanceolate, lanate; bracts 3.19 - 5.93 x 2.0 - 3.43 mm., with a scarious tip 0.99 - 1.7 mm. wide, lanceolate, tomentose. Sepals 6.76 - 8.54 x 2.89 - 3.75 mm., large scarious tip left 0.88 - 1.19 mm., small scarious tip left 0.46 - 0.87 mm.,

72

Fig. 25. Showing the plant and the mitotic metaphase of the taxon V. 2n chromosome numbet= 72. Plant magnification = x.9. Chromosome magnification = x1525.

73

Fig. 26. Showing the plant and the mitotic metaphase of the taxon W. 2n chromosome number = 72. Plant magnification = x.9. Chromosome magnification = x1525. 74 large scarious tip right 0.88 - 1.17 mm., small scarious tip right 0.)1LI - 0.82 mm., scarious tip 0.89 - 1.33 mm. wide, lanceolate, tomentose. Seeds 1.18 - 1.57 x o.84 - 1.22 mm., more or less spherical to ellipsoidal. Pollen grains 32.0 - 43.2µ in diameter, circular. Chromosome number: 2n = 72.

Taxon X. Caespitose to matforming; perennial, with tomentose stems up to 20 cms. Leaves 1.0 - 1.59 x 2.74 - 4.85 mm., lanceolate, lanate; bracts 4.53 - 5.74 x 2.94 - 4.0 mm., with a scarious tip 0.85 - 1.42 mm. wide, lanceolate, tomentose. Sepals 5.91 - 7.72 x 3.33 - 4.48 mm., large scarious tip left 0.95 - 1.6 mm., small scarious tip left 0.6 - 1.28 mm., large scarious tip right 0.95 - 1.53 mm., small sca- rious tip right 0.58 - 1.19 mm., scarious tip 0.95 - 1.28 mm. wide, lanceolate, tomentose. Seeds 1.15 - 1.55 x 0.85 -.1.15 mm., more or less spherical to ellipsoidal. Pollen grains 35.0 - 43.2µ in diameter, circular. Chromosome number: 2n = 72.

75

Fig. 27. Showing the plant and the mitotic metaphase of the taxon X. 2n chromosome number = 72. Plant magnification = x.9. Chromosome magnification = x1525. 76 CYTOLOGY

Review of literature:-

Heitz (1926) published the cytology of the genus Cerastium. He counted more than 100 chromosomes in C. triviale Link. corresponds to a haploid number of about 55. Wulff (1937) working on the flora of Schleswig Holslein, reported 18 small, globular chromosomes in the pollen mother cells of C. tetrandrum Curt. Bocher (1938) working on Scandinavian material, determined the chromosome number of three speciea as follows:-

C. cernstoides (L.) Britton, 2n = 40 C. caespitosum Gibel., 2n = c, 144. C. alpinum L. 2n = 72.

Rohweder (1939) determined the meiotic chromo- some number of seven species from Europe as follows:-

C. arvense L., n = 36. C. candidissimum Correns, n.= 18. C. chloraefolium Fisch. and May, n = 18. C. glcmeratum Thuil.„ n = 36. 77 C. semidecandrum L. n = 18 C. tetrandrum Curt. n = 18 C. tomentosum L. n = 18 C. triviale Link. n = 72

Flovik (1940). determined the chromosome number of C. regellii Ostf from Spitzergen 2n = 72. Hagerup determined the chromosome number of C. subtetrandrum Murb., from Scandinavia n = 36. Hagerup (1944) found the chromosome number of the Scandinavian material of the following species:-

C. glutinosum Fries 2n = 72 C. brachypetalum Pers. 2n = 90 C. glomeratum Thuil 2n = 72 C. triviale Link. 2n = 126

In this account he proposed 9 as the basic chromosome number for the genus Cerastium and com- mented upon the high degree of polyploidy, especially in the polymorphus species C. triviale. He further suggested that C. triviale might have some other chromo- some series. Ldve and LBve (1900 determined the chromosome - 78 number of C. alpinum L. from Abisko and Sweden, 2n = 72. Love and LO've (1948) published a list of chromo- some counts for northern species which included a new count 2n = 108 made by the Danish worker STensen and Westergaerd for C. alpinum L., collected from Green- land. They also determined the chromosome number of C. cerastoides (L.) Brit. from Greenland 2n = 40. Favarger and Sollner (1949) reported the chromo- some number of C. cerastoides L. Brit. from the Alps, 2n = 38. In the same paper they recorded 2n = 38 for C. anomalum Waldst. and Kit., and 2n = 36 for C. latifolium L., C. pedunculatum Gaud., and C. uni- florum Clairv., all from the continent of Europe. Tischler's chromosome list (1950) besides many of the above counts, includes also seven more made by Mattick, as follows:-

C. cerastoides (Le) Brit., 2n = 36 C. latifolium L., 2n = 36 C. uniflorum Clairv. 2n = 36 C. arvense L. 2n = 72 C. brachypetalum Pers. 2n = 52 C. holosteoides Fries 2n = 144 C. fontanum Baum 2n = c. 120 79 Soliner (1950) and Brett (1950) working inde- pendently, each published a short list of chromosome numbers. Soliner reported the chromosome number of C. arvense 2n = 36 and 2n = 72 from different locali- ties of Alps. C. banaticum Fisch. 2n = 38, and C. perfoliatum L. 2n = 38. Brett's list consisted of:-

C. vulgatum L. from Britain, 2n = 136 C. alpinum L. from Britain, 2n = 72 C. alpinum L. from Britain, 2n = 108 C. perfoliatum L. from Britain, 2n = 38

Bdcher and Larsen (1950) published the results of their cytological studies of materials collected by the former during his expedition to Greenland. They reported the chromosome number of C. alpinum 2n = 54 and 2n = 72, C. cerastoides 2n = 34 and 2n = 38. Bdcher considered it possible that the number 2n = 40 (1938) could be interpreted as 2n = 38. Brett (1951) working on British materials re- ported the chromosome numbers of C. arvense L. 2n = 72, C. viscosum L. 2n = 72, C. dichotomum L. 2n = 38, C. tomentosum L. 2n = 72, and C. alpinum L. 2n = 144. 80 This last approximate count was for material collected in Abisko. In 1952, two more papers appeared, one by Soilner (1952) consisted of a list of chromosome counts of which the following are either new counts for the species, or counts for species not previously investigated: -

C. atlanticum D.R. 2n = 72 C. campanulatum Viv. 2n = 36 C. austro-alpinum Kunz. 2n = 36 C. carinthiacum Vest. 2n = 36 C. fontanum Baumg. 2n = c. 144 C. inflatum Link. 2n = 38 C. julicum Schellm. 2n = c. 36 C. macrocarpuill Schur. 2n = 72 C. maximum L. 2n = 38 C. nutans Raf. 2n = 35 - 36 C. pumilum Curt. 2n = 90 C. soleiroli Ser. em. Busch. 2n = 72 C. sylvaticum W. 2n = c. 36

Also a species of uncertain identity, thought to be C. holosteoides Fries 2n = 162. All these mate- rials were of continental origin with the exception of C. inflatum from Iran and C. maximum and C. nutans from N. America. 81 Brett (1952) gave the following counts:-

C. arvense L. 2n = 36 from N. America C. maximum L. 2n = 38 from N. America C. brachypetalum Pers. 2n = 90 for C. holosteoides 2n = 136 - 152 British C. semidecandrum L. 2n = 36 '!material C. tetrandrum Curt. 2n = 72

C. anomalum W. and K. 2n = 38 for C. chloraefolium F. and F. 2n = 38 conti-nental C. davuricum Fisch. 2n = 38 material

She also suggested nine and nineteen the basic chromosome numbers for the genus Cerastium in this paper. Sollner Roland (1953a,b) reported the chromo- some number in C. strictum L. 2n = 36, and in C. ar- vense L. 2n = 72 from central and north-western Europe. Brett (1953) reported the chromosome counts of the following British species:-

C. arcticum Lange 2n = 108 C. cerastoides (L.) Brit. 2n = 38 C. pumilum Curt. 2n = 90 82 Sollner Raland (1954) in a cytotaxonomic study of the genus Cerastium, reported the chromosome num- bers of 43 species, 32 species have chromosome numbers which are multiple of n = 9 (18,36,45,54,72) though species with n = 9 were never found. 8 species have n = 19 and 2 species have n = 17. These may have formed by non-disjunction. One species C. teno- reanum has n = 26. This may have arisen by a cross between n = 18 and n = 36 form. C. pumilum has n = 50 (or 51) chromosomes. One species C. ramosis- simum Boissier has n = 22 - 23 chromosomes. Below is given the name and locality of the 43 species:-

C. cerastoides (L.) Valais 2n = 38 Britton C. anomalum Copenhague 2n = 38 Waldst et Kit. C. dahuricum Fisch. Stockholm 2n = 38 C. chloraefolium Fisch• et Meyer Stockholm 2n = 38 C. perfoliatum L. Espagne 2n = 38 C. maximum L. Canada 2n = 38 C. dichotomum L. Algerie 2n = 38 C. inflatum Link. Iran 2n = 38 C. latifolium L. Valais 2n = 36 C. uniflorum Clairv. Valais 2n = 36

83 C. pedunculatum Gaud, Valais 2n = 36 C. carinthiacum Vest. Autfichc,- 2n = 36 C. austroalpinum Kunz Tessin 2n = 36 C. alpinum L. Canada 2n = 72, 108 C. edmondstonii (Wats.) Murb. Scotland 2n = 108 C. candidissimum Correns Austria 2n = 36 C. biebersteinii D.C. Cluj 2n = 72 C. tomentosum L. Italie 2n = 36 C. decalvans Schloss et Vuk. Switzerland 2n = 72 C. boiss,eri Gren. Corse 2n = 72 C. julicum Schellmann Aulchi, 2n = 36 C. banaticum (Roth.) Heuff Switzerland 2n = 72 C. soleirolii Seringe Corse 2n = 72 C. arvense ssp. suffruticosum (L.) Koch Var 2n = 36 C. arvense ssp. strict7Thaenke) Gaudin Tessin 2n = 36 C. arvense ssp. commune L. Tessin 2n = 36, 72 C. subtriflorum (Rchb.) Pacher Slovenie 2n = 36 C. sylvaticum W.K. Budapest 2n = 36 84 C. macrocarpum Schur. em Gartner Styrie 2n = 144 C. fontanum Baumge Autricho 2n = 144 C. holosteoides Fries ampl.Hylander voir.tableau 2n = 144 C. comatum Desvaux Corse 2n = 34 C. brachypetalum Fers. Copenhague 2n = 90 C. tenoreanum Serin-e Lugano 2n = 52 C. atlanticum D.R. Algerie 2n = 72 C. glomeratum Thuill Copenhague 2n = 72 C. ramosissimum Boisser Portugal 2n = 44; 46 C. campanulatum Viviani Italie 2n = 34 C. pentandrum L. Castille 2n = 36 C. glutinosum Fries Slyrie 2n = 72 C. pumilum Curtis Neuchatel 2n = 100, 102 C. semidecandrum L. Neuchatel 2n = 36 C. nutans Raf Canada 2n = 36 C. tucumanense Pax Argentine 2n = 36 C. beeringianum Cham Canada 2n = 72 Larsen, K. (1954) in his account of the chromosome numbers of the European flowering plants, gave the chromosome number of C.strictum L. asp. sltrictum 2n = 36. Brett (1955) reported the chromosome numbers of British and foreign specico listed below:-

C. alpinum L. British 2n = 72 C. arcticum Lange II 2n = 108 C. arvense L. It 2n = 72 C. brachypetalum Pers. it 2n = 90 C. cerastioides (L.) Britton II 2n = 38 C. glomeratum Thuill it 2n = 72 C. holosteoides Fries. It 2n = 136 - 152 C. vulgatum L. It 2n = 137 - 147 C. pumilum Curt. It 2n = 90 - 95 C. semidecandrum L. o 2n = 36 C. tetrandrum Curt. tt 2n = 72 C. tomentosum L. tt 2n = 72 C. alpinum L. Sweden 2n = 72 C. anomalum Waldst et Kit. Newcastle 2n = 38 C. arvense I. Austria, 2n = 38, 36 Canada C. cerastoides (L.) Britton Switzerland 2n = 38 C. chloraefolium Fisch et Mey. Prague 2n = 38 C. davuricum Fisch. Germany 2n = 38 86

C. maximum L. Canada 2n = 38 C. perfoliatum L. England 2n = 38 C. thomasii Ten. Italy 2n = 72

Multivalents were observed by Brett in three of the the British spp., namely, C. pumilum, C. arcticum and C. holosteoides. In a few spp., certain of the chromosomes in each complement had a satellite. In C. semidecandrum four and in C. arcticum at least three such chromosomes were observed. Mve and Delve (1956b) reported the chromosome numbers of the Iceland material given below:-

C. glomeratum Thuill (= viscosum) 2n = 72 C. fontanum Baumg. Gartner ssp. scandicum Gartner, 2n = 144 C. holosteoides 7r., Hyl.; (= trivale,vulgatum) spp. holosteoides 2n =1L4 C. alpinum L. 3.Str. 2n = 72 C. arcticum Lge. 2n = 108 C. cerastoides (L.) Britton 2n = 38

Blackburn and Morton (1957) published the. chromosome numbers of British and Portugal materials as follows:- 87

C. glomerctum Thuill (=Ariscosilm), British and Portugal, 2n = 72 C.alpinum L. S. Str. 2n = 72, 108 C. arvense L. Gtr. 2n = 72 C. brachypetalum Pers. 2n = 90 C. cerastoides (L.) Britton 2n = 38 C. edmondstonii (Wits.) Murb. 2n = 72 C. glutinosum Fr., Portugal, 2n = 72 C, pumilum Curtis 2n = 72 C. semidecandrum L. 2n = 36 C. tetrandrum Curt. 2n = 72 C. vulgatum (triviale) L. 2n = 72, 126, 144, 18o

Jorgensen, Sorensen and Viestergaard (1958) in of their taxonomical and cytological survey/flowering plants of Greenland, reported the chromosome number of C. alpinum L. 2n = 108. L6've and Chenaveeriah (1959) in the cytotaxonoy of Cerastuim holosteoides reported the chromosome numbers of the following species from Northern Europe and Iceland:-

C. holosteoides Fr. Hyl. (= trivale vulgatum) ssp. holosteoides 2n = 1W1 ssp. globrecens (G.F. Mey) Moschl. 2n = 144 ssp. pseudo holosteoides 2n = 144 88 Larsen, K. (1960) reported the chromosome num- ber of C. glomeratum Thuill (= viscosum) from Canary Islands 2n = 72. Sokolovskaja and Strelkova (1960) reported the chromosome number of C. alpinum L. E. lat from Khibiny mountain in European Russia 2n = 54. 89

Cytological observations All the 24 taxa, A - X, were collected together with seeds from the high plateau of Mount Maiella in the central Italian apewnines 8,000-10,000 ft high. Most of the mitotic counts were made from the root-tipof've/5y young seedlings. In some of these taxa the yo,..ngest- root-tip from the fully grown plant wf,s, used. The meiotic studies of the taxa under in- vestigation could not be done because the plants in any taxon producedno flowers at all. The mitotic counts from two to three plants from each taxon were made for the determination of the chromosome numbers of the taxa. In the mitotic counts the chromosomes of the Cerastium taxa are r eLIZ. very small in size so that their morphology could not be investigated. The mitotic counts of the taxa under in- vestigation are as follows. Taxon A. Mitotic counts in the taxon A determined

2n = 36. Metaphase shows 2n = 36 ' (Fig. 4, p. 39). the chromosomes are all rod-shapAd. 90 Taxon B. Mitotic counts in the taxon B showed p.40 2n = 36, (Fig.5/ metaphase). Chromosomes rod- shaped and of about the same size as those of taxon A. Taxon C. Mitotic counts in the taxon C gave 2n = 36 metaphase (Fig.6,p41). All the chromosomes are rod- shaped, more or less of the same size as those of taxon A and B. Taxon D. Mitotic counts in the taxon D revealed 2n = 36, metaphse (Fig.7p43. All the chromosomes are rod-shaped more or less of the same size as those of taxa A, B, and C. Taxon E. Mitotic counts in the taxon E gave 2n = 36. All the chromosomes are somewhat short thick rods and are shorter than those of taxa A, B, C, and D. (Fig.8 p45metaphase). Taxon F. Mitotic counts in the taxon F gave 2n = 54. All the chromosomes are oval and smaller in size p. L1.6 than that of taxon E. (Fig. 9/ showing metaphase). 91 Taxon G. Mitotic counts in the taxon G showed 2n = 36. All the chromosomes are short thick rods. (Fig.lo, p.48 showing metaphase). Taxon I. Mitotic counts in the taxon I gave 2n = 36. All the chromosomes are short rods, and of about the p.51 same size as those of taxon G. (Fig. 12,/showing metaphase). Taxon K. Mitotic counts in the taxon K gave 2n = 36. All the chromosomes are rod shaped. (Fig. 14, p. 5L showing metaphase) and of about the same size as those of taxa A - D. Taxon L. Mitotic counts in the taxon L gave 2n = 126. The chromosomes are short rods about one-half as ,p. 56 long as those of taxon D. (Fig. 15/ showing metaphase). Taxon M. Mitotic counts in the taxon M showed 2n = 108. Chromosomes are short thick rods, somewhat P. 57 shorter and thicker than in taxon L. (Fig. 16/showing metaphase). 92

Taxon N. Mitotic counts in this taxon gave 2n = 72. Chromosomes are all thick dot like, and are some- P.59 what bigger than in taxon F. (Fig.17/ showing metaphase). Taxon p. Mitotic counts in the taxon P showed 2n = 72 chromosomes. The chromosomes are all rod shaped. 62 (Fig.19/showing metaphase). Taxon Q. Mitotic counts in the taxon Q gave 2n = 72. Each chromosome more or less oval shaped. (Fig. 20, p. 64 showing metaphase). Taxon R. Mitotic counts in the taxon R gave 2n = 72. The chromosomes are all rod-shaped. Most of the p. 65 chromosomes split into two. Fig.2l/ showing split- ting of the chromosomes in two equal halves of about the same size as those of taxon P. Taxon S. Mitotic counts in the taxon S gave 2n = 72 P. 67 chromosomes; each a short rod. (Fig. 22/showing metaphase). 93

Taxon T. Mitotic counts in taxon T gave 2n = 36. Chromosomes are all oval-shaped and more or less p. 69 similar to those of taxon Q. (Fig.23/ showing metaphase). Taxon U. Mitotic counts in taxon U revealed 2n = 63. Each chromosome is more• or less. oval and somewhat similar to those of taxa Q and T. (see Fig.2L, p. 70) Taxon V. Mitotic counts in taxon V showed 2n = 72. Chromosomes are short rods, their size varies. (Fig.25 p.72 showing metaphase). Taxon W. Mitotic counts in the taxon W gave 2n = 72, P. 73 Chromosomes are all short rods. (Fig.26/ showing metaphase). Taxon X. Mitotic counts in the taxon X gave 2n = 72. P. 75 Chromosomes are all rod-like. (Fig.27/ showing metaphase). 94 BIOMETRICS

The genus Cerastium contains many taxa whose exact delimitation presents special difficulty. One such group is that of the perennials represen- ted on Mount Maiella, central appennilaesAItaly. The specimens of Cerastium spp. under in- vestigation were delimitated into twenty-four taxa by eye. For the delimitation of the groups into taxa, fifteen characters were drawn from the fol- lowing five morphological units:- Leaf; Bract; Sepal; Seed; and Pollen Grain. The characters ultimate chosen were those which appeared to be the least plastic under varied growth conditions. The main purpose of this study was to express the limits and variability within and between taxa to quantitative terms. Only the quantitative charac- ters wbich are:readily employed in statistical treatments were considered to be workable. These characters were easy to measure and transform into desirable statistical terms to give taxonomic meaning. The number of specimens from which mea- surements were taken is shown in the appendix to- gether with the means of all measurements. The characters selected are described in chapter 95

