植物研究雑誌 J. J. Jpn. Bo t. Originals Originals 79: 271-280 (2004)

Chromosome Numbers of Fifteen Species of the Genus Saussurea DC. (Asteraceae) (Asteraceae) in the Himalayas and the Adjacent Regions

Kazumi FUJIKA WA ぺHiroshi lKEDA¥ Katsuya MURATA b， Takaaki KOBAYASHl b， b a Tatsuaki Tatsuaki NAKANO ， Hideaki OHBA and Su-Gong WU c

aUniversity aUniversity Museum ， The University of Tokyo ， Hongo Hongo 7-3- 1， Bunkyo-ku ，Tokyo 113-0033 JAPAN; *Present *Present address: The Kochi Prefectural Makino Botanical Garden ， Godaisan Godaisan 420 0- 6，Kochi-city ，Kochi 781-8125 JAPAN; bDepartment bDepartment of Biosphere-Geosphere System Science ，Faculty of Informatics ， Okayama Okayama University of Science ，Ridai-cho 1-1 ，Okayama 700c0005 JAPAN; 'Kunming 'Kunming Institute of Botany ，Chinese Academy of Science ，Kunming 650204 ，Yunnan ， CHINA (Received (Received on March 17 ，2004)

Somatic Somatic chromosome numbers of 15 species belonging to the genus Saussurea col- lected lected from the Himalayas and the adjacent regions are reported. The chromosome num- bers bers of 9 species were examined the for first time. Saussure αleontodontoides was found to to be diploid with 2n = 30; S. globosa ，S. graminea ，S. graminifolia ，S. obv αllata and S. spicata spicata to be diploid with 2n = 32; S. delavayi ，S. forrestii ，S. kingii ，S. medusa and S. yakla yakla to be diploid with 2n = 34; and S. nishiokae ，S. hookeri and S. tridactyla to be dip- loid loid with 2n = 36. The chromosome number of S. hieracioides was 2n = 64 ， which is the first first record of a tetraploid with x = 16 in the genus. A new record of Saussurea nishiokae from from Tibet (Xizang ， China) was confirmed ， which was recognized during our research.

Key words: Asteraceae ， chromosome number ， Himalaya， Saussurea ， Saussurea nishiok4 αe.

The genus Saussurea DC. is distributed Straube (2003) and Kita et al. (in press) sug- mainly mainly in temperate and subarctic regions of gested Saussurea is p紅 aphyletic ，and have Eurasia ，and comprises about 400 species confirmed that Saussurea contains two major classified classified into six subgenera (Lipschitz clades; one is composed of a large number of 1979). 1979). The genus is characterized by its lack Saussurea species including S.α lpina which of of spines on plant bodies; solitary ， is the type species of the genus; the other corymbose or panicu1ate capitula composed consists of section Elatae and subgenera of of red ，purple ，violet ， pink or white disc 臼0- Jurinocera and Frolovia ，which is the sister rets; rets; spineless multi-seriate involucral group to the genus Jurinea. Besides to these phyllaries; phyllaries; usually setaceous receptacles; lin- molecular analyses ， the investigations by e征 branched styles; obovoid to oblong or 4- Raab-Straube (2003) and Kita et al. (in 5-gonal 5-gonal achenes; and usually bi-seriate press) into morphological characters of pappus ， the outer being composed of short ， achenes and pappi show that am 司or group free free scabrid bristles and the inner ， long of Saussurea is distinct from sections Elatae plumose bristles that are connate at the base and Jurinocera which having single pappi (Bremer 1994 ， Haffner 2000). The recently and ， achenes with a horn or dentate crown. molecular molecular phylogenetic analyses of Raab- Raab-Straube (2003) applied the results to

