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Biosystematic Studies on the Genus Polygonatum (Liliaceae) I

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Biosystematic Studies on the Genus Polygonatum (Liliaceae) I _??_1990 by Cytologia, Tokyo Cytologia 55: 443-466 , 1990 BiosystematicStudies on the Genus Polygonatum (Liliaceae) I. Karyotype analysis of species indigenousto Japan and its adjacent regions Minoru N. Tamura Department of Botany, Faculty of Science, Kyoto University, Kyoto 606, Japan Accepted March 9, 1990 The genus Polygonatum (Liliaceae-Polygonateae or Convallariaceae-Polygonateae) includes ca. 58 species and is widely distributed in the northern hemisphere, from the sub tropical to subarctic zone (Bentham and Hooker 1883, Krause 1930, Hutchinson 1934, Dahl gren et al. 1985). According to Tang (1978), the species referred to three series, i.e., Ser. Bracteata, Ser. Alternifolia and Ser. Alte-lobata, occur in Japan, Taiwan and Cheju Island, Korea. The plants of Ser. Bracteata are distributed in northern and northeastern China, Far Eastern U. S. S. R., Korea and Japan, and include 6 species and 2 varieties, among which 5 species and 2 varieties occur in Japan and Cheju Isl. Among the species belonging to this series, chromosome numbers and karyotypes of P. desoulavyi var. desoulavyi and P. involu cratum from China, U. S. S. R. and the Korean Peninsula have been reported (Sokolovskaya 1966, Lee 1967, Abramova 1971, Kim and Kim 1979, Wang et al. 1987), but no reports have been made for the plants in Japan and Cheju Isl. The remaining species of this series are so far unknown karyologically (cf. Table 1). Ser. Alternifolia, which comprises ca. 35 species, is the largest group of the genus, and is distributed in Europe, Asia and North America, among which 8 species, 6 varieties and I form occur in Japan, Taiwan and Cheju Isl. Detailed karyotype analyses have been made on the North American as well as European taxa belonging to this series by various authors (Eigsti 1942, Suomalainen 1947, Therman 1950, Kawano and Iltis 1963b, Nowakowska and Zeglicka 1972, for others see Table 1). However, only scattered information has been avail able for the plants from northeastern Asia. The Bracteata group and the Alternifolia group have been studied taxonomically by Franchet and Savatier (1878), Komarov (1935), Satake (1942), Abramova (1975) and Tang (1978). However, taxonomic concepts of these two groups by the above authors (1. c.) are different, especially as to the identity of P. inflatum. Abramova (1975) used chromosome num bers and karyotypes for the delimitation of these groups in addition to gross morphology. But, her results on chromosome numbers and karyotypes for the Japanese species are different from those of the other authors (cf. Table 1). The purpose of the present study first, is to report the detailed chromosome morphology of all the species belonging to Ser. Bracteata and Ser. Alternifolia occurring in Japan and its adjacent regions through a critical examination of many individuals from different localities, and second, to revise taxonomically these two series based on the results obtained. Materials and methods The sources of all materials examined are given in Table 2. The plants collected from their native habitats were transplanted into clay pots and cultivated in the Department of 444 Minoru N. Tamura Cytologia 55 Table 1. Present and previous cytological studies of the genus Polygonatum * These numbers are first counted . ** Judging from the description of gross morphological characters , this plant should be referred to P. inflatum. 1990 Biosystematic Studies on Polygonatum (Liliaceae) I. 445 Botany, Kyoto University for more than two months . The root tip preparations for the examination of the chromosomes were made by using a modification of the acetic orcein squash method by Kawano and Iltis (1963a) . Excised root tips were stored in 0.2% aqueous colchicine solution at 17•Ž for 4h , then fixed in Farmer's solution for 1-2 min, and subsequently stained in 1% aceto-orcein for ca . 12h, then preserved in an aceto-orcein (1%) -IN hydrochloric acid (1:1) mixture for a minimum of 15 min . Each of the root tips was transferred onto a glass slide , treated with a drop of acetic acid (45%) glycerine mixture (9:1) for ca. 10 sec, and then gently heated over a flame and squashed. Voucher specimens are preserved in the Herbarium of Kyoto University (KYO) . The karyotype descriptions were made according to Levan et al . (1964). The "basic karyo type" examined in the present study was expressed in terms of homologous chromosome pairs, and thus some uncertain or unstable secondary or small constrictions and also B-chro mosomes were excluded from the karyotype formula. Results Chromosomes at mitotic metaphase were examined (Fig. 1), and the results on each taxon were as follows. 