"Methods and Material", (Page 17), and the taxono- mic description of the 24 taxa is given in the pre- vious chapter (Page 34). A close examination of the taxa involved showed that there was considerable variation and that complete correlations 1,:ere seldom, if ever possible. The main difficulty was to find charac- ters unaffected by depauperisation or unusual luxu- riance. By growing plants under various conditions (See Methods & Material, Page 27,) it was found that with the exception of a general greater luxu- riance the growth of all the taxa showed little difference from that in nature. The characters selected for further study were those which remained reasonably constant despite the varying conditions. The next problem was to select a suitable method of utilising the data collected to show the number and statistical affinity of groups within the totality of specimens examined. The requirements of such a method are that it should be as objective as poss- ible, simple in application and capable of revision and independent verification by other workers. The complexity of the taxa involved implied that the method should also have certain precise stati- stical properties. It was important that tho 96 characters chosen should not be merely different expressions of the same genetical determinant but should show some independent variation. If they were not, overweighting of characters would take place and the real extent of variation between samples of the same taxon would be obscured. On the other hand discrimination between taxa would depend upon the recognition and utilization of characters which would be correlated within taxa but show significant mean differences between taxa. It would be a further advantage if the method em- ployed linear functions of the selected measure- ments. Various statistical methods have been developed and employed of which the discriminent analysis Fisher (1936); Cluster analysis Boeke (19L12); factor analysis Sokal (1958); canonical variates Ra0(1948); R- and Q- techniques Cattell (1952) have considerable value for the solution of certain problems. Most methods result in coefficients of similarity ranging between unity and zero, the former for perfect arrangement and the latter for none whatever. These various types of coefficients 97 are employed both in R- and Q- techniques. Re- views of the different types of coefficients which have been proposed and used can be found in Cole (1949 - 1957), and Dagnelie (1960). Some of the methods are usually much simpler to work out than others and each method has its own significance and advantage in the solution of different kinds of problem. It took a considerable time to decide which method should be employed in the present in- vestigation. The application of numerical methods to taxonomy dates from the rise of biometrics in the last century. Heincke (1898) employed a measure of phenetic distance to distinguish between races of the herring. It was early realized that bio- metrics could be used in systematics, but the only outstanding development was that of discriminant analysis (Fisher, 1936) in which each character is given a loading such that there is the least probability of misidentifying an individual taken at random. A very well worked example is given by Whitehead (1954). Related work is that of Rescigno and Maccacaro (1960). The classificatory 98 method of Lockhart and Hartman (1963) also extracts discriminatory characters. Pearson (1926) intro- duced the coefficient of "Racial Likeneas". It was widely employed in anthropology but does not seem to have been used by taxonomists. The C.R.L. was close to being a coefficient of taxonomic simi- larity, and was subsequently developed by Mahalanobis in the form of the "Generalized Distance" statistic, which is also formally a coefficient of this kind (see Rao 1948). Anderson and Abbe (1934) and Anderson and Whitaker (1934) used a similar stati- stic, which was also equivalent to a diagonal in a multidimensional Euclidean space, and which they called the "General Index". These statistics though mathematically quite adequate were mainly developed as discriminant function to aid in the allocation of individuals to existing taxa and not as methods for creating taxa. These methods were in particular based on few rather than many charac- ters. These techniques, with others developed later, have been widely and successfully used for the study of certain limited taxonomic problems, for example the admirable work of Blackith (1957) on sexual and phase variation in locusts. 99 Smirnov (1925) established types on a quantitative basis. His work was evaluated from different points of view by Henning (1950) and Sokal (1962b). Zarapkin (1939) developed a tech- nique called Divergenzanalyse, arriving at a quantity analogous to taxonomic distance. Schilder and Schilder (1951) developed Affinitatsrechnung, that is also a computation of taxonomic distance. These methods could not be used at the time they developed mainly because of the computational difficulties. (The entrenched nature of phylo- genetic systematics also added difficulties in- surmountable at that time). The other early methods specially in- tended for taxonomy are those of Frobes (1933), Anderson and Owenbey (1939), Sturtevant (1939, 1942), Boeke (1942), James (1953), Stallings and Turner (1957), Hudson, Lanzillotte, and Edwards (1959), and Chillcot (1960), based on variations of matching coefficients. These authors also employed a smaller number of characters and did not give equal weight to every character. A larger number of characters would have yielded better results. Although they were correct in 100 not giving equal weight to every character. It is considered here that every character carries weight but not equal weight and thus the weighting is always differential. It is quite illusory and deceptive that every character carries equal weight (Sokal and Sneath 1963) and some others. Since one character may be more useful in classifying a group of specimens than other characters, equal weighting is quite unjustifiable. Kiriakoff (1962) has critised the neo-Adansonian school and argued against equal weighting. Proctor and Kendrick (1963) criticised the equal weighting and produced a worked example in defence of unequal weighting. Unless and until a perfect defensible system for giving equal weight to every character, can be provided, weighting must remain differential. Many papers were given on this subject at the 10th International Botanical Congress, Edinburgh (1964) and examples of the use of various statistics given. This work on Cerastium was referred to by J.N.R. Jeffers in his paper on canonical variates. I am grateful to my supervisor, Dr. F.H. Whitehead, Dr. R.L. Blackitt of Zoology Dept., Imperial College (now at Melbourne, Australia) 101 and Mr. J. N. Jeffers of Alice Halt Lodge, Forestry Research Commission, for suggesting the method of working out 'Canonical Variates' which abbeared to be best suited to the present problem. The Sirius elec- tronic computer of Alice Halt Lodge Was cuito capable of taking such a programme. Dr. J. N. R. Jeffers took a great deal of trouble in processing the data and modifying the method to suit the present data. Canonical Vectors which was first introduced and employed by C. a. Rao (1948) in deriving group cons- tellations of 22 Indian tribes. For details of the procedures and the formulae involved see Rao (1948). The method canonical variates was first intro- duced by Rao (1948) and later fully deScribed by Rao (1952). Relatively few publications have appeared on the application of canonical variates to taxonomic oroblems. However Blackith, Davies and Moy (1963) gave a biometric analysis of development in Dysdercus fasciatus, together with a brief outline of the biometric principles involved. This method is being ab)lied for the first time to botanical taxonomy on such a large and complex taxonomic group involving as many as fifteen variables (characters), showing a fine gradient of characters. 102

The present use of this statistical method is a new extension of its application to taxonomic problem:, The electronic computers have made possible the use of this method in the classification and identification of plant populations. Gardiner and Jeffers (1963) read a paper to the British Association for thei,.dvance- ment of 3cience. They gave an appreciable account of component analysis giving three worked out examples of birch species. Jeffers (1964) read a paper at the 10th international Botanical Congress Edinburgh, further giving a very clear account of the usefulness of this method. This method reduces the large quantities of data, relating to many measurements of individual plants to a small number of essential dimensions and then by computing the means of canonical variates, taxa can be classified and identified. In the present investigation this method not only shows variation between the taxa, but also within the taxa. It also works out the diff- erential weighting to the variables used in the investigation. It is a measure of the usefulness 103 of this method that despite the quite conside- able number of inapplicable samples, discrimina- tion was still obtaired. Most statistic& which have been used in biometric studies so far, have been concern- ed with the type of problem which involves the correct assignment of an unknown individual to an already known grouping. Usually known species involive'd and the statistics require that the basic data for a representative aample from these known species is obtained. The statistics such as; discriminent function then makes a measure of the correlations of the data within and between the known groups and from this pro- vides a means of determining the probabilitiea of an unknown individual belong3to either groups. Canonical variates as, used in this work differs in that data Jf a large number of putative groups was obtained. These did not correspond with already delimited taxa. The groupings of initial taxa were subjective but by means of the statistical treatment an object- ive assessment of their taxonomic rank was- made possible.. Thus some of the original taxa have 104 been merged into a single taxon since the statistic showed that objectively no clear distinction existed between them which was suitable for "keying" by normal taxonomic procedures. By means of the degrees of association between the different taxa within the two major groups, it has been possible to assign them to different species. All the taxa within Group A have been inc- luded in the species C.album. The taxa from Group B have been assigned to four different species i.e. C.laricifolium, C.thomasii, C.matrense and C.ripoi. 105

The total variability calculated from the determinantal matrix derived from between the taxa and within the taxa sums of squares and products was 242, 832.17. Five components were calculated which accounted for 99.72 per- cent wfiriability. 'Z.Lithough the first of these five components alone accounted for 95.97 per cent of the total variability, but the remaining four components were still significant by the criterion developed by Rao(1952). The means of canonical variatosgave comparisons which were taxonomically interpretable. The very high proportion of variability shown by the first of the canonical variates was actually due to the difference between the two major groups of,taxa which were quite recogniz- able even by conventional methods of taxonomy. For this reason, the tinalysls was done separately fore the two groups A and B which are describeA below.

106 The procedure used to work out the canonical variates of nine taxa (P -• X) of group A and fifteen taxa (A - 0) of group B is given below. Measurements of fifteen variables (some pre- viously used as diagnostic and also some new vari- ables not previously used) were made in arbitrary units 1 unit =73 p. except fo2 one variable (pollen grain diameter) where 1 unit = 3.73 The means of variables for each individual within each taxon have been given in the appendix (Page178). From the means of individuals of each taxon) the means of variables for each taxon were computed for both the above mentioned groups. These are given below:

Grou_11 A 74.60 30.01 9.66 5.39 9.28 5.00 8.15 49.38 L ,55 8.24 15.55 12.06 9.95 69.01 22.29 B 78.74 29.97 9.56 5.27 9.10 4.95 9.70 53.25 25.15 8.50 14.84 11.45 10.24 79.85 20.28 C 67.67 29.82 9.30 5.33 8.90 5.12 8.17 43.13 25,11 7.28 14.88 11.34 9.45 72.77 19.83 D 78.11 25.18 6.70 3.10 6.52 4.26 9.)1h 5529 18.74 9.85 16.00 11.48 9.58 76.93 19.50

107

E 83.68 30.16 9.39 3.32 9.17 3.08 11.31 52.83 24.67 11.18 15.56 11.84 11.25 98.35 27.48 F 88.78 32.56 9.95 4.89 9.87 4.60 11.31 55040 24.59 11.01 15.85 11.73 11.29 95.66 33.43 G 79.32 32.47 9.60 3.69 9.47 3.33 10.28 50.14 23.51 9.96 15.14 10.91 11.60 91.60 25.94 H 81.98 33.59 10.15 4.77 9.96 4.46 10.85 50.27 25.29 10.16 15.68 11.74 10.81 97.07 33.56 80.90 34.29 11.07 6.42 10.93 6.24 9.58 52.20 26.48 9.80 15.53 11.98 10.32 101.35 28.35 J 77.25 37.91 11.89 6.64 11.62 6.27 11.04 52.72 28.15 10.93 15.38 12.07 10.86 95.84 32,69 K 83.97 40.22 12.96 7.08 12.77 6.93 12.25 56.10 31.98 11.82 15.65 12.09 10,72 115.04 40.07 L 77.00 38.15 11.85 6.21 11.68 5.99 10.80 49.86 29.97 10.91 16.48 12.57 11.19 136.96 32.55 M 91.79 36.55 11.66 6.59 11.50 6.39 12.46 58.76 32.96 13.21 16.42 12.41 11.37 150.94 30.73 N 86.18 34.77 11.54 6.14 11.15 5.87 11.52 52.63 25.57 10.78 15.88 12,05 11.00 119.45 40.58 O 93.34 37.28 11.38 5.36 11.14 5.16 11.03 56.94 26.53 11.94 16.12 12.08 11.38 111.28 29.03

108 Group A P 94.85 47.01 14.82 8.98 14.45 8.76 lh.01 66.67 35.-65 16,50 17.83 13.56 11.43 155,87 51.22 Q 92.28 45.89 14.79 9.07 14.53 8.66 14.54 61.00 39.98 16.35 17.55 12.60 10.69 151.33 51.23 R 95.83 43.23 14,19 8.22 14.00 7.95 14.40 64.02 16.14 18.28 13.22 10.66 157.65 53.80 s 93.23 51.35 16.15 10.25 15.88 10.00 16.25 65.99 43.49 18.38 17.55 13.11 11.20 153.88 52.09 T 94.20 47.63 15.63 9.57 15.39 9.41 15.61 65.61 42.03 18.19 18.10 13.48 10.44 147.74 65.11 U 93.50 54.71 17.18 10.90 17.08 10.69 16.74 65.71 44.05 18.33 16.99 13.15 11.26 129.94 56.66 ✓ 95.16 41.76 14.29 8.52 14.02 8.19 14.78 67.48 37.74 16.38 18.23 14.05 10.80 139.17 64.55 W 101.31 44.57 14.01 8.41 13.74 8.26 14.44 67.80 35,95 17.76 18.06 13.37 10.74 193.69 53.03 x 89.91 50.81 16.11 11.13 15.95 10.93 14.99 69.47 47.79 16.66 18.21 13.39 10.38 188.27 55.75

In the next step the total variability i.e. variction,buen and within the taxa, was deter- mined in the computer. 109

The total variability for group A was 82520.79 The total variability for group B was 181474.48 Tne total variat nity in group A and in Group B was then divided into fifteen components. The var- iability for each of the fifteen components in both the groups is given below:

Variability divided into fifteen components in group A:

A2 3 A235 A268 A299 R329 Ll .0000806245 9' R14 L2 .0000012907 9 9 A38 'The number A77 indicates that Al 28 8142 the decimal should L3 .0000003355 9 Al9 be read after the R23 L4 .0000001576 9 9th place from A30 R424 the left. L5 .0000000575 A37 A188 R264 L6 .0000000348 9 A153 R197 L7 .0000000135 9 A59 A61 R99 L8 .0000000082 9 Al 52 R270 L9 .0000000016 9 110 A431 A433 R779 L10- .0000000013 9 A206 A338 8451 LII -.0000000006 9 A278 R)489 L12 -0000000000 A679 A733 A786 R827 L13 -.0000000002 9 8123 L14 .0000000001 9 R145 Total _ = L15 r0000000000 9 82520,12 Q-1-9 +.0000306245 97,70 +.0000012907 156% +.0000003355 0,41% First 5 significant +00000001576 0.15% +00000000575 0,07;6 +.0000000348 +.0000000135 +.0000000082 +.0000000016 +.0000000013 -.0000000006 -,0000000004 -.0000000002 +00000000001 +.0000000000

Variability divided LI-145c, fiften _onents in group 13: A21 A37 R69 Ll .000177397 A)43 R52 L2 0000007939 0 111

A48 A71 R83 L3 .000000646i 9 A26 A36 A41 R47 VI_ .0000005108 9 A34 A53 A83 .198 A136 8178 L5 .0000001788 9 .A55 Al 05 A130 R153 L6 0000001 069 9 A36 A50 R55 L7 .0000000913 9 A20 A.25 R30 L8 .0000000547 9 Pt26 L9 .0000000374 9 R.22 L10 .0000000279 9 R12 L11 .0000000196 9 R12 L12 .0000000082 9 R13 L13 .000000000 9 RIO L14 00000000017 R19 Total variability = L15 .0000000001 181474.48 Q-1-9 112

+.0001779897 98.1% +.0000017969 1.070 +.0000006461 0.4% But all still +.0000005108 0.3% significant +.0000001788 0.1% +.0000001069 0.06% +.0000000913 +.0000000547 +.0000000374 +.0000000279 +.0000000196 +.0000000082 +.0000000047 +.0000000017 +.0000000001

In group A the first five components and in group B the first six components were found to be significant respectively. These significant com- ponents of both the groups accounted for c.99.9/07 of the total variability. Within each significant component again, only a few of the fifteen variables were found to be significant carrying high weighting. Below are given the components of group A and group B showing the value of weighting for each variable within each component: Group A components showing weighting:

1 -.0047919677 +.0137925816 +.1000000000 + scarious tip left (large) -.0762972793 - scarious tip left (small) -.0943590347 - scarious tip right (large) 113

+.0642038090 +.0058299896 +.0020000124 -.0008854628 +.0079535376 +.0690368312 -.0692184137 +.0479286073 +.0009933705 +.0001590087 2 -.0439169169 +.0744456177 + sepals breadth -.0499379748 +.1000000000 + scarious tip left (small) +.0505541259 -.0863749602 - scarious tip right (small) +.0219610215 -.0035831086 +.0095293261 +.0003321680 -.0197988538 +.0154970730 +.0351554730 -.0010313611 -.0017244030 3 -.0072084216 +.0271675249 -.0870978491 - scarious tip left (large) +.1000000000 + scarious tip left (small) +.0283853377 -.0608315661 -.0386261738 +.0020917903 +.0016101181. +.0009800060 +.0614843986 -.0585701302 +.0299060890 +.0136322326 .0442305362 114

-.0052901927 +.0141645999 -.0812103728 - scarious tip left (large) +.0607562754 +.0563138833 -.0420392873 +.0021521890 + .0119190179 -.0234997457 +.0010015801 -.0846451566 - seeds length +.1000000000 + seeds breadth +.0445775005 +.0005414051 -.0056869467 5 -.0061558707 -.0074561270 -.0448757714 -.0621894285 +.0585266715 +.1000000000 + scarious tip right (small) +.0073481917 +.0118696639 -.0054152144 -.0116650145 -.0463418923 +.0962459629 + seeds breadth -.0722652128 +.0019425702 -.0004860422 6 -.0017956571 -.0190008598 +.0505760965 +.1000000000 -.0482287188 -.0655671559 +.0155597592 +.0092089425 +.0087583834 -.0409957007 115

+.0375649183 +.1000000000 -.0210111779 -.0578535303 +.0479501289 +.0136040232 -.0025227858 -.0035355411 -.0062252794 -.0009067443 -.0187787524 7 -.0401186768 +.0373889190 -.0026296382 +.0322732424 +.0116303383 +.0022306673 +.1000000000 +.0041746435 -.0045311551 -.0965208868 10 -.0110450408 -.0306837806 -.0049832710 +.0011593932 +.0060338232 -.0046693115 -.0905461729 +.003Wi92415 +.0581160860 +.0584065344 +.1000000000 -.0817092182 -.0813481324 +.0177204200 +.0016671805 -.0008084014 -.0061971155 +.0046030338 -.0078343086 -.0172541765 8 -.0392346956 +.0292236191 +.0058205452 +.0486668265 +.0003382078 +.0027565573 -.0851826442 +.0064962945 -.0320358668 +.1000000000 11 +.0207293735 -.0075332490 +.0018139106 +.0028500932 -.0049157383 +.0024095805 +40529722783 -.0114112053 -.0898869691 -.0145152554 -.0506568231 +.0126846001 +.1000000000 -.0294854666 -.0018808469 -.0005981825 +.0032084073 -.0022880182 +.0006048502 +.0114209479 9 +.0287815418 -.0121788523 -.0039917707 -.0264529911 +.0025973654 -.00166801)14 -.0928822470 -.0047512256 +.0400481946 116

12 14 +.0031605976 +.0012638067 +.0080254196 -.0057838866 -.0275451832 +.0557516345 +.1000000000 -.0959633042 -.0041291998 -.0460616533 -.0976166635 +.1000000000 -.0130964537 +.0002986310 +.0046923176 +.0029114538 +.0018063390 +.0010464032 +.0042798933 +.0087972732 +.0022356171 +.0271117198 -.0231017081 -.0118990547 -.0120862125 -.0237105110 -.00105489H -.0013713138 +.0010717082 -.0047016476 13 15 +.0012897614 +.0026326929 -.0051689860 +.0080784969 +.0517312789 -.0264206664 -.0954961895 +.1000000000 -.0444557654 -.0033513415 +.1000000000 -.0982270647 -.0003893306 -.0125808857 +.0013982770 +.0048208922 +.0012218356 +.0012588949 +.0074295891 +.0042161388 +.0224569251 +.0012216017 -.0057968853 -.0227981938 -.0157952286 -.0123793013 -.0012110669 -.0008315358 -.0043671305 +.0013244098

Group B components showing weighting:

1 +.0050797883 +.0038933713 +.0536880504 -.0124153251 -.0333447946 +.0058040437

117 +.0210857145 +.0041271489 +.0049147787 -.0144602749 +.0831212143 + seeds length -.01 18775974 +.1000000000 + pollen grain diameter +.0003305809 +.0041061926 2 -.0394104601 +.0878886050 + sepal breadth +.0607978439 -.0055882765 -.0150371971 -.0116897335 -.0077420622 -.0109491592 +.0225187389 -.0108267594 -.0916987170 - seed breadth +.1000000000 + pollen grain diameter -.0090418573 +.0003497936 +.0112119255 3 +.0160470751 -.0560914119 -.09180L7661 - scarious tip left +.0074323810 +.1000000000 + scarious tip right -.0048656314 -.0050155939 -.0364767503 +.0503553459 +.0537735131 -.0506384273 +.0600018473 -.0109222694 +.0228600458 -.0527509426