-271- 272 植物研究雑誌第79 巻第5号 平成 16 年10 月 the the generic concept of Saussurea ， and estab- (Mehra et al. 1965 ， Shetty 1967 ， Koul et al. lished lished the new genus Himalaiella ， and the 1976 ， Jee et al. 1983 ，Koyama and Shimizu ， genera genera Frolovia and Lipschitziella are resu ト 1984 ， Gupta and Gill 1989 ， Gupta et al. rected. rected. As such ， the systematics of the genus 1989 ，Kondo et al. 1990 ， Siddique and Wafai is is now considered. 1993 ， Liu et al. 1994 ，Huang et al. 1996 ， The taxonomy of Saussurea species has Amano and Ohba 2000 ， Fujikawa and Ohba proved proved to be problematic due to wide ranges 2003a) ，nevertheless about 160 species are ofmo 叩hological variation and high species known from this region (based on Lipschitz ， diversity. diversity. Species delimitation has been 1979) ， and one of the diverse genera in the based based on a limited number of gross morpho- alpine zone (Ohba 1988). For clarifying cy- logical logical characters (Clarke 1876 ， Hooker tological features of Saussurea in the 1881 ， Kitamura 1969 ，Lipschitz 1979). Since Himalayas and adjacent regions ，more exten- the the latest monograph by Lipschitz (1 979) ， sive cytological investigations on more spe- the the genus has been studied only in floristic cies from various localities 訂 e required. treatments treatments (Chater and Kitamura 1982 ， Hajra The aim of this paper is to report on the 1995 ，Koyama 1995 ， Shih and Jin 1999 ， chromosome numbers of Saussurea species Springate Springate 2001). Although micro- an d/ or flo- collected from various localities in the ral-morphological ral-morphological observations have been Himalayas and the adjacent regions.

made by Saklani et al. (2000) on Indian spe 田 cies ，by Fujikawa (2002) on the Himalayan Materials and Methods species species and by Yang et al. (2003) in a phar- Plants were collected from their native macognostical macognostical study on the species of sec- habitat in the Nepal Himalaya in 1995 ，1997 ， tion tion Eriocoryne ， it still remains desirable to 1999 and 2000 (see Miyamoto 1995 ， clarify clarify issues regarding species delimitation. Wakabayashi 1998 ， Fujikawa 2000) ， Sikkim Previous Previous cytological studies of the genus Himalaya in 2003 ， and in China (including Saussurea Saussurea have been reported by many Tibet) in 1996 ，1997 ，2000 ， and 2001 (see authors ， and the chromosome numbers of ap- Ikeda and Miyamoto 1996 ，1998 ，Miyamoto proximately proximately 104 species have been revealed ， and Ikeda 2001 ，Akiyama and Ikeda 2002). indicating indicating a wide range of variation: 2n = 24 ， Localities and voucher specimens are listed 26 ，28 ，30 ，32 ，34 ，36 ，38 ，39 ，42 ，48 ，50 ，52 ， in Table 1. 54 ，72 and 78 (see F吋ikawa and Ohba Chromosome counts were made on so- 2003b). 2003b). Judging from the numbers ， the genus matic metaphase cells using the squash tech- Saussurea Saussurea is a cytologically diverse consist- nique. Twenty-four populations of 15 species ing ing of diploid ，tetraploid ， and their aneuploid were analysed; within each population at species species with serial basic chromosome num- least two individuals were examined. Shoot bers bers x = 12 ，13 ，14 ，15 ，16 ，17 ，18 ，19 ， and meristems were fixed with Newcomer's fluid 21 ， although the numbers with x = 12 ，19 ， (see Sharma and Sharma 1980) in the field and and 21 were considered to be doubtfu l. after pre-treatment in 0.05 % cholchicine or Therefore ，polyploidization and aneuploidi- 0.002 M 8-hydroxyquinoline solution for zation zation may play an important role in specia- about 3-4 hours. In order to macerate cell tion tion of Saussurea. walls ， the samples were soaked in a solution However ，cytological evidence is quite of 2 % pectinase and 4 % cellulase (1: 1) for limited limited both in species number and localities 20 minutes-3 hours at 37 0 C having been for for the Himalaya and the adjacent regions. cleaned with an alcohol series and distilled There There are only 28 species for which cyto- water. They were then stained with 2 % logical logical information has been published aceto-orcein for 12-20 hours or 2 % lacto- 02 。σRN 。。 Table Table 1. Sources of materials of 15 species of Saussurea D C. hF

