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Cytogenetic Studies in the Genus Iris: Subsection Evansia, Benth. The

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Cytogenetic Studies in the Genus Iris: Subsection Evansia, Benth. The Cytologia 38: 501-514, 1973 Cytogenetic Studies in the Genus Iris: Subsection Evansia, Benth. B. B. Chimphamba1 Department of Botary and Microbiology, University College London, England Received January 25, 1972 Introduction The genus Iris belongs to the family Iridaceae and comprises over 200 species (Lawrence 1953, Lawrence and Randolph 1959) distributed exclusively in the Northern Hemisphere. The subsection Evansia is a small rhizomatous group consisting of 12 known species distributed either in N. America (3 species) or Asia. The Evansias take their name from Thomas Evans of India House, who is said to have introduced I, japonica Thunb. into Europe in 1964. The distinctive feature of the group is the linear crest on the perianth segments, replacing the beard which marks the bearded group. According to Dykes (1913) I. cristata Soland was the first of the Evansias to be recorded in 1764. I. japonica followed in 1794, I. graci lipes Gray in 1859, IL tectorum Maxim. in 1871, I. speculatrix Hance in 1875, I. milesii Fost. in 1883, I. confusa, sealy (originally termed wattii) in 1892 and I. wattii Baker in 1931, whereas I. lacustris Nutt. was accepted generally as a local form of I. cristata Soland. In a recent discussion of the Evansia group by the British Iris Society Species Group (Species Group Bulletin 1966) the 9 most familiar species were arranged into 3 morphologically distinct groups as follows: Group 1. I. confusa (2n=30) I. wattii (2n=30) I. japonica (2n=34, 36, 54) Group 2. I. milesii (2n=26) I. tectorum (2n=28) Group 3 I. cristate (2n=32) I. gracilipes (2n=36) I. lacustris (2n=42) I. speculatrix (2n=44) This grouping is followed in the present investigations and the additional species I. formosana Ohi, which was recently introduced into Europe in 1969 is included in group 1. The 2 species, I. cristata and I. lacustris are restricted to E.N. America and the other 8 species are restricted to E. Asia. In addition to the 8 Asiatic species is I. pseudorossii Chien, a less well known species and which was not available during the present study. The evansias are of interest for several reasons. Firstly 1 Present address: Department of Biology, Chancellor College, University of Malawi. 502 B. B. Chimphamba Cytologia 38 there is very little evidence apart from the crest that the subsection is phylogenetically a natural group. Secondly the subsection has been the recipient of an additional species in recent years as a result of a transfer of the W.N. American species I. tenuis Wats (Lenz 1959) from the series Oregonae of the subsection apogon. This transfer was based solely on morphological characters. The Evansias are also of particular interest cytogenetically because with the exception of one pair of species (I. confusa and I. wattii) and possibly a second pair (I. tectorum and I. tenuis) the chromosome numbers of the various species as compiled by Darlington and Wylie 1955 and as determined by Longley 1928; Simonet 1932, 1934; Randolph 1959 and Randolph and Mitra 1959, are different and form an interrupted aneuploid progression from 2n=26 to 2n=44 with one species having 2n=54. Cytological affinities within the group need therefore to be determined-not only from a com parison of the karyotypes but also from meiotic observations in hybrids. Matherial and methods The plants used in the present study were obtained as transplants or as seed from Botanic Gardens. The sources of the materials were as follows in Table 1. Methods Somatic chromosome numbers were determined from root-tips stained by Feulgen squash method described by Darlington and La Cour (1942). Actively growing root-tips were pre-treated with 0.1% aqueous colchicine solution for 3 to 5 hours, fixed for 12 to 15 minutes in freshly prepared acetic alcohol (3 parts absolute alcohol: 1 part glacial acetic acid), stored overnight in 95% alcohol, hydrolyzed in N HCl at 60•Ž for 8 to 10 minutes and stained in leuco-basic fuchsin for 2 hours. Squashing was carried out in a drop of 45% acetic acid or if necessary in a drop of aceto-carmine in order to increase the intensity of staining in the chromo somes. Slides were temporarily sealed with rubber solution and stored for up to 3 days at about 0•Ž. They were made permanent by the quick-freeze method of Conger and Fairchild (1953). For meiotic study the aceto-carmine fixing and staining technique described by Thomas (1940) was followed. Photographs were taken with a Leitz Orthomat camera using a Leitz planar apochromatic •~100 oil immersion objective. The film was Kodak Recordak Microfile which was developed in Kodak D8 extreme contrast developer for 2 minutes at 68•‹F. For the purpose of cross-fertilization it was often necessary to store pollen. This was done by removing matured anthers and allowing them to dehisce in petri dishes. The pollen was then stored in vials stopped with cotton wool and kept in desiccators. Fresh pollen was used as far as possible. The anthers of the species used in the crossing programme dehisced shortly after the bud opened, and except for a few members of the species all were self compatible. Emasculation was there fore essential and this was carried out on the day before the flower would have 1973 Cytogenetic Studies in the Genus Iris: Subsection Evansia, Benth. 