118

+.0132679221 -.0313724904 -.0839199932 - scarious tip left +.0087409088 +.1000000000 + scarious tip right -.0063574319 -.0093487253 -.0000477890 -.0260056849 +.0424027916 -.0733640975 - seed length +.0373177421 -.0349402388 +.0016452362 +.0361354619 5 +.0147204601 -.0404677938 +.0648023560 +.0456749878 -.0540129006 +.0035105877 +.0038397720 -.0097848297 +.0254592307 -.0224000042 -.0617591352 +.1000000000 + pollen grain diameter -.0432245122 -.0013620867 +.0065308670 6 +.0036503001 -.0285350043 +.1000000000 + scarious tip left -.0180520809 -.0469024002 -.0233161222 +.0288862867 -.0138638670 +.0209966190 -.0437648365 119 +.0167304756 -.0007651242 -.0219525548 +.0163960481 +.0538505191 +.0111325845 -.0005443079 -.0142939365 +.0038009639 -.0113242616 -.0558614644 7 +.0615198990 +.0112403039 +.0130033057 +.0014547393 -.0147752705 -.0017018602 -.0143933638 +.0281947039 10 +.1000000000 -.0440274041 +.0030379336 -.0296210363 -.0041025193 -.0106796980 +.0456930403 +.0014338610 -.0999987534 +.0241566227 +.0290006108 -.0799630276 -.0051745845 +.0242175362 -.0256541467 +.0526873772 -.0045107767 -.0017035227 +.0072414265 -.0081637681 +.0289557336 -.0297136588 8 +.1000000000 -.0879866365 -.0050049763 -.0008994128 -.0026735153 +.0003777160 -.0408554791 -.0356251358 11 -.0189285346 +.0067764520 +.0026086478 +.1000000000 +.0103981981 +.0062885383 +.1000000000 +.0509037557 -.0392890250 -.0217793692 -.0924027164 -.0225698694 +.0067822098 -.0339549037 +.0112929252 -.0084318122 +.0005685602 -.0042041801 -.0000707266 +.0015987324 -.0240010712 +.0167938145 9 -.0212235032 -.0413620726 -.0059198124 +.0007264936 -.0004578072 -.0009132670 +.1000000000 -.0101984887 -.0808528838 120

12 1)4 -.0013754197 +.0032763998 -.0133103173 -.0018038015 -.0417635519 -.0972557)1)19 -.0360784465 -.0296558290 +.1000000000 +.1000000000 +.0117985747 +.0497599214 -.0081843572 +.0361128310 +.0049735666 -.0101165928 +.0027833084 -.0050019276 -.0105411905 -.0030970711 +.0155147572 -.0411211841 -.0202988019 +.0615021554 -.0041090804 +.0118999497 -.0001196980 -.0007704709 +.0006267525 -.0044505064 13 15

-.0005274585 -.0009613714 -.0080028949 -.0228364131 +.1000000000 -.0498323111 -.0045437099 -.0554842933 -.0751705193 +.1000000000 +.0096072023 +.0887194185 -.0058563251 +.0186836092 -.0027694315 -.0098729306 +.0011141328 -.0046912788 +.0255345863 +.0130697077 +.0294373409 -.0089857163 -.0391563952 +.0172104862 +.0044927186 +.0567149022 -.0008783665 -.0009105971 -.0011897973 -.0030237436 Group A The component I accounted for 97.7% variability. In this component, the variable scarious tip left (large) carried very highiweighting, with contrast to this the variables scarious tip left (small) and scarious tip right (large) carried significantly high-weighting. 121 The component II accounted for 1.56% variability In this component the variables sepal breadth and scarious tip left (small) carried high +weighting and in contrast the variable scarious tip right (small) carried high -weighting. The component III accounted for 0.41% vari- ability but was still significant. The variable scarious tip left (small) carried high +weighting and in contrast the variable scarious tip left (large) carried high -weighting. The component IV accounted for only 0.19% variability but was still significant. The vari- able seed breadth carried high +weighting and in contrast the variables scarious tip left (large) and seed length carried high -weighting. The component V showed only 0.07% variability but was still significant. In this component the variables scarious tip right (small) and the seed breadth carried high +weighting.

Group B The component I accounted for 98.1% of the 16W variability. The variables seed length and pollen grain diameter carried very high +weighting. The components II accounted for 1% variability. In this component the variables sepal breadth and 122 pollen grain diameter carried high +weighting and the variable seed breadth carried high -weighting. In component III which accounted for 0.4o variability but was still significant, the vari- able scarious tip right (large) carried high +weighting and in contrast the variable scarious tip left (large) showed high -weighting. The component IV accounted for 0.3% variability, but was still significant. The variable scarious tip right (large) carried high+weighting and in contrast the variables scarious tip left (large) and seed length carried high -weighting. The component V accounted for only 0.1% vari- ability but was still significant. The variable pollen grain diameter carried high +weighting. Lastly the component VI accounted for

-Only 0.06% variability but was still significant. The variable scarious tip left (large) carried high +weighting.

In the next step the canonical variates for each individual of the taxa in each of the five components in group A and in each of the six components in group B were calculated. The means of the canonical variates for each taxon in each 123 of the five components in group A and in each of the six components in group B were calculated. The means of the canonical variates for both the groups are given below:

Means of canonical variates for group A:

I II III IV P 16.405 -0.633 0.801 1a467 0.012 / 0.125 Q 16.045 -0.086 0.602 -0.709 -0.804 0.254 IR 15.756 -2.512 0.242 -0.632 -0.326 0.248 S 17.135 2.458 0.704 0.187 -0.202 -0.026 T 16.258 0.102 -1.696 -0.856 0.166 -0.457 U 16.801 4.186 -0.961 0.638 -0.200 -0.169 V 14.931 -2.879 -2.325 0.075 0.384 0.482 lh 16.199 -3.735 1.874 0.542 0.419 -0.812 X 17.071 2.687 2.015 -0.637 1.487 0.157

)1 VI component 124

Means of canonical variates for group B:

I II III IV V A 21.938 -0.585 -1.123 -1.570 0.971 70.135 B 22.224 -1.713 -0.180 -1.616 0.820 0.247 C 20.903 0.878 -0.164 -2.560 0.723 0.677 D 21.636 -5.626 -0.639 -0►376 -0.840 -1.540 E 23.624 -2.465 0.538 0.455 -0.202 0.819 F 24.268 -2.432 -1.115 1.471 0.116 0.122 G 23.403 -0.868 -0.451 -0.627 -1.762 0.652 H 23.614 -0.081 -0.984 0.734 -0.060 0.504 I 23.030 0.509 0.056 -0.250 0.501 -0.520 J 23.662 3.320 -1.477 -0.383 -0.454 -0.576 K 24.664 3.683 -1.028 0.911 0.288 -0.319 L 24.480 3.647 1.941 -0.692 -0.872 0.015 M 25.327 -0.449 4.032 0.325 0.582 -0.327 N 24.520 0.149 -0.669 2.363 0.745 0.455 0 24.828 -1.067 0.412 0.282 -0.640 -0.752

VI component 125 The values of the means of canonical variates of all the taxa in group A were then plotted taking two values at a time for each taxon, on the graph in the following combinations: I and II, I and III, I and IV and I and V. II and III, II and IV and II and V. The values of the means of canonical variates of all the taxa in group B were plotted in the same way as mentioned for group A in the following combina- tions: I and II, I and III, I and IV, I and V, and I and VI.

The graphical respresentation of all these taxa in group A and in group B in various combinations shaved the degree of relationship of the taxa within the group. The taxa which are closely related would group together. The taxa which are distantly related would fall wide. The taxa in the group A are delimited in three sub-groups, given below.

A' includes taxa V,R and W A" includes taxa T,P and Q Aft includes taxa U,S and X

The group A has been considered to fall within the limits of C. album PhAst 126

GROUP A

VALUES OF CANONICAL VARIATES 19 1 -II COMPONENTS.

s

S 17 D V' 0

P A ❑1.4 AT G. 16 A R 0

15 01/

14

CI - Sub-youp A'

A - Sub-croup A"

0 - Sub-croup A*

13 2. 4 5 - 4 -3 -z q- 0 3

FIG. 28 127

GROUP A

VALUES Of CANONICAL VARIATES 19

I— DI COMPONENTS.

OS 17 ox 0

o w

16 R 0

15 V 0

14

CI — Sub-group

— Sub-group A"

- Sub-group

13 2. 4 5 -4 -3 -2. -1 3

FIG. 29 128

GROUP A

VALUES OF CANONICAL VARIATES 19

I—ISZ COMPONENTS.

18

S 0 17 X 0 0 u

& P T a 0 w a 6. El R

15

14

CI — Sub-you? A'

LI — Sub croup A'

0 — Sub-sroup A'''

13

-4 -3 -2 -1. 0 I. 2. 3 4 5

a FIG. 30

129

GROUP A

VALUES OF CANONICAL VARIATES 19

I -"SE COMPONENTS.

141

S 0 11 ox

u 0

a 16 A

R 0

15 1:1 V

14

0 - Sub-irour A'

A - Sub-youp Au

0 - Sublroup A"'

13 - 4 -3 -z -1 0 1 2. 3 4 S. "3 FIG. 31 130

GROUP A

VALUES OF CANONICAL VARIATES s II — III COMPONENTS.

U 4 CD

3 0" OS

2.

If

1

T A 0

4 P

-1

-2

0 R c3 v -3

W 0 0 - Sub-sroup A. -4 .O. - Sub-sroup A' 0 - Sublroup A"

-5 -3 2 3 -2 -1. 0 1. in FIG. 32 131

GROUP A VALUES or CANONICAL VARIATES 5

13.-33I COMPONENTS. u 4 0

x 0 OS 2

DT 0 1>%

_2

R 0 3v -3

ow El - Sub-group A' - Sub-youp A" - Sub-youp A"' -5 -3 - 0 1 3

FIG. 33 a 132

GROUP A

VALUES Of CANONICAL VARIATE 5 5"

II — = COMPONENTS. ll 4 0

3 x S 0 0

2

11 t

- A r 0 a &

AP

-1

-2

R 0

0 V -3

W 0 CI — St.kb-youp A' -4 ,6, — Sub-croup A"

0 — Sub-group Au'

—5 3 —3 —2. —1 O 1 2. FIG. 34 133

GROUP B VALUES OF CANONICAL VARIATES 26

1 -II COMPONENTS.

M zs

K A N AL F 0 2.4

E H 0 A

GO

23

B 0

A 22. O

21 O C CI - C. laricifollum A - C. ri9oi

(;) - C. G20r110411

• - C. matrense

20 2 4. -5 -4 -3 -2. -1 0 1. 3 IL FIG. 35 134

GROUP B VALUES OF CANONICAL VARIATES 24.

I-131 COMPONENTS.

. M

25

„K N LAL

oF

24

E 7 L> c31 0 I Oct

I 23 D

B O 22 A O

ID

21 - C. lariciiolw.m

- C. rivi

- C. ISomasii • - C. matrenst

20 -5 -4 -3 -2 -1 0 1 2. 3 4 5 III FIG. 36 135

GROUP B VALUES OF CANONICAL VARIATES zb

1.-14 COMPONENTS.

M

25 0

A K L . N

0F 24

T v 0°H 06

I 23 0

B 0 2.2 0 A

21 C CI - C. laricif awn 0 — C. riipi

— C. 6omcxsii

• — C. matrans

- 4 -3 -2 0 2. Ig FIG. 37 136

GROUP B VALUES OF CANONICAL VARIATES

I COMPONENTS.

M

25 0

A L N .66

0 F 2.4

60E0 I co

I 2.3 a

oa

2.2 0 A

0

2.1 C 0 — C. lo.ricifoltom — C. rigoi — C. Momosii • — C. mo.trense_

2.0

-5" -4 -3 -2. 0 1 2 3 4 5-

FIG. 38 137

GROUP B VALUES OF CANONICAL VARIATES 26 I-M. COMPONENTS.

11

2.5 0 AK . N

OF

2.4

7 A H 00 E

G

23 0

B 0 22 m. A

B

- C. loricif oltum 21 O C - C. M9o1

- C. I7•iomnsii

• - C. mo.trense

20 -4 -3 -2. 0 1 2 3 4 5 Sa. FIG. 39 138 The taxa in group B are delimited in four main sub-groups and each sub-group has been considered to fall within the limit of species as follows:-

I C. laricifolium Vill. includes the taxa A.,,B,C,D and I. II C. thomasii Ten. includes L,F,G and H. Porta III C. rigoi Huter et/ includes J,K and L. IV C. matrense Kit. includes M,N and 0. 138a Nomenclature By means of the canonical variates it can be shown that the Cerastium population on Mount Maiella consist of 24 taxa. Also an unknown in- dividual can be correctly assigned to its relevant group by means of an "identification matrix" fed into the computer. Nevertheless it is not possible to provide a conventional key to separate each of these taxa. It was decided therefore that entities cap- able of conventional keying only should be recognised as far as nomenclature is concerned. In the des- criptions (see pages 34 - 75) it has been made clear that the species recognised show considerable varia- tion and that there is some degree of discontinuity in this variation coinciding with differences of habitat and ecological niche but not of sufficient degree to merit the creation of contained subspecies. Accordingly it was proposed that the fol- lowing species should be recognised to contain the taxa referred to throughout this work as 'A', 'B', etc. I Cerastium album Presl contains taxa P, Q,R,S,T,U,V,W, and X. These taxa (of group A) may be grouped according to the canonical variates t into three sub groups A , A , and A- corresponding

13 8b

with the following habitats subgroup A (R, V, and w) is found on the lower slopes of Mount Maiella. subgroup AI Q, and T) is found growing in a .1 III habitat intermediate to A and A . Ii subgroup A (S, U and X) is found just below the shallow cliffs at the edge of the altipiano. These above subgroups are never found on the alti- piano proper and only in sheltered habitats at the higher altitudes. These three subgroups have not been given any nomenclatural status since': it is not possible to separate them on conventional methods. Despite the facts that the canonical variates clearly demonstrate their existence. II C. thomasii Ten.-This contains taxa E, F, G and H. These are found in exposed parts of the altipiano and at the edges of the cliff tops. III C. laricifolium Vill.-This contains Taxa A, B, C, D and I and forms a fairly homogenous group whose habitat is generally in crevices often fairly sheltered in the cliffs at the edge of the altipiano. IV C, rigoi Huter et Porta-This contains Taxa J, K and L and is again a fairly homogenous group. 138c

tat Its habyis at the base of the altipiano cliffs. V C. matrense Kit. This contains the taxa M, N and O. These taxa we in many respects inter- mediate in characters between the C. album group and the C.arvense sensu latu group. It is also one of the groups showing the highest degrees of poly- ploidy and is the only C. arvense sensu latu group which cohabits with the C. album group. It is found below the cliffs of the altipiano and extends a little way down into the valleys below which lead off from the altipiano. One population was found at least 1000 ft. lower down the mountain from the edge of the altipiano and in the middle of the al- titude range of the main population of C.album. It has not yet been possible to see the type specimens of the species whose names have been emp- loyed above. The choice of these names results from extensive examination of the collections at the British Museum of Natural History, and Kew Herbarium together with the appropriate critical floras. Some of these names do not appear in the Flora Europeae account of Cerastium. It is suggested that their names are the best possible provisional ones but may need amendment after their types have been seen. 139

DISCUSSION

Although the oldest, taxonomy is one of the most neglected disciplines in biology. The review of on literature/systematics shows that the aims, and co/17. cepts of classification have gradually changed from artificial to natural systems. This change in out- look and methods has developed parallel with the improvement in our techniques and ideas. New techniques are being developed in studying living organisms, in discovering new characters, in describing new species and revising the systematics of previously known organisms. The advances in cytology, genetics, biochemistry and geographical distribution have provided a wealth of material thereby opening a way for new systematics. Much of this has been used for phylogenetic interpretation. It has not always been used in developing a natural system of classification which could have more objec- tivity. As a matter of fact very little work has been done since the 19th century to explain what should be the conceptual basis of classification. In classification4 the delimitation of taxa is based upon discontinuity, however small and artificial it may be. This practice of taxonomy 140 was based mainly upon intuition or neural perception of such discontinuity which is an art rather than a science. This intuition varies from worker to worker, taking a definite example - taxonomists have never come to any clearcut and defined conclusion as to where the true limit of sipecies should lie in a particular group especially where texa of the problem" or "critical" group are involved. The same group of plants have often been treated taxono- mically in many different ways. During the last few years there has been tremen- dous awareness of the problems in the techniques and principles of taxonomy. Some of the noteworthy and referable publications (see Cain 1956, 1958, 1959a, b, c; Cain and Harrison 1958; Gilmour 1937, 1940, 1951,1958,1961b Gilmour and Walters 1964) that have appeared in recent years have tried to re-eva- luate the logical basis of taxonomy. This has involved the separation of various functions which the science is trying to fulfill. A critical account of the shpaing of angiosperm taxonomy by S.M. Walters (1961) has drawn attention to the early historical and philosophical background. The importance of these factors cannot be overlooked, and there can be no doubt that angiosperm classi- 141 fiction would look rather different if Linnaeus had lived in far east and not in Sweden. Critical accounts on angiosperm taxonomy from Linnaean and post-Linnaean era up to the present can be seen in Heslop-Harrison (1960)9 Davis and Heywood (1963) and others. Turrill (1938a) pointed out that our "alpha" taxonomy is in a healthy and profound state of change. He could visualize the future form under the impact of experimental findings. This he termed as "omega" taxonomy. In the assimilation of the data in taxonomy, it has been usually pointed out that the question is not only of assimilation of experimental data to fit in a hierarchy of nomenclature, but to establish a certain defined reference system (see Erna Bennett 1964). This can be stressed more so in the defini- tion of species in face of new experimental data coming to light. Phenetic classification according to Davis and Heywood (1963) and Sokal and Sneath (19e) is the main aim of taxonomy. This concept of phenetic classification in all respects coincides with the natural classification of the post-Darwinian era which is based upon the correlation of maximum number 142 of attributes. However the phenetic classification may or may not coincide with the phylogenetic system. It is suggested that the phylogenetic classification cannot serve the purpose of natural classification for in most of the cases the phylogeny is unknown and even if it is known we do not have the complete fossil records. Heslop-Harrison (1964) has called it a "romantic speculation" which is long overdue for abandonment. The main aim of experimentation is to understand the taxonomic units rather than to define taxa (see Gregor 1963). Whether a few or all of the characters are ith- portant depends upon the type of problem which we are dealing with in taxonomy. Those with deeper acquaintance with the principles of evolution would appreciate that it is a multidimensional process. The survival value of plants in nature depends upon their adaptability in relation to selection pressure, It has been, therefore, difficult to arrive at a precise definition of species. The species concept has a long involved history behind it. In the pre-Darwinian era it was gener- ally accepted that different kinds of organisms have rigid or divine creation. Linnaeus in his earlier days accepted the idea of divine creation of species, but towards the cloSing years of his life, 143

experiments on hybridisation led him to revise his earlier concepts. He proposed an evolutionary con- cept of the origin of species (Linnaeus 1774): So great was the impact of publication of his sexual system and binomial system of nomenclature that the beginnings of the difficulty of species definition can be traced back to his time. Greater attention was given on the arguments for and against the sexual system by his contemporaries and its extension and application by his successors. As a result of which the later concept of Linnaeus and even the experimental work on interspecific hybrids by Koel- reuter (1761-1766) was overlooked, and as a matter of fact very little challenge to the concept of divine creation of species was offered until about the mid-nineteenth century. The next step in the concept of species was the observations of Alex Jordon (1846) a French botanist. He showed by experiments that several races of one Viola species of Linnaeus collected from different localities re- mained distinct when cultured under standardized conditions. He interpreted each race as a separate species, but his ideas were opposed by his contempo- raries mainly because this would have led to an 144_

immense multiplication of specific names. The publication of the 'Origin of Species' by Charles.