Species Species Locality Voucher specimen

Subgenus Eriocoryne Hoo k. f. s. s. medusa Maxim. China: Sichuan ， around Wuming Shan ，4600 m Wu & al. 19 China: China: Yunnan ，Deqin Xian ， Meili Xueshan ，4700 m Wu & al. 103122 China: China: Xizang (Tibet) ，Baxoi Xian ， around Rawu ， ca. 4000 m Akiyama & al. 105655 s. s. nishiokae Kitam. Nepal: Solukhumbu Dist r.， Panch Pokhari-Seto Pokhari ，5200 m Miyamoto & a l. 9580324 China: China: Xizang (Tibet) ，Nyalam Xian ，Nyalam- Tashigang ，4870 m Akiyama & al. 106293 』。 s. s. spicata Ki tam. Nepal: Mustang Dist r.， around Yak Kh arka ，4470 m Fujikawa 9920119 zgmw s. s. tridactyla Sch. Bip. ex Hook .f China: Xizang (Tibet) ，Nyalam Xian ，Nyalam- Tashigang ，4870 m Akiyama & al. 106291 目。同 s. s. kingii C .E. C. Fisch. China: Xizang (Tibet) ，Nagarze Xian ，Nagarze-Karo La ，4500 m Akiyama & al. 106244 E s. s. delavayi Franch. China: Yunnan ， around M t. Chang Shan ，3720 m Wu & al. 86 宮口 Subgenus Amphilaena (Stscheg l.) Li psch. 20 s. s. globosa Chen China: Sichuan ，Haizi ， near Daocheng ，4460 m Wu & al. 1516 図。窓口可〈。 s. s. hookeri C. B. Clarke Nepal: Solukhumbu Distr. ，Panch Pokhari-Seto Pokhari ，5080 m Miyamoto & a l. 9588174 s. s. obvaZZata (DC.) Edgew. China: Xizang (Tibet) ， Nyingchi Xian ， Sezhaila Shan-Bayizhen ，4500 m Akiyama & a l. 104745 Nepal: Nepal: Solukhumbu Distr. ， Tashing Dingma-Thuli Kharka ，3530 m Wakabayashi & a l. 9730354 --J3z Subgenus Saussurea s. s. forrestii Decne. China: Xizang (Tibe t)， Nyalam Xian ，Nyalam-Zhangmu ，3710 m Akiyama & al. 106310 。・ s. s. gramine αDunn China: Xizang (Tibet) ，Baxoi Xian ， Anjiu La-Rawu ，4430 m Akiyama & al. 105098 u s. s. graminifolia Wall. ex D C. China: Xizang (Tibet) ，Nyalam Xian ，Nyalam-Tashigang ，5170 m Akiyama & al. 106292 Nepal: Nepal: Solukhumbu Distr. ， around Khare ，4780 m Wakabayashi & a l. 9730300 Nepal: Nepal: Solukhumbu Dist r.， Khare-Tangnag ，5010 m Wakabayashi & a l. 9730305 s. s. hieracioides Hook 工 Nepal: Solukhumbu Dist r.， around Tangnag ，4200 m Wakabayashi & a l. 9730232 Nepal: Nepal: Solukhumbu Distr. ， around Tangnag ，4470 m Wakabayashi & a l. 9730242 Nepal: Nepal: Solukhumbu Distr. ，Tangnag-Dig Kharka ，4360 m Wakabayashi & a l. 973025J s. s. leontodontoides (DC.) Hand.-Mazz. China: Xizang (Tibet) ，Baxoi Xian ， Anjiu La-Rawu 4430 m Akiyama & a l. 105097 s. s. yakla C. B. Clarke Nepal: Solukhumbu Distr. ，Chatarwa-Kurke ，4005 m Miyamoto & a l. 9588270 Nepal: Nepal: Solukhumbu Distr. ，Chalem Kh arka- Kh ola Kh arka ，3430 m Wakabayashi & a l. 9730121

NJ 可凶 274 植物研究雑誌第79 巻第5号 平成16 年10 月 propionic propionic orcein for several minutes and are described below. squashed. squashed.

V oucher specimens for all populations in 四 Subgenus Eriocoryne (DC.) Hoo k. f. cluded cluded in this study 訂 e deposited in the 1. Saussurea medusa Maxim. Herbarium Herbarium at the University of Tokyo (TI). The somatic chromosome number was 2n 2n = 34 (Fig. lA). The chromosome counts Results Results and Discussion made in this study disagree with a previous The chromosome numbers of 24 popula- count of 2n = 28 by Huang et al. (1 996). We tions tions representing 15 species of the genus examined the chromosome number of mate- Saussurea Saussurea observed in the present study and rial collected from three populations in previously previously published counts are shown in Yunnan ， Sichuan and Tibe t. All counts of the Table Table 2. Chromosomes as observed in a so- chromosomes in all of these populations

matic matic cell of each species 征 e shown in Fig. were the same ，2n = 34. This disagreement in 1. 1. The chromosome numbers of each species chromosome number could be explained by

Table Table 2. Present and previous studies on 15 species of Saussurea DC.