1. Polygonatum cryptanthum Lev. et Van. -2n=18 (Figs. 2 and 9) This is the first count of chromosome numbers for this taxon. Plants collected from two localities, i. e., Mt. Ariake and Kamisaki Cape, in Tsushima Isis., Nagasaki Pref. all proved to be 2n=18, that is, diploid. The karyotype was composed of one long pair of chromosomes (ca. 10.5 um in length), six medium-sized pairs (ca. 8.2ƒÊm-ca. 5.9ƒÊm) and two short pairs (ca. 3.9ƒÊm). One pair of the long chromosomes had median centromeres. One of the six medium-sized pairs was also metacentric, but the remaining five pairs were submetacentric. One of the two short pairs had median centromeres, while the other pair was submetacentric. One medium-sized pair of metacentric chromosomes (the 2nd pair) and the other medium sized pair with submedian centromeres (the 6th pair) had secondary constrictions at the distal regions of the long arms. The karyotypes from the two localities were identical. 2. Polygonatum falcatum A. Gray var.falcatum-2n=18 (Figs. 2 and 9) Plants collected from ten localities (Table 2) were all diploid with 2n=18 chromosomes, which agreed with previous reports (Table 1), although 2n=20 was also reported for this taxon. The karyotypes of these plants were identical with one another, and basically the same as that of P. cryptanthum described above. 3. Polygonatum falcatum A. Gray var. hyugaense Hiyama-2n=18, 2n=18+4B-8B (Figs. 3 and 9) Plants from two localities, Mts. Iwaudo and Sekigan, Kumamoto Pref., were first ex amined karyologically in the present study. Plants from Mt. Iwaudo proved to be 2n=18, diploid, and those from Mt. Sekigan were also diploid, but with B-chromosomes, 2n=18+ 4B-8B. The number of B-chromosomes varied from 4 to 8 in the same individual. The basic karyotype of this variety was identical with that of var. falcatum. 4. Polygonatum involucratum (Franch. et Savat.) Maxim. -2n=18 (Figs. 3 and 9) Plants from four localities, Irimizu, Fukushima Pref., Daibosatsu Pass and Koarama, Yamanashi Pref. and Koidzumi, Shiga Pref. were karyologically examined. All plants were diploid with 2n=18 chromosomes, which is in agreement with previous reports (Table 1). Table 2. Sources of materials 1990 Biosystematic Studies on Polygonatum (Liliaceae) 1. 447 448 Minoru N. Tamura Cytologia 55 However, Lee (1967) reported two different chromosome numbers, 2n=20 and 22, for this taxon from Korea. The basic karyotype of this species was identical with those of P. crypt anthum and P. falcatum. Fig. 1. Somatic chromosomes of Polygonatum species. A, P. falcatum var. hyugaense from Kuma moto: Mt. Sekigan. B, P. involucratum from Yamanashi: Koarama. C, P. odoratum var. maxi mowiczii from Hokkaido: Abashiri. D, P. macranthum from Yamagata. E, P. desoulavyi var. azegamii from Tokyo. F, P. miserum from Nagano: Narakawa. G, P. domonense from Fukushima. Detailed localities shown in Table 2. Bar=5ƒÊm. 1990 Biosystematic Studies on Polygonatum (Liliaceae) I. 449 Fig. 2. Somatic chromosomes of Polygonatum species. 1, P. cryptanthum from Tsushima Isis.: Mt. Ariake; 2, from Tsushima Isis.: Kamisaki Cape. 3, P. falcatum var. falcatum from Yamanashi; 4, from Mie; 5, from Yamaguchi; 6, from Fukuoka; 7, from Ohita; 8, from Kumamoto; 9, from Nagasaki: Mt. Gongen; 10, from Tsushima Isis.: Mt. Shiradake; 11, from Tsushima Isis.: Kami saki Cape; 12, from Cheju Isl. Detailed localities shown in Table 2. Bar=5ƒÊm. 450 Minoru N. Tamura Cytologia 55 Fig. 3. Somatic chromosomes of Polygonatum species. 13, P falcatum var. hyugaense from Kuma moto: Mt. Iwaudo; 14, from Kumamoto: Mt. Sekigan; 15, P. involucratum from Fukushima; 16, from Yamanashi: Daibosatsu Pass; 17, from Yamanashi: Koarama; 18, from Shiga. 19, P. tri chosanthum from Nagasaki: Mt. Gongen (A); 20, from Nagasaki: Mt. Gongen (B); 21, cultivated in Tokyo Univ. 22, P. humile from Hokkaido; 23, from Aomori. Detailed localities shown in Table 2. Bar=5ƒÊm. 1990 Biosystematic Studies on Polygonatum (Liliaceae) I. 451 Fig. 4. Somatic chromosomes of Polvgonatum species. 24, P. humile from Niigata; 25, from Shiga. 26, P. lasianthum var. lasianthum f. lasianthum from Yamagata; 27, from Fukushima; 28, from Nagano; 29, from Hiroshima; 30, from Okayama. 31, P. lasianthum var. lasianthum f. arnabile from Saga. 32, P. lasianthum var. coreanum from Tsushima Isls.; 33, from Cheju Isl.: Mt. Hanla (A); 34, from Cheju Isl.: Mt. Hanla (B). Detailed localities shown in Table 2. Bar-5ƒÊm. 452 Minoru N. Tamura Cytologia 55 Fig. 5. Somatic chromosomes of Polygonatum species. 35, P. odoratum var. maximowiczii from Hokkaido: Abashiri; 36, from Hokkaido: Haboro; 37, from Hokkaido: Mt. Kannoniwa; 38, from Hokkaido: Tomakomai. 39, P. odoratum var. pluriftorum from Fukushima; 40, from Ibaraki; 41, from Sado Isl.; 42, from Yamanashi: Koarama; 43, from Nagano; 44, from Fukui: Yokogaki; 45, from Fukui: Awara; 46, from Oki Isis.; 47, from Okayama. Detailed localities shown in Table 2. Bar=5ƒÊm. 1990 Biosystematic Studies on Polygonatum (Liliaceae) I. 453 Fig. 6. Somatic chromosomes of Polygonatum species. 48, P. odoratum var. pluriflorum from Yamaguchi; 49, from Kumamoto; 50, from Cheju Isl.; 51, from Yamanashi: Daibosatsu Pass. 52, P. odoratum var. thunbergii from Ishikawa; 53, from Fukui: Namimatsu; 54, from Fukui: Gamo.
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