Darwin (1859) changed the entire outlook of the bio- logical thoughts at that time by giving rise to the importance of phylogenetic studies. The phylogenetic studies are mainly philosophic and speculative and based on deduction or comparative

observations rather than on experimentation. The development of genetics right from the beginning of the 20th century further led to much discussion

about the species problem. Larly geneticists sought the origin of species in single mutations. With the advances in population genetics, many geneticists consider that speciation is a slow pro- cess of population drift aided by mutation, selection and isolation resulting into several forms having a degree of qualitative distinctness. By the post- Darwinian era, on the other hand the morphological species concept had become much refined. It re- cognizes variations and tries to contain them. As has been referred to at appropriate places in this thesis most of the investigations in experimental taxonomy revolve at or around assessment of the objectivity of taxa at the specific or infraspecific level. As to the concept of species as taxonomic 145 and biological unit there have been always varied. and various opinions. Whether taxonomic units should coincide with biological units or biological units should be fitted into taxonomic hierarchy. As pointed out by Bennett (1964) our biological units when fitted into taxonomic hierarchy are looked upon as something which haiephylogenetic significance. There are people who argue that the biological classi- fiction, although important in its own way should not interfere with the taxonomic classifications. Where taxonomic coincides the biological concept of species there is no difficulty, but difficulty arises when we come across groups which form morphological clusters despite their biotype distinctness. Some would ad- vocate a synthetic taxonomy where the units in the related group of plants should be assessed not only on their morphological attributes but also upon other intrinsic potentialities (Turrii11942, Heslop- Harrison 1959). The concept of species actually is a challenge to the biologists and it has been debated from time to time without any definite conclusion. What actually constitutes a species? Whether they are created by nature or nurture? Whether they exist in the mind of taxonomist or whether they are 146 units whose objectivity can really be assessed by the sophisticated experimental techniques? To elucidate these and associated conceptual conflicts a symposium was held in 1957 (see Mayr). Several definitions of species have been proposed (see Mayr loc. cit. & Beaudry 1960). The twc main classes are as follows: a) taxonomic (including the orthodox, typological, morphological, morphogeo- graphical, etc.) b) biological (including the bio- systematic, genetical, cytogenetical, non-dimensional, multidimensional, etc.) However most of the practising taxonomists have employed the first concept. Since the actual, though not always acknowledged, justification for classifi- cation is that it imposes order on a naturally dis- orderly array, these specific bounds should be placed so that disorder is at a minimum, that is at the point of greatest discontinuity. It is obviously impractical to use genetic criteria for the delimitation of species for it demands considerable long time for experimental work to know the genetic constitution. The same objection may be raised at the biometric procedure advocated in this thesis that this method does not give a readily available identification matrix for 147 which a field taxonomist may look for his ready re- ference. The, aim of the present taxonomic method is not only to aid in a quick and precise allocation of herbarium specimen to their respective taxa in terms of non-dimensional units, but also to find out whether all, one or a few attributes are the deter- minants in shaping of taxa. In the course of the experimentation, however, it was discovered that these taxa in terms of variation reveal attributes which deserve differential weighting. Such deter- minations have the advantage of being unaffected by personal prejudices and intuitions which differ from worker to worker. When morphological differences are to be employed it must be borne in mind that these are relatively unaffected due to variation in the environmental con- ditions. This will need experimentation and study of variation preferably in biometric terms. The present taxa though apparently morphologi- cally and cytologically related, do not fit into a straight-jacket'. They are all local collections from Mount Maiella, Italy, and therefore their re- lationship had to be looked into from the point of view of their morphological expressions in different ecological niches. The selection of morphological 148 characters in relation to their adaptive values is of a paramount importance in taxonometrics. It is not the characters as a matter of fact, but their states or expression which is more meaningful. As explained earlier in Chapter V on page 94 care was taken in selection of characters (variables) to be employed in the computational work. Only those characters were employed which were considered as suitable attributes from the point of view of their plastic range. Biometric methods because of their exact nature can be of greater help in making the taxonomy more objective. These methods should be looked upon as useful tools in the hands of taxonomists, rather than a universal answer to all taxonomic problems. It must be pointed out that all the problems are not suited to treatment by this method and in fact many problems do not require such complex labour for their solution. The present method is most suitable for the solution of the problem presented by 'critical groups' like the one under investigation. It can disclose the existence of minor grouping, their relationship to the main groups and above all, it can give a reliable objective estimate of the degree of resemblance between and within groups.

149

All characters employed did not appear to have equal value in their contribution towards total variability either within or between taxa. This further shows that to give equal weight to all characters will give incorrect values for the delimi- tation of taxa. As mentioned in Chapter V the twenty four taxa were subjected to biometrical analysis using fifteen variables. The method measured the"generalised 'distance" between all the preceding taxa, and has sorted out five major groups with minor sub-groups included in them. This generalised distance be- tween each major group with sub-group within them, each has been graphically represented in figs. 28-39 on pp. 126-137. The five main groups have been taken as five species which with infraspecific clines are noted below: -

I. Cerastium laricifolium Vill. including the taxa A,B,C,D, and. I.

II. C. thomasiiTen. E,F,G and H. et Porta III. C. rigoiHuter/ J, K and L. IV. C. matrense Kit. M, N and 0. V. C. album Presl P,Q,R,S,T,U,V0'ii, and X.

C. album includes all the taxa which have been referred to es group A. 150

The other four species, namely C. laricifolium, C. thomasii, C. rigoi and C. matrerIse include all the taxa which have been hitherto referred as group B. In order to test the validity and correctness of the method employed these specific taxa were mat- ched with material bearing names in Herbarium British Museum and Kew Herbarium. It was found that these taxa in general facies correspond to the materials mentioned above but in their greater details they differ to an extent as may call for further infra- specific treatment. However, this question was left open for further confirmation. As mentioned earlier all the taxa were collected from Mount Maiella and whether they are sympatric or allopatric is still a matter of detailed morpho- geographical investigation. So far as the taxa included in C. album are concerned, these appear to be true distinct lumps including P,Q, and T in one group, R,V and W in an- other group and S,U and X in the third group. At the first look one is tempted to treat them either as different species or sub-species. 151

To test the validity of the findings only the cytological investigations (mitotic counts) were carried out. Our intended programme of cytogenetical investigation in relation to taxonomy could not be done due to the failure of the plants to produce flowers under the standard conditions of the green- house as well as in the field. So our cytological studies find much limitation in their approach to this problem. The chromosomes being dot like and short rods could not clearly show either centromere or the arms length in the karyotype. Our karyo- typic investigations were mainly based on the somatic counts. However these somatic counts could give valuable information which could provide as a check to our biometrical findings.

"Chromosome tree" GROUP A (Cerastium album)

A ►►

2n=72 2n=72 2n=36 113 2n=72 2n=63 2n=72 2n=72 2n=72 2n=72

152

Vrom the tree shown above it can be seen that in this species we have three 2n chromosome numbers 36, 63 and 72. However in the sub-group A' above, all the taxa have 2n = 72, in the group A" 2n = 36 and 72, and in the sub-group A''' 2n = 63 and 72. It therefore can be inferred that the group I, (Co album) shows a polyploid series ranging from tetraploid to octoploid. The taxa with the same somatic number have tendency to form separate clusters on the graph and it may be probable that these taxa with same chromosome number, may have different genic balance due to the introgressive hybridization. The cyto- taxonomical data did not appear to be conclusive for treatment of the three sub-groups as different species or sub-species. For having said so detailed meiotic investigations in conjunction with morpho- geographical studies (as pointed out before) would be necessary. "Chromosome tree" GROUP B

C. laricifolium C. thmasii C. Ligoi C. matre'ase ! I ; ! , !1 ir I 1 L 6 t_ ) , i 3F a I., H i r\ =3( :•?• • ':R.T -- - 7), 7_5 t :fv.. t .1., .77.). 153

An examination of the chromosome tree of the Group B indicates that the chromosome number alone cannot be said to be sufficient or conclusive in itself for the delimitation of the species. That these species form polyploid series is quite clear. The specific delimitation however is based on boi- metric analysis in conjunction with somatic chromo- some number. So far the taxa within C. laricifolium are concerned they all show the same somatic chromo- some number. The morphological variability is to the extent which permits their treatment in one specific limit. The somatic chromosome number in C. thomasii ranges from 2n = 36 to 2n = 5L and it can be con- cluded that cytological mode of speciation is more active in this group. In the species C. rigoi the somatic chromosome number ranges from 2n = 36 to 2n = 126. It can be concluded that polyploidy has been most active in this group. C. matreAse however is the group characterized by highest somatic chromosome number which indicates that polyploidy has been most active in this group. 154 The biometric findings of all the taxa under in- vestigation indicate that the taxa of this 'critical group' which morphologically appear to be related can be delimited on the basis of the biometrical method employed in this investigation. These find- ings should aid in understanding the mechanism of evolution together with the information supplied from other disciplines of studies. As pointed out earlier the aim of experimental taxonomy is more to understand and investigate the causes of variations rather than using the experimental data for taxo- nomic delimitation. 155

CYTOLOGICAL DISCUSSION Basic chromosome number in the genus Cerastium 4.1.•••••••••,*0 The genus Cerastium belongs to the family Caryophyllaceae. It is a very large genus. The chromosome numbers of many spp. from various localities have been determined by various workers (see cytological review). The somatic chromosome numbers were deter- mined of the 21 taxa under investigation. These were found to be all multiples of n = 9 (18, 36, 54 and 63). The cytological observations of the taxa showed many degrees of polyploidy, the 2n chromosome number ranging from 2n = 36, 2n = 54, 2n = 63, 2n = 72, 2n = 108 to 2n = 126. All the taxa were found to be fertile; their seed viability and germination percentage was very high. It thus appears that the basic chromosome number for Cerastium is nine, although the lowest chromosome number so far known is n = 18 and the plants having chromosome numbers 2n = 36 are perhaps tetraploids. Rohweder (1939) and Hagerup (1944) also have suggested that nine may be the basic chromosome number for the genus Cerastium. 156

Brett (1952) has also indicated that nine is the basic. chromosome number- for Cerastium from the mitotic preparations of C. semidecanderum where she found four nucleolar organizing chromosomes, each with a satellite. Although most spp. whose chromosome numbers have been reported, come in the n=9 group directly, there are a few spp. in this genus said to have n=17 and n=19. Brett (1952) has suggested two basic numbers i.e. nine and nine- teen. LOve and LOve (1961) have given three basic numbers; 9, 17 and 19 for example spp. with 2n=3Li_ have basic number n=17 and spp. with 2n=38 have basic number 19. Brett (1952) in support of 19 being another basic number, has reported 19 bivalents and two nucleolar-organizing chromosomes clearly visible during prophase and claims that the plants are truely diploids with 19 as the basic, number. About their origin she has suggested that n=19 group has a polyphyletia origin and that the app. comprising it have arisen possibly as a result of amphidiploidy between a dipoid species with n=9 and one possibly from another genus or from a putative extinct 157 section of Cerastium with n = 10. This conclusion be does not appear to/the most likely hypothesis and simpler origins can be postulated. The groups with n = 17 and n = 19 might have arisen by non-disjunction (a phenomenon of aneuploidy); a suggestion already pro- posed by Soliner (1)54). Brett (1952) herself included three species C. cerastuides 2n = 34, 36 and 38, C. arvense 2n = 36, 38 and 72, and C. tomentosum 2n = 36, 38 and 72, to the n = 9 group, and she was of the opinion that the races of these species with n = 17 end n = 19 are as a result of aneuploidy. The genus Cerastium, represents many degrees of polyploidy and so far no diploid is reported. .It is suggested here that some future investigations may show either some diploid species with 2n = 18 in this fenus from some other localities which have not yet been explored. Usually the poly- ploids have different habitats than their diploid ancestors. If no plant with 2n = 18 is reported then there may be a possibility that amphidiploidy might have also played a major role in the diploids containing two more or less non-homolous chromosome sets AB; had its chromosome number doubled and it 158

gave rise to an allototraploid (amphidiploid; of constitution AABB. The amphidiploidsonce formed, the conditions would tae favourable to their continued survival, for amphidiploids are according to some authors to be more adaptable and better able to withstand competition than their diploid relatives, and thus in the course of time the diploid species in the genus Cerastium became extinct and replaced by amphidiploid species with 2n=36. It is there- fore suggested here that the diploid species could not survive and become extinct and as a result only the polyploids have survived to re- present the genus Cerastium. The few species of the genus which have n=17 and n=19 may all have been formed by non-disjunction. It is also possible that further cytological investigations of these few species from other localities may reveal plants with n=18 as it occurred in the case of C, cerastoides, C. arvense and C. tomentosum. It is therefore proposed that it is more likely that the genus has 9 as a basic chromosome number and not 9, 17 and 19. It is proposed that the species with n=19 and n=17 are possibly the end product of non-disjunction a case of aneuploidy. 159

Polyploidy Polyploidy appears to occur in all groups of plants with the possible exception of the fungi. About 50% of the angiosperms are said to be polyploids. The estimate of 5c% polyploidy among angiosperms is not very precise, as relati- vely few species have been critically examined. Some cytologists, however, estimate the percent- age ranging between 30 - 35. Most wild poly- ploid plants are alloployploids or sometimes com- plexes of al20-and autopolyploids. These may reach higher degrees of polyploidy; tend to have greater distributions than their diploid ancesters; and usually have different habitats than their diploid ancesters, and higher poly- ploids are usually found in the perennial entities. There is also a corelation between polyploidy, latitude and glaciation. The creation of new and open habitat as a result of glaciation would appear to be the main reason for these regions having high ratio of polyploid entities. It is also possible that other factors which lead to the creation of open or varied habitats will also be found to favour polyploidy. 160 The genus Cerastium shows high degrees of polyploidy. The highest polyploid in this genus reported so far by Blackburn and Morton (1957) in C.vulatum (triviale) where 2n = 180. Eric Hulten (1956) has shown and admirably discussed the introgressive hybridization in C.alpinum complex. This taxonomic account shows that C. alpinum group forms a chain of species connected not only by hy- brids but also by hybrid swarms, dominating the Cerastium flora. The 24 taxa of this genus Cerastium under investigation also show high degrees of polyploidy. An analysis of the taxa shows that:- Nine taxa (A,B,C,D,E,G,I,K and T) are tetraploids 2n = 36. One taxon (F) is a hexaploid with 2n = 54. One taxon (U) is a septaploid with 2n = 63. Eight taxa (N,P,Q,R,S,V,W and X) are octoploids with 2n = 72. One taxon (M) is a 12 ploid with 2n = 108. One taxon (L) is a 14 ploid with 2n = 126. Since the formation of polyploids in Cerastium has played a major role in the evolution of this genus, the question arises what type of polyploidy has taken place? From the information supplied by various 161

workers who have raised artificial autopolyploids, the followinw, ways 'of distinguishing betwen auto- polyploids and allopolyploids may be considered. (1) By studying the external morphology of the chromosomes to see if there are more than two sets of chromosomes morphologically similar. But there may be a possibility that false conclu- sions may be drawn. Besides this, in Cerastium species the chromosomes are too small to study their morphology. (2) Polysomic inheritance has been held to show autopolyploidy; but it is not possible to apply this method to high polyploids. This would also not necessarily distinguish autopolyploids from segmental allopolyploids.

(3) Generally reduced fertility has been found in artificially produced autopolyploids. It is therefore possible that a degree of sterility may be due to autopolyploidy. But such con- clusions must be drawn very carefully, for steri- lity may be due to some other causes norrelated to polyploidy. However in Cerastium taxa 162 under study, th, sterility was not observed et all and certainly there was no cytological evi- dence of this so far. (4) The presence of multivalents during meiosis is an indication of autopolyploidy, since the multivalents are very rare and when they do occur, they can be made out with difficulty. However Brett (1952) has reported multivalents in three species with high chromosome number, C. holo- steoides 2n = 144, C. arcticum 2n = 108 and C. pumilum 2n = 90,95. There may be a possi- bility that autopolyploidy might have played the role in the evolution of the above mentioned three species. But this possibility may also not hold good, for, if the two diploid species are closely related, the amphidiploid may form multivalent as a result of homology between pairs of the chromosomes of the two species es- pecially when the two species are from the same genetic stock as for the x = 9 species and have chromosome numbers which are multiple of nine. This phenomenon of multivalent formation was reported in Crepis by Poole (1931). He found that all the associations could be tetra- valents in amphidiploids. 163 (5) This is the most obvious and perhaps the most used method for distinguishing autopoly- ploids. Usually the autopolyploids resemble closely with the diploid parents so if a plant is found to resemble closely to another with half its chromosome number then it is generally said to be an autopolyploid race. It may, therefore, be quite possible that autopolyploidy may have taken place in the formation of intra- specific races of the genus Cernstium / C. album 2n = 36, 63 and 72; C. rigoi 2n = 36 and 126; C. matrense 2n = 72 and 108; C. thomusii 2n = 36 and 54; C. arvense 2n = 36 and 72; C. alpinum 2n = 54, 72, 108 and 144; C. tetrandrum 2n = 36 and 72; C. pumilum 2n = 54 and 90, etc.';, for the higher chromosome numbers resemble the lower chromosome races closely and may have a similar distribution. But even in these intraspecific races it cannot be assumed with certainty that these races are autopolyploids. There is certain amount of circumstantial evidence suggesting that these are not true autopolyploids and per- haps they may not be even intraspecific allo- polyploids. As more information is being 164 accumulated concerning the polyploid origin of the plant species it is becoming more clear that autopolyploidy is not a ccAmon phenomenan. The plants which were analysed as autopolyploids when subjected to further analysis have proved to be allopolyploids. Clausen, Keck and Hiesey (1945) working on Media species, showed that a species with three chromosome races, all of which resembled each other morphologically very closely were actually derived by hybridization and polyploidy between at least three diploid spedies. This demonstrates strongly against assuming autopoly- ploidy. Autciplyploids are less adaptable to changing condition on account of their lower mutation rate. Haldane (1930) hag', reported that the chance of a tetraploid acquiring homozygosity for a set of four allelomorphs is much less for a hexaploid and almost none for higher polyploida. 165

Lt'ia, therefore, suggested that probably allo- polyploidy has played the main part in the evo- lution of Cerastium pirticularly since multiva- lents are very rare. There are also many tetraploid species with 2n = 36. They mor- phologically differ from each other, indicating that they might have been evolved as a result of amphidiploidy amongst the variable group of diploids. It also seems probable that aneuploidy has also played a part in the evolution of this genus. The species, whose 2n chromosome num- ber is not multiple of nine (see basic chromosome number page l55), are probably formed as a result of aneuploidy. Intraspecific races also might have formed in some species because of the aneuploidy. For example C. holosteoides has three races with 2n = 34, 36 and 38. The race with 2n = 38 is most wide spread in Europe and the races with 2n = 34 and 36 have restricted distribution. This also suggests that the group with 2n = 38 is a polyploid and not diploid. If these species would have diploids then it was highly improbable for these species 166 to survive with chromosome missing. It there- fore canbe concluded that polyploidy has played a very important part in the evolution of this genus. This may be mainly due to the allo- polyploidy and to some extent to aneuploidy. 167

ACKNOWLEDGEMENTS

It is with great pleasure I express my gratitude to my supervisor, Dr. F. H. Whitehead for his stimulating interest, supervision, suggestions and healthy criticism throughout the course of this investigation; Dr. R. E. Blackith of Zoology Department, Imperial College and Mr. J. N. R. Jeffers of Forestry Research Commission for helping me with the statistical analysis; Mr. P. Sell of Botany School, Cambridge for helping me in the identifica- tion of Cerastium taxa; Dr. A. Melderis and Keeper of herbarium of N.H.B.M. for the permission to use the library and herbarium; and the Director and Keeper of herbarium Kew for permission to use the library and herbarium.