Present Present counts Previous counts Species Species References (2n) (2n) (n) (2n)

Subgenus Subgenus Eriocoryne Section Section Eriocoryne s. s. medusa Maxim. 34 28 Huang et al. 1996 S. S. nishiok αe Ki tam. 36 36 Amano and Ohba 2000; Fujikawa Fujikawa and Ohba 2003 S. S. spicata Kitam. 32 S. S. tridactyla Sch. Bip. ex Hook 工 36 36 Amano and Ohba 2000; Fujikawa Fujikawa and Ohba 2003 Section Section Pseudoeriocoryne S. S. kingii C. E. C. Fisch. 34 Section Section Cincta S. S. delavayi Franch. 34 Subgenus Subgenus Amphilaena S. S. globosa Chen 32+3B S. S. hookeri C. B. Clarke 36 36 Amano and Ohba 2000 S. S. obva l! ata (DC.) Edgew. 32 ，32 + 3B c.16 ， 16 32 Shetty 1967; Gupta and Gill Gill 1988 ， 1989; Gupta et l. al. 1989; Amano and Ohba 2000 Subgenus Subgenus Saussurea Section Section Jacea S. S. forrestii Decne. 34 Section Section Pycnocephala S. S. graminea Dunn 32 + 1B S. S. graminifolia Wall. ex D C. 32 32 Amano and Ohba 2000 Section Section Saussurea S. S. hieracioides Hoo k. f. 64 Section Section Cyathidium S. S. leontodontoides (D C.) Hand.-Mazz. 30 S. S. yakla C. B. Clarke 34 October October 2004 Joumal of Japanese Botany Vo 1. 79 No. 5 275 either either misidentification of plant material or covering of brown-black hairs on its miscounting miscounting of the chromosomes in the ear- capitula ， and by its brown-black coloured lier lier study. Saussurea medusa is widely dis- pappus. Though Springate (2001) refers all tributed tributed in the Himalayas ， from Central Asia material from Bhutan to S. spicata ，all speci- to to S.W. China ， and exhibits great morpho- mens from Bhutan seen by the first author in logical logical variation throughou t. There therefore the course of this and other research were remains remains a need for more sampling across the identified as S. l αniceps. Saussurea spicata is range range in order to better cl 訂 ify the systema- therefore considered endemic to the Western tics tics of the taxa ， and so confirm possible pat- p紅 t of the Nepal Himalaya ， while S. terns terns of cytological variation. laniceps is distributed throughout the Bhutan 2. 2. Saussurea nishiokae Ki tam. Himalaya ， Tibet and S.W. China. The somatic chromosome number in this 4. Saussurea tridac 砂la Bip. Sch. ex Hook. f. species species was 2n = 36 (Fig. 1B). Counts made The somatic chromosome number in this from from two populations collected from the species was 2n = 36 (Fig. 1D). Chromosome Tibet Tibet and Nepal Himalaya agreed with a pre- counts from Tibet made in this study support vious vious report collected from the Nepal previous reports for the plants in Nepal Himalaya Himalaya by two of the authors (Fujikawa Himalaya (Amano and Ohba 2000 ，F 吋ikawa and and Ohba 2003a). Saussurea nishiokae was and Ohba 2003a). This species is widely described described by Ki tamura (1 969) based on col- distributed in the Himalayas and Tibe t. lections lections from Bhutan and western Nepal ， and 5. Saussurea kingii C. E. C. Fisch. the the species has since been regarded as occur- The somatic chromosome number was ring ring only in the Himalayas. However ，a col- 2n = 34 (Fig. 1E). This is the first record of lection lection of S. nishiokae was made in Tibet a chromosome count in this species. (Xizang ， China) by the second author in Saussurea kingii belongs to section Pseudo- 2000 (see Miyamoto and Ikeda 2001). The eriocoryne ， and is distributed in Tibet and plants plants collected on this occasion bore arach- the Bhutan Himalaya. noid noid hairs on leaves ，3 -4 -seriate involucral 6. Saussurea delavayi Franch. phyllaries ，clavate papillae on the surface of The somatic chromosome number in this the the achenes ，alveolate receptacles and a bi- species was 2n = 34 (Fig. 1F). The chromo- seriate seriate pappus. This combination of charac- some number was examined for the first ters ters distinguished the plants from the closely time. Saussure α delavayi is distributed in related related S. gyacaensis ， which has naked re- Yunnan ， and belongs to section Cincta of the ceptacles ceptacles and a uni-seriate pappus (though subgenus Eriocoryne. also also described from Tibet (Liu 1984)) ， and confirms confirms the collection as a new record of Subgenus Amphilaena (Stscheg l.) Lipsch. S. S. nishiokae from Tibe t. 7. Saussurea globosa Chen 3. 3. Saussurea spicata Kitam. The somatic chromosome number was The somatic chromosome number in this 2n = 32 with three B-chromosomes (Fig. species species was 2n = 32 (Fig. 1C). This is the 1G). The somatic chromosome number was first first time a count has been reported for this examined for the first time. This species is