503 Table 1. Materials and sources * The complete addresses of the donors of the above materials can be obtained from The Iris Year Book 1967. 504 B. B. Chimphamba Cytologia 38 opened. The buds were bagged to avoid insect contamination. Pollination was achieved by dusting the pollen onto the lip of the style with a clean camel hair brush. Results a) Cytology Ten Evansia species were available for this comparative cytological survey, and their chromosome numbers as determined in this investigation and by previous workers are listed in Table 2 where the species are arranged in groups on the basis of their morphological similarities. There is good agreement between the chro mosome counts from this investigation with those reported by earlier workers except for 2 species in group 1, IL formosana and I. japonica. The former species was Table 2. Chromosome number in 10 Evansia species recently imported (1969) into Europe from Formasa as representing the typical wild form of this species, but in view of the different and uneven chromosome number of 2n=35 it is doubtful whether this is so. In the case of I. japonica the count of 2n=54 agrees with previous determinations, and this chromosome number was found in the hardy type known horticulturally as Ledger's variety and in material recently imported from Japan. On the other hand the 2 counts of 2n=31 and 2n=33 have not previously been recorded. Both numbers have been found in non hardy forms grown as ornamentals, and in forms obtained by Marchant from Japan as representing the wild species. No types of I. japonica were encoun tered with the previously reported chromosome numbers of 2n=34 and 36. The somatic chromosomes varied in length in the various species from 1ƒÊ to 7.5ƒÊ. The smallest chromosomes were recorded from I. lacustris and the largest 1973 Cytogenetic Studies in the Genus Iris: Subsection Evansia , Benth. 505 chromosomes from I. wattii and I. cristata . Meiotic study was made in 5 species (I. confusa, I. wattii, I. japonica, I, milesii and I. tectorum) and all except I. wattii and I. japonica, had regular pairing of chro mosomes and chromosome segregation at anaphase was equal. Both I. wattii and the 2 cytotypes of I. japonica (2n=54 and 2n=31) had trivalunts and univalents at metaphase I apart from the ordinary bivalents (Table 3) and this chromosome ir regularity corresponded with the difficulties experienced in pairing the somatic chromosomes in these species. Except 2 species (I . wattii and I. japonica) all had normal pairing of chromosomes and segregation at anaphase was equal . Table 3. Meiotic chromosome pairing in 2 Evansia species For comparison the idiograms of all the species investigated are shown in Figs. 1 and 2. For the species in group 1 the complete chromosome complement is represented because of the difficulties in pairing the chromosomes in most of the types. For the species in groups 2 and 3 where pairing was possible only -the haploid complements are represented. The main features evident from the idio grams can be summarized as follows: 1) The karyotypes of I. confusa and I. wattii show more similarity to each other than any other pair of Evansia species. This affinity was later confirmed from the hybridization studies which are discussed in the next section. 2) There was no evidence in groups 2 and 3 that external morphological simi larities were paralleled with karyotypic similarities. In group 2 for instance, the karyotype of I. tectorum showed more resemblances to some chromosomes in the 2n=54 cytotype of I. japonica than it did to I. milesii while I. milesii showed more correspondence with I. cristata in group 3. Also in group 3, I. lacustris showed more similarities to I. speculatrix than to I. cristata, yet in this instance the 506 B. B. Chimphamba Cytologia 38 Fig. 1. Idiograms of somatic chromosomes in Evansia species. Fig. 2. Idiograms of somatic chromosomes in Evansia species. 1973 Cytogenetic Studies in the Genus Iris: Subsection Evansia , Benth. 507 Fig. 3. Somatic and meiotic chromosomes in I. confusa and I. wattii. a, somatic metaphase in I. confusa, 2n=30, b, somatic metaphase in I. wattii, 2n=30. c, diakinesis in I. confusa, showing 15 bivalents and a nucleolus (arrowed).
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