I should like to thank all those who have helped me in the preparation of this thesis; Mr. A. Horne for the photographic work; Mrs. M. G. Peterson, Miss M. Wilkins, Miss A. Leathers and Miss F. Mawani for typing the thesis; Messrs. R. P. Sinha 168

and M. Hasan for help in various ways; Messrs. J. Bunning and R. Adams for occasional assistance in the lab. I should like to thank Mrs. F. Baig for her help and encouragement throughout the course of this work. Finally, I should like to thank Professor W. 0. James and Dr. F. H. Whitehead for tho generous financial support. 169

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APPENDIX Taxon A

a b c d e f g h i j k 1 Ea

Aa 65.4 25.2 10.( 6.0 9.0 5.0 8.0 42.5 20.5 8.0 14.1 12.0 9.4 59.5 18.8 Ab 69.2 29.6 7.0 3.0 7.0 203 802 47.0 23.0 9.5 17.0 13.0 10.6 73.9 23.4 Ac 70.2 28.0 9.6 5.0 9.0 5.3 5.0 LC.0 19.5 5.0 14.5 11.7 9.1 75.7 23.2 Ad 84.0 31.0 IC.3 5.5 9.5 5.3 8.0 50.0 28.0 9.0 15.0 12.c 9.6 68.2 23.5 Ae 79.0 29.4 10.0 6.0 9.5 5.6 4.4 51.5 26.5 4.0 17.6 11.7 9.2 70.7 26.2 Af 80.2 34.8 9.0 3.0 9.0 2.5 10.8 149.0 27.0 11.0 16.0 12.4 10.3 64.5 19.6 4,g. 65.2 25.0 9.0 7.0 8.0 6.0 7.-1; 4a,.3 23.0 6.8 15.5 10.5 iC.0 77.8 22.0 Ah 73.0 31.2 11.c 7.5 lo.o 7.o 6.8 51.0 24.5 4.5 15.0 12.0 9.14 70.9 20.8 Ai 78.8 32.6 12.0 5.5 11.0 5.3 13.0 56.0 25.0 11.5 15.c 11.0 10.8 67.5 22.8

j 80.2 35.0 8.5 3.3 9.0 3.0 10.0 4500 22.0 10.0 15.0 11.6 10.3 74.0 23.0 Ak 7716 31.0 10.3 3.0 10.0 2.3 12.0 53.8 25.2 11.2 17.0 13.3 10.9 71.6 20.3 Al 77.6 31.0 10.0 7.0 9.5 7.0 3.8 56.5 27.0 7.0 14.8 11.0 10.0 76.4 22.7 im 83.6 34.4 11.6 9.6 11.3 8.3 12.0 59.6 30.0 12.6 15.7 12.4 9.3 90.4 25.6 Continuation

Taxon A

a 1

An 67.8 26.0 8.5 5.0 8.3 4.5 9.0 58.0 28.5 8.0 16.3 13.1 9.8 70.1 20.6 Ao 72.2 28.5 9.6 5.0 9.0 5.0 7.0 hi .0 20.0 6.0 1 5.0 i 2.8 10.0 73.0 2)4.0 Ap 78.0 30.0 1 0 .0 5.0 IC.0 5.0 15.0 ;3.c 28.0 9.0 14.5 1 2.0 9.5 66.5 20.0 Aa 70.2 30.0 7.3 3.3 7.0 2.5 8.0 46.5 23.0 9.3 16.5 13.C 10.5 71.0 23.0 Ar 75.0 32.1 1C .0 7.0 1 C.0 7.0 7.3 .6 25.0 6.0 1 5.2 1 2.1 9.7 68.0 21 .0 As 78.4 30.2 1L.5 7.0 11.0 7.5 7o0 7.3 23.0 9.0 16.2 12.6 10.3 79.2 25.0 At 66.4 25.2 9.0 11.0 8.5 3.5 6.0 45.6 22.3 7.3 15.0 11.0 10.0 71.2 20.2 Au Av Aw Ax Taxon

a b c d e f g h i j k 1 In

Ba 91.0 34.2 11.6 3.0 11.0 3.0 11.0 45.3 22.6 9.0 15.4 11.8 11.3 98.0 23.1 Bb 80.2 32.2 11.0 6.5 10.5 6.0 12.6 58.0 29.5 6.0 14.3 10.5 10.2 91.5 21.6 Bc 75.8 30.2 10.0 8.0 10.0 8.3 9.8 55.3 26.0 7.6 14.3 10.4 9.2 73.6 17.6 Bd 81.0 30.6 11.0 7.0 10.0 6.3 12.2 58.0 27.5 13.5 11.2 9.2 84.7 22.0 Bc 80.2 3C.2 10.0 6.6 9.5 6.3 14.0 56.5 26.5 13.5 12.6 9.6 9.0 85.0 23.1 Br 81.8 30.6 9.6 5.5 9.0 5.0 9.6 53.2 27.7 6.7 17.0 13.3 11.2 80.4 24.3 Bg 6006 30.4 3.6 3.0 9.0 2.6 9.4 50.0 23.0 6.0 17.0 12.3 11.3 75.0 19.0 Bh 73.4 27.0 8.6 3.0 8.0 2.6 9.2 53.6 24.3 6.3 16.2 12.0 11.2 74.5 16.9 Bi 82.6 29.2 8.6 5.0 8.0 5.3 6.2 53.0 25.7 7.2 14.9 10.9 9.5 766 20.9 Bj 80.6 2994 9.0 3.3 8.5 3.0 10.4 24.0 9.2 14.3 11.0 11.7 72.6 17.8 Bk 76.8 27.0 80 3.0 7.0 2.6 11.6 51.5 23.5 12.2 16.2 12.6 12.0 81.7 20.3 Bi 87.2 32.2 11.r 6.0 10.0 6.0 6.4 58.0 27.0 7.0 14.8 11.0 9.2 88.0 22.8 Bin 71.6 31.2 11.0 7.5 11.0 7.3 8.2 52.0 23.0 6.0 12.3 1C.6 9.8 79.4 19.4 Continuation Taxon

a k 1

Bn 75.2 30.6 10.5 7.0 i0.0 7.0 9.0 54.3 25.3 9.0 13.0 10.8 10.0 76.0 18.5 Bo 70.4 27.8 8.0 4.0 8.0 3.6 9.0 50.3 23.0 9.6 12.6 9.5 9.5 70.5 16.5 Bp 71.0 27.6 9.0 5.5 8.0 5.0 7.6 57.5 21.c 7.0 111.3 12.4 9.3 71.6 19.2 Bq 81 .2 31 .2 10.5 7.0 10.5 6.5 11.0 56.0 28.0 7.0 1 5.0 1 2.0 10.0 88.2 22.4 Er 78.6 30.0 904 505 9.0 4.0 9.6 57,4 27.6 8.0 16.c 12.3 10.2 78.0 19.0 Bs 79.2 29.6 00J 5.0 8.0 5.0 7.8 51.3 25.3 7.2 1 5.2 11 .6 9.6 77.2 20.2 Bt 80.0 29.8 8.5 5.3 8.0 5.0 9.0 L:6.0 23.5 9.0 15.c 11.5 10.4 71.3 18.0 Bu 75.2 28.4 8.4 4.0 8.0 3.5 10.0 P,0.5 24.2 11.5 16.3 13.2 11.2 83.0 21.2 By

By;

Bx Taxon C

a 1 ,,, „ci 65.6 29.0 8.6 6.0 8.0 5.6 6.0 39.0 26.0 7.0 14.4 10.4 9.0 82.2 20.7 Cb 68.0 27.2 10.6 5.5 9.0 5.3 8.0 44.5 21.5 7.5 13.7 11.0 9.3 76.5 19.0 Cc 60.4 26.8 9.0 ,.() 8.5 5.0 9.8 33.7 22.2 7.2 11.5 9.0 1000 55.2 15.6 Ca 67.4 27.2 8.3 3.0 8.0 3.0 9.8 41.0 26.0 9.0 15.6 12.1 9.2 75.9 23.7 Ce 81.4 39.6 14.0 8.5 14.0 7.6 13.0 46.0 23.0 12.0 14.2 11.6 11.4 68.2 19.2 -:f 65.0 26.4 7.6 3.3 7.5 3.c 5.0 L7.5 26.0 2.3 16.2 12.3 10.7 95.1 15.8 ,,,. ,, 54.0 25.0 6.0 2.0 6.0 2.0 8.c 36.6 22.3 7.0 14.4 11.2 9.2 69.5 25.8 Oh 81.2 35.4 10.3 6.0 9.0 6.3 10.6 L 8.7 28.7 9.2 15.9 11.5 9.9 78.6 19.6 ci 76.6 34.8 11.0 7.c 10.6 6.0 9.2 52.5 26.0 7.0 16.7 1200 9.2 81.2 20,1 - j 62.0 27.8 8.6 5.5 9.0 5.0 7.2 43.3 26.0 4.0 16.7 11.0 9.4 57.4 20.2 31k. 65.8 27.0 9.0 6.5 8.0 6.0 3.8 44.5 24.5 5.0 1L.5 11.2 8.5 56.4 20.2 Cl 66.2 30.4 9.0 6.0 9.3 6.0 6.7 40.2 26.2 7.0 15.0 11.2 9.2 81.0 20.1 Cm 64.2 28.6 8.5 5.3 8.3 5.3 6.5 42.3 25.6 7.5 14.0 10.5 9.2 79.2 19.5 Continuation Taxon C

a b c a e f g h i j k 1 m n o On 28.2 69.4 9.6 5.4 9.0 5.3 8.0 43.0 22.5 7.5 14.5 11.3 9.2 75.0 19.0 Co 62.0 27.4 9.0 5.0 9.0 5.0 8.5 36.2 24.0 706 14.2 11.1 9.6 61.0 16.3 Op 66.6 28.0 9.0 6.0 8.5 6.0 7.5 42.0 25.2 7.0 15.2 12.1 9.0 66.2 20.0 Cq 60.8 28.2 9.2 5.5 9.c0_,. 7,5 L,.0 26.2 6.3 15.0 11.3 9.3 68.0 20.2 Cr 68.0 29.4 8.8 4.0 8.0 4.0 9.0 42,5 27.0 7.8 16.2 13.0 9.2 80.8 23.6 I-co, cs 78.6 37.2 1c.3 5.6 1c.0 5.5 10.3 4:;02 25.0 1(.3 14.5 11.2 9.3 71.0 18.6 Ct 70,2 32.8 9.6 5.5 9.3 5.5 9.0 50.0 27.3 7.3 15.2 11.8 9.2 77.0 19.4 9°0? 0°09 9°9 O'El- 9°9 1- oeol, 0024 00 05 00 6 c'E 0°9 0°9 '17°CE TG

0°64 o°9L 0'01, 0"04 "6 0°61, 0*-,;G_- 006 GoE G-9 Go o°L oocz o'oi_ 31G o°aE t7°6 9°t19 0°94 9°6 O'F, 0'09 E"6 0°9 g°E o°9 9°t1 47°GL ca g°14 trOE 0°01. 5'6 0°04 0°EE z°95 9°6 o°c G°9 o°C o*L i7°5E o°LL Ta E°17 9'0L c"6 00z1, g*cl, 1.*E L't-g t7°04 0°C 0°9 o'c 9°9 oo9E oe2 uc 9°9 0•22. 2-6 2°14 z L4 7z 4 0 e'L G -- 0°L9 00 6 a°L o°6 o°L 00 6 9°Lz 47°99 2a S'OE 9°L9 go6 o°54 o°o4 cyoz c°G,Ti 9°W4 o°C- o°L C°c 7.,°L? if\ z° 46 HH o°64 o°22. c°6 9°9 1, C'6 G°64 5°o9 z"6 Goz 5o9 aoL °GE 9°9L as 5°64 z0 o9 0*O4 z° 9l, E"ol, E°1_4 E"og 0°6 C° o°9 G°E °9 17°Cz E'EL PG 0°94 L°oL 2°6 G'1 ,1- o°L4 G'6 ooLl, ooLty ooz Gog o°?, 9°S aozz o'69 ou 9°24 °92_ 9°6 0°14 4 0°94 G°o4 0°64 la°G C° o°9 G° o°9 9*t7 *17°Z. as o'oz 9'92. 9°6 0°14 2°54 crzz 5o9G 47°6 g'E 0°9 g" o°9 t7°9? *02

0 u T P z u 2 0 p 0 Continuation Taxon D

a k 1

Din 80.6 26.0 8.0 5.3 8.3 5.0 9.5 55.L 19.3 9.3 111.2 10.0 10.5 72.6 19.0 Dn 85.5 26.6 7.0 3.0 7.0 3.0 9.5 60.0 20.0 9.0 16.2 11.6 9.2 80.0 20.0 Do 75.6 24.0 6. 2.7 6.0 2.5 9.5 )43.0 17.0 9.2 17.0 1205 9.5 70.4 17.2 Dp 81.2 27./1 6.0 2.5 6.0 2.3 10.0 21 .3 9.8 16.6 12.0 9.7 77.0 19.2 Taxon E

a 1

Ea 93.2 34.4 12.0 7.0 12.0 6.7 11.2 60.5 26.7 10.5 16.5 11.9 9.7 85.2 25.5 2b 90.8 33.2 10.3 3.0 10.0 2.3 10.6 53.7 28.5 14.5 13.8 10.9 11.9 117.7 33.6 ,,c 76.0 27.8 8.0 4.0 8.0 3.3 10 0 L6.0 10.0 8.3 15.2 12.0 10.8 100.6 29.2 Ed 93.6 33.0 9.0 2.0 9.0 1.0 12.0 65.0 28.5 12.0 16.2 12.3 12.5 103.6 28.2 Ee 8.2 30.4 9.6 2.0 °_.. 0 2.6 16.2 17.2 19.0 13.2 15.1 11.6 11.6 105.0 23.5 Ef 74.0 27.8 7.6 3.6 7.0 3.0 10.0 1;.6.2 22.7 10.5 16.0 12.0 11.8 102.7 27.8 (_,cr 87.0 31.6 ,0.1.5 3 2.0 9.0 11.0 56.5 25.2 11.0 16.4 11.5 12.2 119.0 30.7 _11 79.0 30.2 7.6 4.3 8.0 4.0 10„2 50,0 22.0 10.0 15.6 11.6 11.3 85.2 24.7 --,-; _ 78.2 28.4 9.0 2.0 9.0 2.0 11. 5, 62.2 27.5 12.0 15.2 11.8 11.3 90.6 25.2 -]j 78.0 28.0 10.5 3.0 9.5 3.0 10.7 57.0 26.0 11.5 15.5 12.0 11.4 100.0 26.2 2k 84.0 30.4 9.0 2.0 9.0 2.0 11.3 .0 20.3 12.0 16.2 12.4 11.0 110.6 28.4 11 75.8 26.5 9.0 3.0 9.3 3.0 10.3 48.6 21.3 9.0 14.8 11.2 11.2 112.7 29.2 _.,,-1 74.0 26.0 8.4 3.8 8.0 4.0 11.0 4.2 21.0 10.0 15.5 11.8 11.5 92.0 24.2 Continuation Taxon E

a k 1 En 90.2 31.0 10.2 3.6 10.2 3.0 12.8 60.0 27.2 12.0 16.2 12.0 12.0 108.2 28.0 94.0 Jo 33.2 10.2 3.0 10.o 3.0 11.5 55.6 26.0 11.5 15.8 12.2 11.6 115.6 30.2 ]13, 82.0 30.0 10.0 3.0 10.0 3.0 12.0 F0.7 24.0 10.3 14.5 11.2 11.4 90.3 24.0 94.2 34.E 11.4 4.6 10.5 4.5 11.2 55.3 31.5 11.3 16.2 11.8 10.5 97.7 26.0 .±]/, 77.0 28.6 9.3 4.6 9.0 4.5 10.0 42.0 19.3 9.7 14.2 11.0 10.4 107.0 28.2 jls 85.4 30.0 9.3 2.3 9.0 2.0 12.0 7-0.0 22.3 13.5 16.o 12.2 11.4 105.6 25.8 :t 85.2 29.6 9.0 3.0 9.o 2.3 11.5 52.7 30.0 11.5 15.4 12.0 9.8 110.0 30.2 Ou 80./1 28.6 8.6 4.0 8.0 4.0 10.5 56.0 30.2 10.5 16.h 13.2 10.4 106.0 28.3 Taxon F

a b c d e f g h i j k 1 Fa 98.2 35.8 11 .3 6.5 11 .0 6.0 11 .8 53.0 30.0 1 2.0 1 5.4 11 .2 9.8 113.0 35.7 Fb 87.0 31.8 E.6 2.0 9.0 2.0 11.2 61.0 24.5 12.2 1L.3 11.0 12.2 92.7 30.9 Pc 82.0 31.8 10.0 3.3 10.0 3.0 12.6 55.0 25.0 9.5 12.0 9.0 11 .4 80.3 27.8 Fd 85.8 31 .6 9.3 6.0 9.5 5.3 14.6 52.0 27.0 13.0 11 .9 9.0 1 2.1 111 .3 43.1 Fe 92.0 35.2 12.6 3.5 13.0 2.6 11 .2 50.4 1996 12.2 17.1 13.2 12.2 94.7 35.6 Ff 87.9 35.2 10.35.0 10.0 4.3 14.6 54.3 29.5 11.2 16.2 12.1 12.8 75.4 35.8 Fg 87.0 33.1.9.0 6.3 9.0 5.6 1 ?.6 51 .5 21 .0 1 5.5 1 = .0 13.4 10.0 83.5 36.3 Ph 86.8 37 .2 11 .0 6.0 10.5 6.3 10.2 8.0 26.0 9.5 16.3 11 .5 9.0 99.1 29.2 Fi 96.2 29.8 9.6 5.0 9.5 4o6 10.4 6L1.0 21.0 7.6 15.9 12.511.3 98.9 31.2

87.0 30.E 8.0 LL. 0 8.0 2.0c, 9.0 51 .5 23.0 9.5 1 7.8 1 2.0 1 0.9 1 03.0 36.0 Fk 96.6 31 .8 8.6 4.5 9.0 4.3 1; .0 56.0 25.5 10.0 16.3 11 .3 11 .8 96.0 33.2 Fl 87.2 36.0 10.5 5.3 10.0 5.0 10.2 50.2 20.6 11.2 16.2 12.1 11.7 95.6 32.0 Pim 86.2 3)4.2 11.3 6.0 11.0 5.6 11.2 55.3 2)4.0 11.5 15.6 12.0 12.0 96.0 34.2 Continuation Taxon F

a b c ri e C ah i j k 1

Fn 95.2 31.0 9.0 5.0 9.0 4.5 10.2 58.2 26.0 10.3 17.0 12.4 12.0 100.0 35.2 Fo 90.0 31.6 9.0 4.5 8.5 4.3 9.5 50.2 23.0 10.0 16.8 12.1 11.3 1 01 .0 36.2 Fp 94.2 30.0 10.0 5.0 9.5 4.6 11.0 60.3 23.3 9.2 16.0 12.1 11.5 1 01 .2 33.0 Fq 85.0 32.2 10.5 5.5 10.5 6.0 9.8 56.0 25.0 9.5 15.8 11.3 10.2 96.1 28.1 Fr 86.2 34.4 11.0 5.0 10.5 5.0 12.0 50.0 25.2 11.0 16.5 12.2 12.3 80.2 31.7 Fs 87.2 34.2 10.3 5.0 10.0 4.6 11.3 56.0 24.2 10.2 15.2 11.3 11.7 85.0 30.6 H Ft 85.8 31 .0 1C.0 5.6 10.3 5.3 1 2.0 58.0 26.5 11.5 1 5.6 11 .8 .4 90.0 32.2 kc8 Fu 92.2 33.2 11 .0 6.0 11 .0 6.0 11 .3 56.2 28.3 1200 16.0 11 .3 10.2 108.6 34.6 Fv 86.0 30.2 9.0 3 , 5 9.3 3.3 11.5 57.2 23.3 11.7 15.5 12.0 12.2 95.6 31.2 Fw 80.2 30.6 9.0 4.0 9.0 3.5 11.0 60.0 24.0 13.0 17.2 13.0 9.7 103.0 35.2 Taxon G

a j k 1 Ga 91.6 36.2 11.3 8.0 11.3 7.0 12.8 41.5 21.7 9.0 16.o 12.3 11.6 113.4 34.8 Gb 79.4 33.8 11.3 6.0 11.0 4.3 11.0 50.0 27.0 9.0 16.5 12.0 10.1 100.9 24.8 Gc 74.0 29.6 7.0 3.0 7.0 2.6 8.4 149.0 25.0 11.0 14.6 10.2 11.7 87.5 21.1 3-(1 30.0 71.6 7.3 3.0 7.0 2.6 9.0 35.0 18.6 9.6 15.4 11.7 11.8 78.0 24.4 0To 72.0 29.2 7.6 3.0 7,0 2.6 8.2 43.3 22.6 8.0 16.2 11.3 12.1 75.8 20.5 3f 84.2 34.6 10.6 2.3 10.3 2.5 10.4 55.5 25.5 10.0 15.3 11.0 12.0 94.9 25.5 Gg 83.2 35.2 10.0 4.0 10.0 4.0 11.0 61.0 25.5 10.0 13.6 9.5 12.3 88.4 29.1 3h 80.4 35.0 9.0 4.5 9.0 3.6 10.4 45.0 17.5 9.5 14.2 9.5 11.8 78.3 25.1 Gi 86.8 35.4 10.0 2.0 10.5 2.0 11.E 6 .5 26.5 13.0 11.0 7.7 13.0 100.1 35.0 Gj 73.4 30.2 9.5 3.0 9.0 2.5 9.8 48.2 24.0 9.8 14.8 10.2 11.8 100,0 27.6 Gk 75.6 33.0 10.0 3.5 10.0 2.3 8.8 bi.0 21.3 10.0 15.4 12.4 11.0 88.2 25.9 84.0 34.0 12.0 5.5 12.0 5.6 12.0 57.5 23.0 10.0 16.9 11.6 10.8 83.5 28.5 rm 83.0 34.0 10.3 3.0 10.0 2.5 11.6 55.0 26.0 10.2 15.0 10.5 12.5 90.8 22.0