species. species. Saussurea spicat αhas， sometimes distributed in China (Qinghai ，Gansu ， Shensi been been confused with S. laniceps on account of and Sichuan). its its spicately arranged capitula distributed 8. Sα ussurea hookeri C. B. Clarke along along the upper p紅 t of the clavate stem ，a The somatic chromosome number in this character character shared by the two species. species was 2n = 36 (Fig. lH). This chromo- However ， S. laniceps is distinguished by a some number agrees with that reported in a 276 276 植物研究雑誌第79 巻第5号 平成16 年10 月

騨覇 F

F

Fig. Fig. 1. Somatic chromosomes of fifteen species of Saussurea. A: Saussurea medusa. B: S. nishiokae. C: S.

spicata. spicata. D: S. tridac りJla. E: S. kingii. F: S. delavayi. G: S. globosa. H: S. hookeri. B 征= 10flm. Arrows in- dicate dicate B-chromosome. October October 2004 Journal of Japanese Botany Vo l. 79 No. 5 277

K

韓

Fig. Fig. 2. Continued. 1: S. obvallata. J: S. forrestii. K: S. graminea. L: S. graminifolia. M: S. hieracioides. N: S. leontodontoides. leontodontoides. 0: S. yakla. Bar = 10μffi. 278 278 植物研究雑誌第79 巻第5号 平成16 年 10 月 previous previous paper (Amano and Ohba 2000). It is chromosome numbers within the genus distributed distributed in the Himalayas ，Tibet ， Sichuan Saussurea ， the somatic count 2n = 64 is re- and and Qinghai. garded as a tetraploid ， with x = 16. This is 9. 9. Saussurea obvallata (DC.) Edgew. the first record of a tetraploid with x = 16 The somatic chromosome number in this within the genus. The species is distributed species species was 2n = 32 ， with or without three B- in open subalpine meadows in the eastern chromosomes (Fig. 11). Chromosome counts Himalayas and S. W. China. It belongs to from from two populations agree with previous re- section Saussurea. ports ports (Shetty 1967 ， Gupta and Gill 1989 ， 14. Saussurea leontodontoides (DC.) Sch. Gupta Gupta et al. 1989 ，Amano and Ohba 2000) Bip. and and confirm the chromosome number of The somatic chromosome number was 2n = 32 for this species. Saussurea obvallata found to be 2n = 30 (Fig. lN) ， and is the first is is widely distributed in the Himalayas ， Tibet record for this species. The species belongs and and China (Qinghai ，Gansu ， Sichuan and to section Cyathidium ， and is distributed in Yunnan). Yunnan). the Himalayas ， Tibet and S. W. China. 15. 15. Saussurea yakla C. B. Clarke Subgenus Subgenus Saussurea The somatic chromosome number of s担任 10. 10. Saussurea forrestii Diels ples taken from two populations were 2n = The somatic chromosome number in this 34 (Fig. 10). The chromosome numbers species species was 2n = 34 with one B-chromosome were examined for the first time. This spe- (Fig. (Fig. 1J). The chromosome number for this cies is characterized by having a uni-seriate species species is reported for the first time. This plumose pappus and a conspicuous ，muricate species species belongs to section J acea and is dis- achene surface. It belongs to section tributed tributed in the Himalayas ， Tibet and Yunnan. Cyathidium. Saussurea yakla ， as circum- 11. 11. Saussurea graminea Dunn scribed by Springate (2001) ， is distributed in The somatic chromosome number was the Himalaya and Tibe t. 2n = 32 with three B-chromosomes (Fig. lK). lK). The chromosomes of this species were Among the 15 species in the genus examined examined for the first time. This species be- Saussurea collected from the Himalayas and longs longs to section Pycnocephala and is distrib- adjacent regions examined in the present uted uted in Tibet and S.W. China (Sichuan ， study ， the somatic chromosome number of Gansu ， Yunnan). 2n = 30 was found in one species ， 2n = 32 in 12. 12. Saussurea graminifolia Wall. ex DC. five species ， 2n = 34 in five species ，2n = 36 The somatic chromosome number in this in three species and 2n = 64 in one species. species species was 2n = 32 (Fig. lL). The chromo- The chromosome numbers for Saussurea some counts from three populations support spic αta ， S. kingii ，S. delavayi ，S. globosa ， S. a previous report for this species (Amano forrestii ， S. gramine α， S. hieracioides ， S. and and Ohba 2000). Saussurea graminifolia is leontodontoides and S. yakl αwere first similar similar to S. graminea ， but differs in having counted in the present study. Based on these basalleaves basalleaves with a broad sheath. The distri- results as well as those given elsewhere (see bution bution of this species is wide ， ranging Fujikawa and Ohba 2003b) ， S. hieracioides ， throughout throughout the Himalayas and Tibe t. with 2n = 64 ，is considered to be te 住aploid 13. 13. Saussurea hieracioides Hook. f. with x = 16; the other species with 2n = 30 ， The somatic chromosome number was 32 ，34 ， and 36 紅 e diploids with x = 15 ，16 ， 2n = 64 (Fig. 1M) ， and was examined the for 17 ， and 18 ，respectively. first first time. Considering previous reports of Chromosome numbers representing 15 October October 2004 Joumal of Japanese Botany Vo l. 79 No. 5 279