Continuation Taxon G

a b c d f g h i j k 1 m n o

Gn 90.0 36.0 11.3 6.5 11.3 6.5 12.0 43.2 22.0 9.0 16.0 12.0 11.5 108.3 33.0 Co 8206 33.0 12.0 6.0 11.5 5.6 12.0 55.0 23.0 10.0 16.5 12.0 9.6 90.2 25.6 Gp 75.2 30.0 8.5 3.0 8.0 3.0 9.5 50.2 25.0 11.0 15.2 11.2 12.0 85.0 22.3 3q 74.4 30.0 10.0 3.0 10.0 3.0 9.7 50.0 25.0 10.2 14.6 10.3 11.2 92.2 25.0 `ter 70.4 29.2 8.0 2.5 8.0 2.5 9.0 -0.0 19.2 9.6 15.7 11.3 12.0 86.6 25.2 ,-, ..,, 84.8 1--, 33.4 9.0 2.5 9.5 2.0 10.7 59.3 25.0 11.2 14.7 10.0 12.2 105.2 27.4 iv 84.0 34.0 10.0 2.5 10.0 2.5 10.0 5-02 25.2 10.5 15.0 11.0 12.0 98.2 26.0 72.0 29.0 8.0 2.5 8.0 2.5 9.7 46.2 22.89.5 15.3 11.0 11.7 80.2 21.6 Tv 76.8 32.0 7.5 2.5 7.5 2.0 8.7 52.0 25.3 9.0 14.3 10.4 11.4 85.0 21.2 "w 75.0 30.0 10.0 3.0 10.0 3.0 10.0 5_.6 24.0 10.0 16.0 11.8 10.8 96.2 25.0 Taxon H

a 1

Ha 90.2 39.2 12.0 5.6 11.0 5.3 9.2 58.8 32,0 11.6 15.7 11.0 9.0 9.6. 34.3 Hip 74.0 32.0 10.0 4.5 9.0 3.5 13.6 43.P 23.4 12.6 14.6 11.5 11.4 97.5 33.0 He 80.0 32.6 9.0 2.0 9.0 1.5 9.6 47.6 20.6 9.6 11.8 8.4 10.3 94.0 34.5 Hd 76.6 31.4 9.3 4.0 9.0 4.0 9.6 L'0.0 18.3 4.3 16.6 12.0 10.6 104.6 37.8 He 80.6 31.6 10.0 6.0 10.0 5.0 9.6 51.3 28.6 8.6 15.0 12.6 1 1.1 73.8 28.6 Hf 76.2 32.6 10.3 6.0 10.0 5.6 10.6 511..0 23.0 11.0 17.4 12.6 10.0 13',.1 48.8 Hg 87.0 36.4 11.3 6.0 11.0 5.6 10.6 50.E 25,0 11.6 16.2 11.5 10.9 125.9 35.2 Hh 76.2 32.0 10.3 6.5 10.0 6.0 1 1 .0 50.0 23.0 10.0 16.0 11.6 9.5 102.0 29.9 Hi 100.8 38.8 10.3 3.0 10.5 3.0 12.2 49.5 26.5 10.5 15.5 11.7 11.5 92.1 32.4 Hj 87.6 35.2 10.3 5.0 10.0 4.3 14.6 51-L..7. 29.5 11.2 15.6 11.8 12.8 75.4 35.8 78.0 32.4 9.3 4.0 9.0 4.0 10.6 52.0 27.3 10.2 15.8 12.0 11.2 99.2 33.0 H1 77.8 32.0 10.0 6.0 10.0 5.5 12.6 50.0 25.7 11.0 16.5 12.6 i0.8 105.0 30.8 im 76.4 32,0 10.9 5.5 10.0 5.0 10.4 52.0 24.8 11.5 16.6 12.3 10.6 95.2 33.0 Continuation Taxon H

rrz

Hn 85.4 36.0 9.5 4.5 9.3 4.5 12.3 4/1.5 25.0 8.6 14.5 11.0 12.2 80.4 36.0 Ho 96.6 37.6 10.3 3.0 10.0 3.0 11.7 50.0 24.8 10.2 15.8 11.8 11.2 90.1 31.3 Hp 76.0 31.8 10._'J 5.5 10.0 5.3 10. 55.3 24.2 11.0 16.5 12.2 10.3 110.2 35.3 Hq 88.b 56.4 11.3 5.3 11.0 5.3 9.0 57.2 31.3 11.0 16.2 11.2 9.6 95.6 33.4 Hr 85.6 35.2 11.3 5.0 11.0 5.3 1C.5 13.2 25.6 11.3 1 6.0 11.5 10.6 110.0 34.5 H Hs 79.2 31.6 9.0 2.5 9.0 2.5 9.3 a7.0 23.0 12.0 1.2 11.7 10.8 96.5 32.2 hb 75.0 30.2 10.5 5.0 10.3 t4.. 5 9.0 50.2 27.6 9.0 16.2 13.0 11.2 80.2 27.3 flu 80.0 31.0 10.0 5.5 10.0 5.0 10.0 50.0 27.2 9.3 15.5 12.3 11.4 78.0 27.0 Hy 76.0 31.2 9.0 4.5 10.0 6.5 9.3 4--61 20.0 7.5 15.7 12.0 10.8 100.2 34.3 Hw

Ftx

Hy Taxon I

a b c d a f g h i j k 1 In n o

I[l 69.8 28.4 9.0 5.0 9.0 4.3 10.8 49.5 26.0 10.0 16.9 12.9 9.1 86.9 38.8 Ib 89.2 35.6 11.6 7.0 11.5 7.0 8.8 52.3 27.0 10.0 16.6 13.2 12.0 93.5 37.9 Ic 7900 33.6 13.6 8.0 13.0 8.3 5.6 5303 26.6 7.3 11,05 12.2 10.4 103.9 20.2 Id 93.2 L;2.6 12.3 7.0 12.0 7.0 13.14 14..0 29,0 9.5 15.1 108 11.3 143.5 33.5 lc 75.4 32.0 10.0 5.5 10.0 5.3 7.4 'h.0r 23.3 9.0 16.2 12.7 9.6 79.9 23.2 If 76.2 34.6 11.3 8.5 11.5 8.0 8.0 5-.0 27.6 13.3 15.0 11.5 9.9 90.9 26.6 Ig 81.2 36.4 11.3 7.0 11.0 7.0 11.6 57.0 31.3 11.0 13.5 10.9 9.5 111.4 23.6 H t,.31 III 80.8 36.4 11.0 6.3 10.5 6.0 8.7 r_.5 .6 24.5 10.3 14.6 11.6 9.7 99.2 24.0 Ii 76.0 32.(--- 10.6 5.5 10.0 5.0 8.0 L7.0 21.2 8.5 15.7 11.0 10.2 92.5 32.6 II 90.0 53.2 11.6 5.6 11.3 6.3 12,0 50.4 29.2 9.3 16.6 11.3 11.0 125.2 30.6 11 47.4 34.2 11.0 6.6 11.0 6.3 10.0 55.4 2 ).0 10.0 1 6.0 13 .0 1 2.1 100.5 37.0

17 Ti 70.8 28.0 9.0 _).0 9.3 4.5 103 40.0 25.2 10.3 16.5 13.0 10,2 96.8 36.2 ID 80.4 33.6 11.6 6.0 12.0 7.0 9.6 517).0 28.4 8.7 15.2 11.6 10.3 100.2 22.3 Continuation Taxon I

a b c d 0 f g h i j k 1 m n o

In 81.2 36.0 10.5 6.5 11.0 5.0 11.0 5-=',.0 30.0 10.0 15.2 11.8 9.7 105.2 26.4 To 70.8 28.0 10.0 5.0 9.5 5.0 10.0 50.2 26.0 10.3 17.2 13.0 9.3 90.7 32.6 Ip 90.3 37.5 12.0 7.0 11.3 7.0 9.8 24J.0 27.3 9.0 15.8 12.3 11.0 95.3 23.2

Ta 81..3 514.0 11.0 705 11.0 8.C° 10.3 52.0 23.7 12.0 14.3 11.0 10.3 101.3 25.3 Ir 77.4 33.6 ic.c 5.3 10.3 5.0 8.0 ;c.c 23.0 10.c 16.0 12.3 1(.2 97.3 24.3 1--- ko, Is 80.3 34.0 12.6 7.5 13.0 7.3 7.0 54.2 26.0 8.0 15.0 12.o 10.8 92.3 20.3 0, It 85.2 36.2 11.5 5.5 10.3 4.5 11.0 F-2).0 25.3 9.5 14.6 11.C' 9.7 112.4 27.3 ill

Tv Tw Taxon

a

Ja 76./4 37.2 1 2.3 9.0 11.5 8.6 12.3 46.5 23.5 10.5 13.3 11.3 11.5 93.5 31.5 Jb 83.4 /43.6 14.3 9.0 13.5 9.0 13.4 52.0 25.0 11.0 16.4 13.6 11.1 93.6 30.3 Jc 85.4 40.0 13.3 6.0 13.0 6.0 15.0 64.6 35.3 13.3 15.8 13.0 11.5 115.4 39.2 Jd 76.8 38.0 1 2.0 6.5 11.5 6.3 8.0 45.0 28.3 8.6 15.6 11.6 11.4 117.6 1+7.3 J6 79.8 37.0 12.0 6.3 11.0 6.0 8.0 43.0 24.0 10.0 15.3 10.3 12.4 108.8 26.0 Jf 64.4 324.8 13.0 10.3 12.8 8.0 11.6 68.0 35.0 12.0 14.5 10.5 10.1 99.7 35.1 Jg 79.4 37.2 13.0 7.0 13.0 6.0 1 2.8 47.0 23.0 10.0 16.4 13.0 9.6 117.7 35.0 Jh 74.0 36.6 11.3 3.0 11.5 2.0 1 0.2 Lir .0 27.0 11 .3 13.1 10.1 1 2.1 91.1 28.5 Ji 74.6 36.0 10.3 /4.3 11.0 4.0 11.6 55.0 3000 13.0 114.1 10.3 11.5 103.7 40.6 Jj 72.4 36. 10.6 3.0 10.3 3.0 9.0 41.5 23.5 10.5 15.3 12.1 10.1 77.0 30.2 Jk 76.2 38.6 12.3 8.0 11.5 7.3 10.4 58.5 29.0 9.5 17.6 13.8 10 0 69.9 28.0 79.4 40.0 11.3 7.0 11.0 6.3 11.4 .0 28.0 9.0 15.6 13.1 9.9 67.9 25.1 Jm 76.2 36.6 11.6 6.0 12.0 6.3 10.6 43.0 27.5 9.0 15.1 12.1 10.2 80.0 27.8 Continuation Taxon J

a b c d e f g h i j k 1

Jn 87.0 38.4 12.0 8.0 11.0 7.0 11.2 63.6 33.0 13.6 16.9 12.0 10J) 100.1 34.9 To 82.6 38.2 10.0 7.5 10.0 7.0 10.6 63.6 33.3 10.0 17.5 13.4 9.2 96.7 29.8 Jp 76.0 37.4 1 2.0 8.0 11.5 8.0 12.0 48.0 23.5 10.5 14-0 11 .5 11 .2 95.5 32.0 70 75.2 37.0 11.0 4.0 11.0 4.0 1 0.5 47.2 27.0 11.0 12,01 11.2 11.5 *5.0 35.5 Jr 74.0 37.0 11 .0 4.5 11 .0 4.5 1 0.2 60.0 31 .2 1 2.0 1 5.2 11 .3 11 .5 1 00.2 37.5 Js 81.0 40.0 12.0 6.6 13.0 7.0 12.0 140 . 2 214.0 10.0 16.0 13.0 10.6 110.2 36.2 st 66.2 34.0 1 2.0 8.0 11.3 7.6 11 .6 60.0 29.5 1 2.0 1 5.0 1 2.0 1 0.8 90.8 30.2 Ju 75.0 38.0 12.0 8.0 12.o 8.0 12.0 48.2 214.5 11.0 14.2 11.5 11.3 95.2 32.0 Jv 81.2 39.3 13.0 6.0 -12.0 5.5 12.0 (5.0 35.6 13.0 15.3 12.8 11.3 105.0 36.2 Jw 82.0 40.2 11 .9 6.0 11 .0 6.0 9.0 50.0 25.0 11 .0 16.3 1 2.3 11 .7 100.3 28.3 Jx 75.3 3= 03 11 .5 7.3 11 .5 7.0 ,11..0 56.3 30.0 10.5 16.5 13.5 10.2 75.2 27.3 Taxon K

a i j k 1 Ka 88.0 43.5 14.0 8.5 13.5 8.0 13.3 53.3 32.3 12.5 16.5 12.3 11.2 125.0 42.2 Kb 85.2 43.2 15.0 900 14.0 9,3 -14.2 52.5 29.5 13.5 16.2 11 .1 9.5 158.2 54.2 Kc 73.2 35.2 13.6 8.0 13.5 8.0 14.), 61 .3 34.0 12.6 15.1 11.6 8.2 109.0 42.6 c, Kc? 81.0 38.3 13.0 7.0 13.0 7.0 12.3 v., .,, 34.3 12.3 16.4 12.6 11.0 112.0 40.2 Kc 85.4 43.6 14.6 9.5 15.0 9.0 10.6 62.0 40.3 13.0 17.3 13.2 10.0 105.1 43.1 Icy' 74.2 34.8 11.6 6.o 11.0 6.0 9.4 51 .6 32.3 11.3 16.8 12.3 9.6 108.7 35.2 H Kg 92.6 43.2 13.6 7.0 1 4.0 6.6 13.4 6!--.3 37.6 13.0 15.0 -12.2 11.3 117.3 42.4 Kh 75.L 39.2 15.0 7.0 13.0 6.6 12.2 51.0 33.5 13.5 15.3 12.3 11.5 141.6 41.0 Ki 87.8 41 .6 13.3 6.0 13.0 6.0 11.2 46.0 27.0 10.0 16.7 13.0 10.2 103.0 37.7 KJ 86.2 39.2 12.0 6.0 11.6 6.0 6.4 43.5 25.6 6.0 14.8 ii .4 11.1 90.2 30.7 Kk 85.6 39.4 12.0 7.5 12.0 7.3 12.0 245.5 22.5 8.5 16.3 12.5 11.3 103.2 35.5 1 86.4 42.6 12.3 6.0 11.5 6.0 11.6 50.6 30.3 12.0 15.6 12.3 11.2 108.6 37.0 Kin 80.0 40.2 12.3 6.0 12.0 6.0 1203 60.2 35.2 12.3 16.5 13.2 11.3 112.0 40.2 Continuation Taxon K

a b c d e f g h i j k 1

Kn 86.2 38.4 12.6 8.0 12.5 7.3 13.4 55.5 31.0 13.0 17.6 12.1 11.5 131.8 39.9 Ko 94.0 33.4 10.6 4.0 10.0 4.0 12.4 65.0 38.0 12.0 15.5 12.2 10.7 100.7 29.5 Ki.) 84.6 41.4 13.0 7.0 12.5 6.5 13.2 57.3 32.3 12.0 16.0 12.8 11.2 108.2 37.2 Kq 85.2 40.0 13.3 7.0 13.0 6.3 12.6 52.0 34.6 12.3 13.6 10.6 11.0 120.6 45.1 Kr 86.6 39.8 13.3 7.0 13.5 6.6 12.0 53.3 34.3 11.6 10.2 8.6 11.8 128.5 46.0 Ks 86.0 L3.0 13.0 8.0 13.3 8.0 11.5 60.0 35.2 12.0 16.3 13.0 10.3 110.2 41.3 iv 0 lYt 75.0 37.0 13.0 7.0 13.0 7.6 13.3 64.3 35.2 12.5 15.5 12.2 9.3 115.8 40.3 Ku 87.2 44.0 1/1.0 8.5 13.5 8.0 13.5 50.0 28.5 13.0 16.0 12.0 9.8 120,0 43.0 Kv 88.4 45.0 13.0 6.7 13.5 7.0 13.0 63.0 38.0 13.0 15.0 12.0 11.4 120.2 42.0 Kw 80.0 38.2 13.0 7.0 12.5 7.0 13.2 50.2 25.0 10.3 16.0 12.4 11.5 100.6 35.0 x 81.0 40.0 12.0 6.3 12.0 6.3 12.6 57.6 31.0 11.5 15.5 12.3 11.3 110.2 40.3 Taxon L

a 1

La 68.6 34.0 1 0.0 7.0 1 0.0 6.3 1 2.0 50.0 28.0 9.0 15.5 12.0 11.4 146.1 22. Lb 714.2 36.6 1 2.0 5.5 11 .5 5.3 10.6 40.0 24.9 9.0 17.0 1 2.0 1 0.8 114.3 31 .3 Lc 75.2 33.2 11.0 5.5 11.0 5.0 10.6 50.0 31.0 10.0 14.0 12.0 11.0 11 )4.0 26.9 Ld 80.8 4.0.2 12.6 6.0 12.0 5.6 8.4 43.0 29.0 11.0 17.7 12.0 11.2 15.5 39.6 Le 82.2 40.8 12.0 6.0 11.5 5.3 10.6 Y. L1 32.4 13.0 17.0 12.3 10.8 120.2 27.3 Lf 63.6 314.0 13.0 7.0 13.0 7.8 12.2 51.0 37.0 11.0 18.5 13.5 10.6 1)1),.1 43.5 LET, 85.2 43.0 1/4.0 8.0 13.5 8.0 8.8 55.0 32.0 13.0 18.5 16.2 12,5 160.0 36.0 L1-1 83.2 41 .4 12.0 6.5 11.5 6.0 10.8 56.0 34.0 1 2.6 1 5.4 11 .3 10.2 173.2 38.8 Li 77.4 38.0 11.5 6.0 11.5 6.0 10.6 52.6 32.2 11.0 16.0 12.0 11.0 114.0.2 35.8 Ll 82.8 /40.2 11 .6 5.0 11 .0 4.3 1 2.4 44.2 27.2 1 0.2 1 6.7 11 .9 11.8 11 9.2 50.0 Lk 70.8 35.0 11.3 6.0 11.0 5.5 10.4 50.0 30.2 11.0 16.2 12.0 11.1 130.2 30.0 la 74.0 36.0 12.0 6.0 12.3 6.3 10.4 55.7 31.8 10.5 15.7 11.5 11.41557 32.6 Lrn 75.0 37.8 12.0 6.5 11.5 6.0 10.7 56.0 32.0 10.4 16.0 12.3 11.3 160.2 38.7 Continuation

Taxon L

a b c g d a f . h i j k 1 in n 0

Ln 70.2 34.6 10.5 6.3 10.0 6.0 11.2 48.6 28.3 10.3 16.2 12.3 11.2 130.6 23.3 Lo 83.6 41 .0 12.0 6.3 12.0 6.0 12.2 46.2 27.0 11.3 15.3 11.7 11.0 120.0 27.8 Lp 73.6 36.0 11.0 I , r 5.5 11.0 5.3 11.0 4- .0 25.0 10.0 16.7 12.4 11.1 116.0 2-.2 Lq 80.0 39.6 12.3 6.0 11.5 6.0 11.2 5;.0 34.0 10.8 16.2 12.0 11.1 135.6 33.3 Lr 75.4 35.4 3 . ry 11.0 5.5 11.3 5.0 10.7 f2-.6 0 2 10.3 15.2 12.7 11.0 120.2 25.7 0 NI Ls 84.0 43.6 1 2.5 7.0 1 3.0 7.0 1 0.0 5-.0 31.6 1 2.0 17.2 14-0 11 .8 140.6 30.3 Lt 81 .6 40 2 11 .5 6.0 11 .5 6.0 11 .0 50.0 30.3 12.2 17.5 13.6 10.5 120.2 27.3 Iu 66.4. 34..0 12.0 6.0 12.3 6.0 10.3 /4 5.0 25.3 11.0 18.0 13.8 1 2.0 1450 31.0 Lv 86.2 44.6 13.0 7.0 13.0 7.0 11.6 13.7 26.0 10.4 16.1 13.0 11 .4 152.0 35.6 Lw Lx