species species of the genus Saussurea in the present Nepal ， in quest of Saussurea. Newslet t. Him. Bo t. study study suggested a diploid state except in one (26): 6-7. species. species. It seems that the Saussurea species ー- 2002. A taxonomic study of Saussurea goss 伊i- phora and allied species (Asteraceae) in the Nepal in in the Himalayas and a司jacent regions have Himalayas. Himalayas. In: Noshiro S. and R 司bhand 訂 i K. R. diversified diversified at a diploid leve l. (eds.) ， Himalayan Botany in the Twentieth and

Twenty 同 first Centuries ， pp. 112-116. The Society We 紅 e grateful to Professor Michio of Himalayan Botany ， Tokyo. Wakabayashi ，Makino Herbarium ，Tokyo 一一一 and Ohba H. 2003a. A cytological study of Metropolitan Metropolitan University and D r. Makoto Saussurea subgenus Eriocoryne (Asteraceae) from the the Nepal Himalaya. 1. Jpn. Bo t. 78: 135-144. Amano ， N atural History Museum and 一一一 and 一一一 2003b. A list of chromosome numbers Institute ， Chiba for their technical advice in of Saussurea (Asteraceae). Newslet t. Him. Bo t. the the cytological analyses and field sampling. (32): 9-14. Gupta Gupta R. C. and Gill B. S. 1989. Cytopalynology of Thanks 紅 e also due to Dr. Shinobu Akiyama ， N ational Science Museum ， north and central Indian Compositae. J. Cyto l. Gene t. 24: 96-105. Tsukuba; Tsukuba; Dr. Futoshi 民1iyamoto ，Tokyo 一一，一一一 and Garg R. K. 1989. Chromosomal con- University University of Agriculture for providing us spec 回 s of western Himalayan Compositae. Aspects with with materials; and to Mr. Madhu Sudan Pl. Sci. 11: 427 -4 37. Bista ， the ex-Director General ， Department Haffner E. 2000. On the phylogeny of the subtribe of of Plant Resources ， the Ministry of Forest Carduinae (tribe Cardueae ， Compositae). Englera and and Soil Conservation ，HMG Nepal ， for his 21. 209 pp. Botanic Garden and Botanical Museum Berlin-Dahlem ，Berlin. help help in ca 町 ying out the field surveys. We Hajra P. K. 1995. Saussurea. In: Hajra P. K.， Rao R. thank thank Mr. Stephan Gale ， the Kochi R.， Shigh D. K. and Uniyal B. P. (eds.) ，Fl ora of Prefectural Prefectural Makino Botanical Garden ， for India 12: 186-216. Botanical Survey of India ， checking checking the English manuscript; and also Calcutta. the the reviewers for their constructive and ef- Hooker J. D. 188 1. Saussurea. The flora of British India India 3: 365-377. L. Reeve & Co. ， London. fective fective criticism. Huang R. F.， Shen S. D. and Liu X. F. 1996. Studies This This study was partly supported by a on the chromosome number and polyploidy for a Grant-in-Aid Grant-in-Aid for Scientific Research (A) to number of plants in the north-east Qinghai-Xizang H. H. O. from the Japan Society for the Plateau. Acta Bo t. Borea l. -Occiden t. Sin. 16 (3): Promotion Promotion of Science. 310-318 (in Chinese with English abstract). Ikeda Ikeda H. and Miyamoto F. 1996. A botanical research in in the Hengduan Mountains ，SW China ， in 1996. References References Newslet t. Him. Bo t. (20): 1-5. Akiyama S. and Ikeda H. 2002. Sino-Japanese research 一一- and 一一一 1998. 1997 botanical research in the in in northwest Yunnan and Tibet ，200 1. Newslet t. Hengduan Mountains ，SW China. Newslett. Him. Him. Bo t. (29): 8-13. Bo t. (23): 1-5. Amano M. and Ohba H. 2000. Chromosome numbers Jee R.， Dhar U. and Kachroo P. 1983. IOPB chromo- of of some alpine species of Saussurea (Asteraceae) some number reports LXXI X. Taxon 32 (2): 32 1. in in Nepal Himalaya. 1. Jpn. Bo t. 75: 178-184. Kita Y.， Fujikawa K 叶It o M. ，Ohba H. and Kato M. In Bremer K. 1994. Saussurea. In: Asteraceae Cladistics press. Molecular phylogenetic analyses and & Classification ， pp. 147-148. Timber Press ， systematics of the genus Saussurea and related Portland. Portland. genera (Asteraceae ，Cardueae). Chater Chater A. O. and tamura Ki S. 1982. Saussurea. In: Kitamura S. 1969. Compositae of southeastern Asia Hara H. and Williams L. W. J. (eds よAn Enumera- and Himalayas IV. Acta Phytotax. Geobo t. 24: 1- tion tion of the Fl owering Plants of Nepal 3: 37 -41. 27. British British Museum (Na t. His t.)， London. Kondo K.， Taniguchi K.， Tanaka R. and Gu Z. J. 1990. Clarke Clarke C. B. 1876. Saussurea. In: Compositae Indicae ， Karyomorphological studies in Chinese plant- pp. pp. 220-236. Thacker ， Spink and Co. ，Calcutta. species involving the J apanese floristic elements ，1. Fujikawa Fujikawa K. 2000. A short trip to Yak Kh arka ，West La Kr omosomo 11-57: 1889-1915. 280 280 植物研究雑誌第79 巻第5号 平成16 年 10 月

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藤川和美aj ，池田 博b ，村田勝哉b ，小林隆晃b ，

中野辰晃b ，大場秀章a，武 素功C' ヒマラヤおよ び周辺地域に産する卜ウヒレン属15 種(キク科) の染色体数 ヒマラヤおよびその周辺地域に産するトウヒレ 36 の種はそれぞれ x= 15 ， 16 ， 17 ， 18 を基本数と ン属植物 15 種の染色体数を報告した.このうち 9 する 2t 音イ本であり S. hieracioides の2n=64 は x= 種については今回が初めての報告である.染色体 16 の 4 倍体であると考えられる. 数は Saussurea leontodontoides が 2n = 30 ， S. globo- また，これまでヒマラヤ地域のみに分布すると sa ， S. graminea ， S. graminifolia ， S. obvallata とS. されていた S. nishiokae がチベットで採取された. spica 仰が2n = 32 ， S. delavayi ，S. forrestii ， S. kingii ， これは本種の，中国における初めての報告である. S. S. medusa と S. yakla カf 2n = 34 ， S. nishiokae ， S. (a 東京大学総合研究博物館， hookeri と S. tridac ηlα カf 2n = 36 ， S. hieracioides *現所属:高知県立牧野植物園，

が 2n=64 であった.これまで報告されているト b 岡山理科大学，

ウヒレン属植物の染色体数から， 2n = 30 ，32 ，34 ， c中国科学院見明植物研究所)