Ly Lz

Taxon M

a b c d e f o• 1

Ma 101.2 40.8 13.3 8.0 13.0 7.3 16.5 61.5 30.5 13.5 15.0 1e.b 11.1 179.2 35.5 Mb 89.2 34.0 10.3 6.0 10.0 5.3 17.6 L, 5. 5 27.5 16.0 13.5 11 .5 9.5 116.5 25.2 Lc 99.6 39.8 14.0 3.0 13.c' 705 11.0 5C.0 140,0 15.0 16.7 12.8 11 .4 147.9 3c.7 :Id 37.6 34.0 10.6 6.0 10.5 5.6 11.6 6.0 30.0 14.0 17.2 13.`-' 14.6 138.1 33.0 iv Me 100.4 36.0 -1-1 . 7.c 11 .5 7.0 11.8 75.c 35.0 15.0 19.0 15.0 11 .4 204.9 27.9 W0 Mf 82.0 37.6 11.3 8.0 11.0 7.6 52.5 38.7 8.7 1 7.0 1 2.3 11.5 217.8 30.2 r Mg 90.0 34.0 1 0.0 4.0 10.0 4.0 13.49'4 .0 293 13.6 17.0 12.0 10.1-1 126.1 24.2 Mh 85.0 34.0 11 0c- 6.c 11 .0 6.3 12.0 5(.5 35.5 12.5 17.2 12.6 11.3 163.033.9 'Ti 95.8 38.0 11.6 7.0 11.3 6.5 11.7 57.c 37.0 13.0 16.8 12.8 11 .7 165.0 32.3 ':j 93.2 37.8 12.0 7.0 12.0 7.3 11.2 6O.8 33.7 12.4 17.0 13.o 11.3 150.2 31.0 Mk 87.8 34.4 10.0 5.5 10.5 5.5 13.7 50.2 29.0 114 14.2 11 .2 11 .2 120.8 224.8 Ml 98.7 38.2 13.0 7.0 13.0 7.3 12.0 57.7 34.7 13.8 16.2 1 2.3 11 .7 150 .3 35.8 111 86.4 33.6 12.5 7.0 12.5 7.0 12.2 65.3 33.7 13.0 16.8 13.0 11.1 142.7 32.3 Continuation Taxon M

a 1

Mn 96.2 3-.0 12.C;7.0 12.0 6.5 12.3 65.0 34.5 1205 17.0 13.2 11.2 1110.6 2803 Mo 97.6 40.0 12.5 7.3 12.0 7.0 13.6 57.0 30.0 13.0 1603 12.0 11 .3 155.0 32.5 Mp 84.6 34.2 11 .5 6.0 11 .0 600 1 2.0 62.0 34.3 1 2.0 1 7.3 1 3.0 11 .0 1 60.2 35.0 Mq 90.8 35.0 11.0 6.0 11 .3 600 1)4.2 1,8.( 2702 1)4.3 16.0 12.8 10.5 12000 25.6 Yr 90.2 311.6 11 .G 5.3 11 .0 5.0 13.0 12,1 .6 2803 1h .0 16.3 1 2.3 11 .2 1 25.8 23.7 99.0 40.0 13,0 7.0 13.0 7.0 11 .0 5u . 0 3703 14-.3 16.4 11 .8 li .3 i 5C .7 31 .2 Mt 81.8 3,4.0 11.3 6.5 11.0 6.0 10.5 37.3 11 .3 17.0 13.0 11.6 165.0 36.4 Mu 88.2 3 504 11 .0 6.0 11 .0 6.3 1 2.3 56.0 29.3 1 3.0 16.5 1 3.0 1 2.6 1110.6 32.6 'dv 92.8 37.2 1 2.0 7.0 1 2.0 6.5 II .5 514.0 30.6 1 2.6 15.3 11 .5 1 1 . 2 138.0 32.6 Mw 93.0 37.0 11 .6 7.0 11 .0 6.3 1 2.: 62.3 34.6 11 .d 1 5.7 11 .0 11 .3 1 53.3 31 .8 Saxon N

a 1

_a 101 34 42.0 13.0 7.0 12.5 7.0 12.6 51.0 31.n 13.0 15.3 11 .LL 11.5 124.1 39.0 no 82.4 32.0 12.3 7.0 12.r 7.0 12.0 55.0 27.2 11.3 16.3 12.2 11.7 116.2 38.0 c 70.2 30.8 906 4.0 9.5 3.6 9.0 30.2 20.2 7.7 16.2 12.5 12.3 113.8 35.1 -d 76.2 3c.5 11.0 C.c 10.o 6.0 1c.6 L7.3 21.0 1000 46.9 13.8 10.0 106.8 32.7 ry le 38.6 39.n 12.6 8.5 13.0 8.0 14.6 54.5 30.5 13.0 15.2 11.0 9.6 114.5 34.0 ui ,f 86.8 33.0 11.0 7.0 ic.0 6.0 14.8 52.5 23.5 12.0 14.7 11.4 11.6 113.5 41.0 „ID- ._, 91.2 35.2 I, .6 3.0 10.0 3.c 1..-5 53.3 29.0 12.6 15.2 11.7 11.1 112.4 3).2 h 90.0 35.8 12.3 6.8 12.0 6.8 12.2 59.0 26.6 10.3 15.4 11.6 10.1 141.6 52.4 -1 86.2 33.8 12.0 7.0 11.0 7.0 11.4 59.5 27.7 12.5 16.c 12.2 11.0 151.5 47.5 -J 79.6 34.0 13.0 7.0 13.0 6.0 13.0 5.0 25.0 11.o 16.7 12.6 9.4 144.3 48.1 --k 88.0 36.8 12.6 6.5 12.o 6.0 12.6 5L,.0 24.0 9.8 16.2 12.7 10.0 96.1 38.3 , 75.0 32.6 9.6 5.0 9.o 4.6 5.2 b,6.6 22.3 8.0 14.7 11.4 11.0 97.6 44.9 , 83.2 3)4.2 12.3 6.0 11.5 6.0 9.2 46.c 25.0 6.3 17.4 11.9 11.0 95.1 39.7 Tiontinuation 2axon N

k 1

Tn 100.2 40.6 12.5 7.0 12.5 7.0 13.0 55.2 28.0 12.5 16.3 12.1 11.9 130.2 44.0 -;(:) 95.0 31 .8 9.0 5.c 9.0 5.0 10.4 59.3 24.3 8.0 14.9 12.1 11.7 147.5 45.0 .dp 100.0 41.0 12.5 7.0 12.0 7.0 1 2,0 54.0 25.3 12.0 15.6 11.5 10.8 122.3 40.2 ,,c1 93.2 36.4 1 2.0 7.0 1 2.0 5.5 1 '2.3 58.3 27.6 11.3 15.0 11.2 10.3 146.0 51.3 34.2 32.8 11.3 6.0 11.0 5.5 11.5 56.3 27.0 11 .0 16.1 12.3 11 .5 1 20.3 39.0 :s 89.0 35.0 11.0 5-0 11.0 5.3 12.r 5-0 28.7 11.7 1 5.3 11.6 11.2 116.2 3(3.3 c),o -t 73.6 31.6 -icou 4.5 10.0 4.0 1C.0 /.2.0 21.0 9.6 16.7 12.3 12.0 1u8.3 35.2 "ru 77.2 31./1 10 05 5.5 1 0.0 5,8 1 0.7 h5.3 21.3 10.0 17.1 13.4 1C.4 102.0 33.9 4V 86.0 33.0 12.( 6.5 1 1 .5 6.0 11 0C 59.0 29.0 11 .4 1 4.1 11 .0 11.3 130.0 43.4 4w 83.0 34.4 12.0 6.0 11.0 5.5 10.( 48.6 22.8 11.2 17 3 12.4 11.5 100.8 34.2 -x 88.2 36.6 1 2.3 7.0 12.0 7.0 12.G 56.3 25.6 1 13.0 11.0 11 5.0 39.6 "axon 0

a 1 -Da .0 6.0 11.0 94.4 39.4 11 6.3 10.0 56.3 25.6 11.0 16.3 12.0 9.3 110.5 30.0 Dip 90.2 36.L1 12.3 7.0 12.0 6.5 11 .0 57.0 30.0 11 .0 1 6.1 1 2.0 1 2.1 1 22.6 31.1 c 85.6 35.4 , 10 0 3.0 10.0 3.0 10.6 53.0 24.5 14.0 1 5.0 11 .3 1 2.2 112.9 31 .9 :d 9)4.8 38 6 12.5 9 7.0 12.0 5.5 10.2 53.0 29.0 12.6 16.7 12.3 11 .7 86.6 23.7 e 50.0 35.2 11 .3 2.3 1100 2.3 11 .8 25.0 10,0 -16.0 12.0 12.6 100.8 26.8 :4' 94.8 39.8 12.3 8.0 12.0 7.0 12.2 5705 33.0 12.3 17.8 12.1 9.1 13.3.0 33.9 .;g 95.0 38.0 12.0 7.0 12.0 7.0 11.6 613_ .6 32.0 12.2 17.',1 12.2 12.3 125.0 32.6 ,38.2 35.0 11 (-;.0 11 .0 6.0 11.6 52.8 22.4 13.0 16.8 12.3 11.9 113.2 30.4 i 99.8 35.4 10.6 10.0 3.0 2.6 8.6 43.7 19.5 7.5 14.7 11.9 12.2 115.1 31.0 j 109.8 38.8 12.0 3.5 11 .0 2.6 14.c 6':.3 29.3 18.0 15.0 -11.E 12.6 97.5 26.2 Lk 98.7 38.7 6,0 11.0 11.0 6.0 11.5 60.5 27.3 13.3 16.6 13.0 11 .8 107.0 27.3 1 88.2 36.7 11.5 5.0 11.0 5.3 10.5 55,.2 31.7 11.5 17.1 13.2 11.4 122.0 30.7 93.6 37.0 1 0.0 )4.5 10.3 5.0 9.5 45.8 20.6 10.0 15.7 11.3 10.8 110.6 28.6 Continuation Taxon 0

a U c cz e f h i j k 1 On 93.0 37.0 10.5 5.c 10.0 5.5 10.3 4 .3 22.0 10.3 15.0 11.0 11.0 115.2 28.0 Oo 93.2 38.6 11.0 5.5 11.0 5.0 11.0 55.3 24.6 11.3 16.0 12.1 10.4 117.3 30.3 Op 100.0 4r.4 12.5 5.3 12.0 5.3 12.0 62.6 28.0 14.3 17.2 12.6 11.2 105.3 25.0 3q 91.6 37.0 12.0 6.5 12.3 6.0 11.6 56.0 26.3 10.8 15.6 1 1.8 11.3 108.2 27.6 Or ':)C.4 36.0 11.0 5.0 11.3 5.0 11.3 62.8 29.3 11.3 16.6 13,0 11.3 118.3 30.0 Os 86.2 35.0 11.E 5.0 11.5 5.c 11.0 - 0.6 23.3 12.0 16.5 12.7 12.0 116.6 32.3 Ot 94-0 38.0 12.0 6.5 11.5 6.0 10.3 6;:.3 28.7 12.6 16.0 12.0 10.8 102.3 27.0 0u 8 .6 36.4 1 0.5 5.5 10.0 5.5 10.6 502 25.0 11.7 14.6 11.2 11.0 96.8 25.3 Ov Ow Taxon P

a j k 1 Pa 81 .0 13.6 7.0 40 13.0 7.0 13.0 67.6 36.3 18.0 19.5 1 5.5 11 .4 1 57.0 56.6 Pb 80.4 42.2 11.0 7.0 11.0 6.0 10.4 66.3 36.3 11.0 17.7 13.3 11.3 188.4 57.1 Pe 99.4 4 8.8 17.0 8.0 17.0 8.0 13.0 56.0 31 .3 12.66 17.3 12,7 12.0 156.6 53.1 Pd 87.6 47.2 1 5.6 8.5 1 5.5 8.5 13.8 6C .3 33.3 16.3 1 6.6 1 2.5 11 .7 161 .6 50.0 Pc 81.6 42.8 14.0 1C.0 14.0 10.0 12.2 622,.3 33.6 15.0 16.7 12.5 11 .8 140.0 57.0 Pf 96.0 48.6 1n.0 1(.0 13.0 9.3 14.8 64.3 37.0 17.0 14.6 11.8 162.5 52.0 Pg 111.4 56.2 17.9 9.0 15.0 9.0 17.8 64.6 36.3 1 9.0 20.0 1 6.0 11 .8 164.0 56.0 Ph 97.6 118.0 1'.3 10.0 1 5.0 9.6 1 5.3 76.0 37.0 17.5 1 9.2 1 5.5 11 .4 1 78.5 54.8 2± 106.6 51.6 15.3 11 .0 15.3 10.3 14.8 75.0 40.2 25.0 17.0 12.6 10.9 164.9 55.4 2j 110.2 51 .8 16.3 10.5 16.0 10.6 18.0 76.3 41.6 22.3 19.7 14.2 11.5 1 40 .5 46.0 Pk 79.2 36.2 10.6 5.5 10.0 5.6 8.2 1,6.L)-- 28.4 11.4 15.0 14.8 .11.2 109.8 33.1 1108.4 53.4 15.3 9.5 15.5 9.3 18.6 75.0 39.0 15.5 17.0 12.0 10.8 149.6 48.1 Pm 109.2 5C .4 1 5.0 1 0.5 13.5 9.6 14.6 67.0 34.5 1 9.0 1 9.1 13.9 11 .4 1 87.5 57.0 Continuation -L axon P

a 1

±n 94.4 47.4 16.e 8.0 16.5 8.0 13.8 61.0 32.3 46.0 18.1 1 1, . 11.6 137.0 46.0 Po 89.6 44.c 15.5 9.0 15.0 9.0 1300 65.0 33.5 18.0 17.5 12.8 11.6 164.9 49.7 Pp 90.8 46.4 16.5 10.0 16.0 10.0 14.2 64.5 37.0 22.0 1 5.0 12.0 11.2 161.0 54.1 Pg 106.0 52.8 15.5 9.5 15.0, 9.0 17.3 77.3 h( 16.6 17.1 12.5 11.3 14t _.2 48.8 Jr 86.4 44.2 1 4.5 9.3 1 4.5 9.0 13.3 c 2.3 33.0 1 6.0 16.G 1 2.2 11.0 1 55.3 50.1 0 2s 1 C0.6 /0.2 16.5 9.5 1S.c 9.3 1 4.6 60.0 33.6 13.6 1 7.5 13.0 11 .8 1 60.0 52.3 ot 81 .4 39.c 1 2.6 7.3 1 2.3 7.0 1 .3 57.3 23.6 12.0 1 9.6 1 5.2 11 .3 1 25.3 39.7 ?u 10c,.2 52.4 16.0 1C.5 1 6.0 IC.3 17.0 75.3 40.r 20.0 19.0 I 4.0 11 .6 1 46.0 48.3 ?v 96.6 47,2 15.0 10 .0 15.3 10.3 15.0 72.6 38.3 16.3 1 3.6 15.0 i0.8 166.0 52.3 Pw 82.0 42.6 11.8 7.0 11.5 6.5 11.3 6-0( 36.6 12.3 17.0 13.0 11.1 170.3 56.6 :Dx 91 .5 46.0 1 4.3 9.0 1 4.o 9.0 1 2.0 6-.3 37.0 1 3.6 1 .0 1 5.2 1 2.1 1 50.0 55.2 Taxon Q

a b c d e f g h i j k 1

Qa 90.2 44.5 15.0 8.0 15.0 8.0 13.4 56.0 33.5 17.0 17.5 13.0 11.5 149.0 47.1 Qb 94.0 44.0 13.3 7.0 13.0 6.0 14.8 69.7 143.5 19.5 18.1 13.2 11.2 138.1 52.8 Qc 89.6 45.4 124.6 10.0 14.0 10.0 114.2 61.6 41 .3 20.0 17.3 11.4 9.5 15/4.9 52.2 Qd 89.0 14.6 15.0 8.3 15.0 8.0 14.7 39.6 26.3 12.6 18.0 12.5 10.1 136.2 40.4 Q.e 98.4 47.5 16.0 10.5 15.0 10.3 15.3 65.0 )44.0 18.0 16.5 11.6 10.7 107.6 40.3 Qf 104.8 52.2 17.0 11.0 17.0 10.6 16.2 71.0 47.3 19.6 17.3 13.0 11.o 179.3 68.6

Qg 98.0 46.4 16.3 11.0 16.0 1 0.5 1 6.0 60.6 42.0 18.0 17.2 11.5 10,6 15/4.7 56.8 Qh 101 .0 49.6 16.0 11 .0 16.0 10.3 17.0 77.0 50.0 24.0 18.2 13.3 10.8 140.0 46.3 Qi 100.2 49.0 1 5.0 11 •5 16.0 11 .3 13.2 60.0 44.0 19.0 19.3 13.6 11.0 133.7 43.7 Qi 94.3 45.8 14.0 8.o 14.0 8.3 15.3 52.0 33.0 15.0 17.6 12.3 10.6 165.8 50.3 Qk 100.6 49.2 1 6.0 13.5 15.0 12.6 13.8 88.0 56.5 18.0 j5.0 10.5 10.2 184.4 67.6 za. 86.2 42.8 15.3 8.5 .15.0 8.0 14.6 67.6 41.0 16.3 15.1 10.2 10.0 171.0 6c.o Qm 89.8 42.8 13.0 8.0 12.5 7.6 13.0 53.6 36.3 114.6 21.0 15.1 11.0 144.6 44.3

Continuation Taxon Q

a 1

ran 79.0 41 .0 13.3 7.0 13.0 7.0 13.6 51.0 34.0 13.0 6 11 .1 10.8 156.5 51 .0 C)o 78.8 39.6 1 4.6 7.0 14.0 5.6 12.8 52.5 33.0 16.5 ).0 14.4 10.7 133. ) 45.0 Qp 91.2 45.0 15.3 11 .0 1 5.0 10.3 13.2 68.0 44.0 15.5 - `3 .6 14.3 9.5 1 o9. ; 38.0 83.2 42.0 14.3 8.0 1 4.0 7.5 13.2 51 .0 33.5 16.0 18.0 13.1 10.6 122.3 36.4 ")r 100.2 53.0 17.6 13.0 17.5 12.0 15.4 62.0 38.0 15.0 17.0 127 10.0 173.1 52.4 Qs 83.0 43.2 13.4 10.0 13.0 9.0 13.4 55.5 39.5 13.5 20.7 15.8 10.3 160 0 61 .0 Qt 89.6 45.2 13.6 8.0 1 3. 8.0 13.4 39.' 2C .6 2C .7 14.5 11 .6 137.0 46.0 Qu 92.8 48.6 13.6 8.0 13.0 7.3 18.8 56.0 4)4.3 15.0 17.4 12.9 10.7 164.9 60.9 Qv 113.0 55.6 16.3 11 .0 16.5 11.6 16.6 86.3 53.6 18.6 14.7 10.0 11.7 192.9 62.1 90,2 45.6 13.3 7.0 13.0 6.5 14.3 53.6 40.0 16.0 16.1 11 .3 10.5 121.0 4). .0 8)4.4 40.8 13.0 6.0 13.0 6.0 13.3 a .0 35.0 14-.3 18.1 12.7 10.7 174.0 51 ,o .Y 96.6 49.0 16.0 10.0 16.0 10.0 15.2 55.0 34.0 13.0 163.5 13.1 11.0 176.0 6).3 7,z 90.2 /44.0 13.3 8.0 13.0 7.3 11.6 53.6 38.8 12.5 18.0 13.14 12.0 156.6 55.0 86.6 43.0 14.3 6.0 14.0 5.3 11.8 53.6 35.0 13.0 14.0 11.0 10.5 140.3 45.2 89.0 45.2 14.6 7.6 14.3 7.5 13.6 57.3 39.0 13.6 17.0 11.3 10,6 160.3 55.6 Taxon R

a b c d e f g h i .1 k 1 m n n Ra 104.4 47.6 15.0 12.0 16.0 12.6 13.8 79.0 51.0 18.0 18.2 14.1 10.8 152.4 58.4 Rb 84.2 40.2 13.0 7.0 12.0 7.0 13.8 72.5 40.0 17.0 16.5 14.5 10.5 156.3 51.4 Rc 95.6 424-.6 15.3 9.0 15.0 8.3 13.4 57.0 39.0 14.o 20.0 13.5 10.7 170.0 59.1 Rd 104.8 )1)1.6 16.0 8.0 16.5 7.3 16./4 77.0 41.0 180 8 20.3 14.1 12.9 158.2 52.9 N) Re 79.2 37.8 12.3 7.0 12.0 7.0 11.6 54.6 29.0 12.0 19.4 1 4.0 10.0 145.0 Rf 90.2 40.0 12.0 7.0 11.6 6.0 14.2 49.8 29.2 14.8 15.1 11.3 11.3 131.3 3451 :60 1--ju4 Rg 97.8 45.2 14.3 9.0 14.0 9.0 -1,40',) 68.7 35.0 20.0 17.5 12.9 10.5 113.0 43.5 Rh 94.2 43.8 13.0 6.3 12.5 6.0 13.0 58.5 31.5 19.0 16.2 13.2 11.0 191.2 59.2 Ri 97.8 41.0 12.3 7.5 12.5 7.0 15.6 52.0 34.0 13.3 16.5 12.5 11 .4 183.3 66.6 Rj 104.2 43.4 16.3 8.5 16.0 8.3 16.3 69.3 37.0 16.0 15.2 10.2 9.5 154.0 53.7 Tr,C. 104 2 43.4 14.0 8.0 14.0 7.6 14.6 65.0 39.0 14.5 20.5 14.2 9.5 228.5 84.8 Ri 112.6 50.6 16.0 0 10.0 16.3 J0o 15.0 75.3 38.3 16.3 19.0 12.2 10.5 142.6 48.2 Pm 79.2 35.4 12.6 6.5 12.0 6.5 13.2 58.8 28.6 15.4 18.3 13.2 10.2 149.8 48.2 Continuation Taxon R

a d e h 1 m n 0 Rn 90.4 40.6 13.5 8.c 13.0 8.0 14.0 55.3 36.5 14.7 20.3 14.6 10.6 147.3 45.6 Ro 94.6 45.8 13.6 8.0 13.5 7.3 13.0 60.6 40.3 16.o 19.4 15.2 9.7 118.4 42.7 Rp 110.6 49.8 16.0 lox 16.o 10.0 1 5.14 77.5 49.5 16.0 16.1 10.8 11.0 189.2 RQ 95.2 43.8 14.3 9.0 14.0 8,5 13.7 59.3 40.6 15.3 19.7 12.6 10.5 165.3 6456: Rr 99.6 h6.0 15.0 9.0 15.0 8.5 15.0 64.3 37.3 17.6 18 , 3 12.8 10.6 125.7 44.6 ND Rs 89.2 7.3 41.4 13.5 7.5 13.0 lb.o 69.0 40.3 17.3 17.o 13.8 10.3 152.3 1 .0 Rt 97.4 44.0 15.0 9.0 15.0 9.0 14.3 59.3 40.6 15.6 20.0 14.2 10.6 160.3 5;5 05 Ru 102.6 46.0 16.o 8.5 16.0 8.5 15.6 (31.0 56.3 18.0 18.6 13.0 10.7 155.4 51.3 R77. 82.4 36.8 13.3 7.0 13.0 6.5 12.0 55.6 31.6 14.3 21.2 14.1 11.7 156.2 55.2 Rw 93.6 42.2 13.0 7.3 13.0 7.0 13.0 53.0 33.3 17.3 17.2 13.0 10.6 180.3 57.0 Taxon S

a b c d e f r i j k 1 m

Sa 95.2 47.6 16.0 8.0 15.5 8.5 1 6.1_L 70.0 41.0 19.0 18.8 12.9 11.8 133.1 42.7 SID 83.8 45.6 18.0 1C.0 17.5 1000 15.2 60.0 44.0 17.0 19.2 14.0 11.5 107.6 33.9 Sc 105.0 54.0 16.0 12.0 15.5 11.0 17.6 77.0 4.6 21.0 17.0 12.7 12.0 102.5 51.) Sd 87.6 50.4 15.3 11.5 15.0 10,6 15.4 6').0 47.5 18.0 16.4 1 201 9.7 ',69.5 62.5 Se 94.6 53.2 16.0 1 2.5 16.0 1 2.0 17,2 67.3 45.6 21.6 16.5 12.4 10.1 152.2 54.0 SC 82.6 48.4 17.1% 8.0 16,0 7.6 14.6 50,0 32.5 15.5 20.3 14.2 11.8 126.2 ',4.2 3g 86.2 48.0 14.3 9.0 14.0 9.3 1L.4 57.5 37.0 i6.0 16.5 13.0 11.0 195.9 )6.2 Sh 89.8 51.2 16.5 9.5 16.0 9.0 17.0 80.0 60.0 20.0 17.3 13.7 11.5 206.6 ,-;6.6 Si 102.6 55.2 17.0 11.3 17.0 11.0 15.3 60.5 38.6 1700 16.7 13,0 11.3 160.3 52.0 j 90.2 48.0 14.5 8.5 14.3 8.5 17.0 60.0 41-.5 16.3 17.1 13.3 11.5 170.3 60.2 (3,c- 102.4 56.8 18.5 11.0 18.5 1C.6 19.2 5.5 27.0 25.5 16.0 11.3 175.5 60.2 Si 91.6 53.8 1i6,6 14.0 15.0 13.3 14.0 59.0 40.0 17.0 15,2 16.4 9.2 156.0 55.2 93.0 53.2 16.0 12.5 16.0 12.3 16.2 61.6 4 .3 16.7 16.3 11.6 10.1 150.7 55.1 Continuation Taxon S

a 1 Sn 101.4 57.4 16.3 11.6 1 6.5 11.0 17.6 75.3 54.0 19.3 18.3 14.5 11.7 160.3 57.6 so 97.2 5008 15.5 8.0 15.o 8.o 16.0 65.3 42.0 18.5 17.6 12.4 11.7 140.2 453 Sp 88.4 48.3 i 5.0 8.5 1 5.3 8.3 15.3 62.6 41 .3 17.0 17.2 13.7 11.6 1 55.7 52.6 Sc 85.6 47.0 16.0 8.0 16.3 803 55.7 30.3 1 5.0 1 9.6 1L.7 11 .6 130.7 46.3

Sr 86.2 47 6 17.3 9.5 17.0 9.5 16.6 65.3 46.3 17 6 17.6 1 300 11 .3 25.3 34.3 Ss 100.2 52.4 15.3 11.5 15.0 11 3 17.0 73.3 44.6 20.3 16.0 11 .7 11 .7 1 !-,2.3 52.3 St 98.c 56.o 17.0 12.5 17.0 12.0 18.o 62.3 40.3 18.0 17.3 13.1 10.8 10.0 56.0 Su 96.2 53.4 1 5.5 8.0 1 5.0 8.0 1 5.3 67.3 42.6 18.2 17.2 1 3.0 1 2.0 110.6 45.1 Sv

Svc ?axon T

a 1

Ina 98.6 52.6 17.0 11 .5 17.0 11 .3 17.2 67.3 42.0 17.0 16.2 11.8 11.6 155.2 714.3 7b 96.0 49.6 16.3 9.0 16.0 9.0 16.6 67.3 48.0 20.5 19.0 14.5 10.4 141.5 68.5 re 93.0 43.6 16.0 8.3 15.5 8.0 16.3 50.0 45.7 15.5 16.5 12.7 10.8 177.5 85.5 -12d 87.2 44.6 15.0 9.0 14.5 8.5 10.6 62.0 34.6 12.3 20.0 16.0 10.5-173.7 93.7 Te 96.2 51.4 -17.0 11.0 16.5 11.0 15.4 69.0 46.5 18.5 16.6 -13.0 10.5 -178.5 69.4 :f 95.2 49.8 15.6 11 .0 15.0 11 .0 16.8 42.2 21 .0 17.0 14.3 10.3 162.5 72.7 Pg 93.2 145.6 15.0 12.0 15.0 12.3 1 2.C- 78.3 43.6 15.0 23.0 17.0 10.5 138.8 56.8 Th 100.8 52.6 17.6 9.0 18.0 9.0 16.0 63.5 39.0 19.0 18.0 12.0 11.5 152.6 68.1 i 106.2 55.2 17.0 10.0 17.0 10.3 17.3 7)4.0 47.3 24.3 18.3 114.0 10.2 145.9 62.8 96.6 47.8 15.6 8.0 15.5 6.o 16.6 66.0 42.5 19.0 17.9 14.1 9.4188.7 80.2 ik 83.6 39.6 13.6 10.0 13.0 9.6 11.8 61.o 40.5 14.5 19.6 14.9 11.0 148.2 71.7 a 95.2 45.8 16.0 9.5 16.0 9.0 15.2 77.0 40.0 25.0 15.0 12.1 9.5 120.0 54.5 2n 81.8 39.4 13.0 8.0 13.0 3.3 16.6 55.0 34.0 20.5 i6.3 11.6 10.0 122.9 47.7 Continuation Taxon T

a j 1

Tn 101.4 50.2 16.0 8.5 16.0 8.0 15.3 60.0 40.0 15.0 19.5 13.5 10.3 159.0 67.9 To 95.8 49.0 16.0 9.0 16.0 9.0 15.6 68.0 44.5 18.0 17.3 12.7 11.7 145.8 60.0 2p 1 04.2 52.4 5.6 1 2.0 1 5.5 11.5 20.6 66.5 43.5 22.0 1 6.8 13.2 11 .1 1 09.0 44.5 7ci 91.2 44.2 13.6 7.0 13.0 7.3 17.2 64.3 39.0 15.3 17.8 12.6 9.6 135.8 54.5 fr 104.0 49.6 1 7.3 11.0 16.5 10.3 15.0 72.6 38.3 19.6 19.4 /4.4 10.7 143.0 55.6 7s 87.8 46.4 14.3 1 0.5 14.5 9.6 1 2.0 63.3 44.3 1 7.0 16.0 11 .0 9.3 141.2 56.7 2t 93.2 46.6 16.3. 8.5 15.5 8.3 18.0 59.6 42.0 18.3 17.1 12.6 10.4 150.0 61.0 -11 90.4 44.6 17.5 10.5 17.0 10.0 17.0 60.3 41 .3 17.6 17.3 1 2.3 10.0 142.3 56.2 88.2 43.6 1 4.3 9.5 14.5 9.5 15.8 60.o 38.8 19.0 17.0 12.6 9.8 130.7 60.1 97.0 50.o 16.o 9.5 15.5 9.5 16.3 64.0 46.6 20.1 20.1 15.o 10.2 155.3 74.3 7x 90.0 46.2 15.3 9.0 15.0 9.0 1)4.6 63.6 45.3 18.7 17.6 12.1 10.5 130.3 57.0 ;7y 85.8 42.0 14.0 9.5 13.5 9.5 12.6 57.0 38.6 14.3 18.3 13.8 11.0 160.3 74.0 96.4 50.0 16.0 9.5 16.0 9.3 16.o 70.0 47.0 19.0 20.1 15.0 1 0.7 150.1 73.3 94.4 48.6 1 5.0 8.0 1 5.0 8.0 1 4.6 65.6 39.7 1 5.0 21.0 1 5.1 10.5 130.3 57.0 Taxon U

a b c d. e f g h j k

TTa 94.4 54.0 16.3 11 .0 16.0 11.o 16.0 57.0 40.0 17.0 1 9.0 14.1 1 2.3 1 25.8 63.4 b 92.24 50.8 18.3 10.5 18.5 10.0 13.8 65.0 40.0 1 5.5 16.6 13.6 11 .5 106.4 52.1

C 91.2 57.8 17.3 1 2.5 1 7.0 1 2.0 17.2 65.3 148.0 23.3 17.6 1 2.9 10.3 101 .8 63.5 tTd 93.6 62.6 16.6 12.0 17.0 12.0 15..6 68.5 )4)i.2 17.5 17.9 14.9 9.8 116.5 51.7 93.2 53.2 1 6.0 11.0 1 6.3 10.5 17.3 60.o 45.0 1 8.0 1 0.7 1 5.0 11.5 107.6 40.3 Uf 95.8 54.6 17.6 9.5 18.0 9.o 18.o 70.0 46.5 17.5 17.3 13.0 12.0 143.6 57.9 7Jg 82.8 47.0 13.6 10.0 13.5 9.0 16.4 64.0 424.3 23.6 16.5 1 3.4 1 0.1 1 52.4 64.2

;:rh 88.4 49 2 114.6 10.0 14.5 10.0 1b.8 5..3 43.0 20.3 18.6 13.6 10.5 170.5 69.3 104.0 57.6 18.6 11 .0 17.5 11.0 1 9.0 66.0 45.0 17.0 17.0 13.1 11.7 178.3 68.9

Uj 95.2 54.4 1 8.0 11.0 18.0 11 .0 16.0 71.6 46.3 17.6 -16.7 13.2 10.3 107.5 51.o Jk 99.0 56.2 17.3 10.5 17.0 10.3 18.3 65.0 37.0 18.9 1 5.3 11.4 11.8 1 29.1 57.0 is i 83.2 57.8 19.3 11 .0 1 9.5 11.0 1 5.8 52.6 34.6 1 2.6 13.0 10.0 11 .3 1 21.0 49.0 94.0 55.2 17.0 11.0 17.0 11 •3 18.2 82.o 55.0 19.0 1 5.8 12.0 12.1 123.1 48.3 axon U .

a b c d e f g h i j k 1

-fn 90.4 53.4 19.0 11.0 19.0 11.0 16.6 53.0 34.3 16.3 14.0 11 .1 11.5 120.0 47.2 91.0 52.6 18.5 11.5 18.3 11.0 17.0 59.0 41.3 18.0 18.0 13.7 12.0 130.0 60.3 'Pi 95.0 55.0 17.6 10.5 17.0 10.5 16.0 72.0 45.7 18.0 17.0 13.4 10.8 115.0 50.1 7g 100.6 56.4 19.3 11.0 19.0 11.0 19.0 70.3 50.6 17.5 18.1 14.0 11.3 170.6 70.0 Jr 98.0 57.0 18.0 11.5 18.3 11.5 18.4 66.6 49.0 20.6 18.3 13.6 11.8 121.0 59.3 7,s 103.2 60.6 16.0 11.3 15.3 11.3 16.0 6,,.7 46.5 17.0 19.3 14.6 11.3 127.9 51.7 -it 86.8 50.2 1 5.3 1.0.5 15.o 10.0 1 5.8 66.3 42.7 21.0 14.5 i1.5 10.7 148.3 64.o -al 91.2 53.4 16.5 10.5 16.0 10.0 16.3 69.7 46.0 18.7 17.6 14.0 11.8 11 2.3 50.7 Taxon V

a b c d e f g h i j k 1

Va 98.6 )1)1.0 15.6 9.5 15.0 9.0 18.2 63.5 34.5 18.0 19.8 14.5 8.5 149.0 73.8 Vb 82.6 38.4 11.0 7.5 11.0 6.6 9.0 56.0 23.3 11.3 22.1 16.7 11 .4 11 9.5 74.2 Vc 92.2 40.2 14.0 7.0 13.0 7.0 i3.4 71.0 36.0 14.0 16.0 12.5 11 .9 111 .0 55.0 Vd 107.8 W1.8 13.6 8.0 13.5 7.3 18.6 62.3 38.0 17.3 21.0 16.4 10.4 102.0 46.0 ye 94.0 39.0 12.3 6.0 12.5 4.0 16.3 67.2 40.5 17.2 16.8 13.8 11.0 141.2 64.3 ye 98.4 43.6 15.0 9.0 15.0 9.0 17.4 71.5 43.6 21.5 20.6 15.4 9.5 135.7 56.5 Vg 82.0 36.8 13.3 9.0 13.0 9.3 12.6 60.0 38.0 15.0 16.0 12.8 8.3 132.3 55.0 93.4 40.2 14.6 8.0 14.3 8.0 i4.6 50.0 39.5 16.5 16.2 13.1 11.7 107.6 40.4 'Ti 105.6 47.0 16.6 9.0 16.0 8.6 15.4 74.5 44.5 18.0 17.5 14.2 11.3 168.2 71.7 7i 102.4 46.6 14.6 10.5 14.0 10.3 13.8 79.3 41 .1 16.3 14.4 11.0 10.8 175.5 74.8 Ik 97.4 40.2 -15.0 8.5 15.0 8.3 16.6 66.5 35.0 20.0 18.6 13.3 12.1 190.5 77.1 fl. 85.0 38.0 13.5 8.5 13.0 8.3 11.5 70.3 40.3 15.0 20.5 16.0 11.6 140.6 68.3 vm 102.0 45.4 15.3 10.0 15.0 10.0 14.4 74.6 38.6 16.0 15.8 11 .6 11.3 160.3 74.i Continuation Taxon V

a i j k 1

Vn 97.4 44.2 15.0 9.3 15.0 9.0 17.4 65.6 34.0 17.0 20.3 15.1 10.3 155.3 74.9 7o 106.6 45.6 16.3 9.5 16.0 9.3 15.8 68.3 35.6 16.5 18.0 13.7 10.8 110.4 50.3 7p 90.8 40.0 14.0 8.3 14.0 8.0 13.0 77.5 38.8 14.5 17.0 13.1 11.3 135.6 75.6 vq 93.2 4.1.4 13.5 7.5 13.0 7.0 15.4 71.0 40.0 16.6 17.5 14.0 11.2 147.3 66.1 Vr 101 .4 4)1.6 16.o 9.3 16.0 9.3 17.0 74.6 41.6 19.0 19.6 15.1 10.6 130.3 60.2 Is 84.6 37.2 14.0 8.5 14.0 8.0 13.2 65.3 37.0 15.3 16.7 13.6 10.8 131.1 57.6 Vt 87.8 38.0 12.5 7.5 12.0 7.5 11 .4 60.6 29.8 12.5 20.1 15.0 11 .1 140.0 75.1 Taxon W

a b c d e f g h i 3 k 1 m n o

Via 117.0 51.4 16.0 12.0 16.0 11.0 15.6 68.6 38.3 18.0 18.3 12.8 10.2 214.7 50.7 Wb 95.0 42.6 12.0 8.0 12.0 7.5 11.4 81.3 47.0 17.0 21 • 5 14.5 1 0.1 209.3 67.3 Vic 103.6 40.6 13.0 8.0 12.0 7.3 15.0 72.0 33.7 16.0 15.4 11.5 10.8 195.6 53.8 Wd 93.2 43./4 14.6 8.5 15.0 9.0 15.2 50.5 29.0 16.0 16.2 12.0 11.0 157.9 39.4 We 96.6 42.2 14.0 8.0 13.5 7.5 13.0 68.0 34.0 12.5 i7.3 13.5 11.6 214.0 52.0 Wf 93.4 39.6 12.3 7.5 12.0 7.0 13.0 77.0 36.5 20.0 19.5 14.0 10.8 155.5 /43.5 N) wg, 94.2 43.8 13.0 6.3 I 2.5 6.0 13.0 58.5 31 .5 19.0 16.2 1 3.2 11.0 1 91.2 59.2 k.W,4 Wh 96.8 42.0 13.5 7.5 13.0 7.5 13.6 68.0 34.5 20.0 16.6 13.0 10.7 176.0 49.0 wi 107.2 46.4 1)4.3 9.0 15.0 9.0 18.2 71.3 42.0 23.3 18,0 12.7 8.6 205.3 61.2 Wj 111./4 50.0 16.3 9.0 15.0 9.3 1 5.2 71;..2 38.2 1 7.0 21.5 16.7 11.2 190.3 52.1 Wk 114.6 51.2 1 5.5 9.0 1 5.0 9.0 1/4.6 68.3 35.0 1 8.0 20.1 1 6.0 11.4 200.1 54.6 Wi 97.4 44.2 13.0 8.0 12.5 8.0 12.3 73.0 43.6 17.5 19.3 14.0 10.7 201.0 60.1 Wm 105.8 47.6 1)4.0 8.5 114.3 8.5 17.4 73.5 40.6 20.3 19.5 13.1 10.1 210.14 65.5 Continuation Taxon W

a b o d e f g h i j k 1

Wn 110.6 48.2 15.3 11.5 15.0 11.3 15.2 70.3 38.0 17.3 19.6 13.2 10.7 207.1 51.3 Wo 98.0 43.4 14.3 8.5 14.0 8.o 14.0 7(.5 36.3 18.6 16.5 12.1 11.0 160.7 40.1 Wp 98.2 41.0 12.5 8.0 12.3 8.o 14.6 53.6 29.3 16.5 17.1 13.2 11.2 200.0 52.6 dq 96.8 44.2 13.3 7.0 13.0 7.o 13.0 60.3 32.o 20.3 17.0 12.5 11.4 203.6 57.3 wr 97.4 43.6 14.0 7.5 14.0 8.0 14.4 70.3 35.0 13.6 17.5 13.1 11.4 207.1 50.6

Ws 92.6 4i.4 15.3 8.0 15.o 8.0 15.6 0 28.6 16.5 16.1 13.0 10.2 180.3 47.2 -Pr axon X

a b c d e f g h i j k 1 La 85.2 50.4 1 5.6 10.0 1 5.5 10.0 14.6 6)1.6 40.6 11.6 1 5.8 11.6 11.1 145.4 42.0 :a 90.6 51.8 15.3 14.5 15.5 1)4.0 15.2 72.5 46.5 18.0 17.2 13.5 9.4 217.7 66.4 :c 86.2 47.0 15.0 9.5 15.0 10.0 15.0 67.5 )46.5 18.0 19.2 13.7 10.2 136.9 37.5 al 87.8 46.0 1)4.3 8.5 15.0 8.0 15.2 72.0 50.5 18.5 17.6 12.9 10.7 203.5 62.5 _le 53.0 32.4 13.0 7.o 13.0 7.0 12.8 65.0 )1)1.6 16.6 19.0 124.0 9.5 212.0 58.4 f 105.8 61 .14 22.0 17.5 21.0 16.3 15.14 67.5 48.0 15.0 17.7 13.5 10.0 192.3 53.2 103.2 57.4 16.3 11 .5 16.0 11.3 14.8 76.0 50.0 18.0 18.1 13.7 10.1 186.0 53.o a 96.14 55.0 16.0 ii .0 16.0 11 .0 15.4 72.0 50.3 17.3 20.1 14.6 9.7 490.1 56.6 ai 91 .0 53.2 16.3 12.0 16.0 12.0 16.6 75.5 52.5 16.5 16.1 .4 10.3 211 .1 61.9 Ti 100.2 56.8 17.5 12.3 17.0 12.0 15.2 69.3 248.6 16.6 19.3 1)4.2 10.5 180.1 49.7 :k 81.0 45.6 1/t.0 8.3 13.5 8.0 13.0 62.0 40.3 12.5 16.7 12.0 9.9 182.0 57.4 11 95.8 52.4 18.5 13.3 18.3 13.3 17.6 78.6 55.0 13.0 21 .3 15.7 11.3 201 .7 61.2 Lla 88.4 50.0 15.5 10.3 15.5 10.0 11,-.2 63.3 44.6 19.5 18.7 14.1 11 .0 190.1 55.4 Li 86.24 49.6 16.0 10.3 16.0 10.0 15.6 66.0 48.3 17.3 17.3 12.1 10.4 178.0 55•4 ;o 97.6 53.2 16.3 11.0 16.0 11.0 14.2 70.3 50.5 16.5 19.0 13.8 11.6 197.1 60.7