J. Jpn. Bot. 85: 157–165 (2010)

Cytological Studies on in the Nepal Himalaya I. Chromosome Counts of 14 Species Collected from the Manaslu Himalaya, Central Nepal

a a, b Okihito Yano , Hiroshi Ikeda *, Mark F. Watson , c a Keshab R. Rajbhandari and Hideaki Ohba

aDepartment of Botany, University Museum, University of Tokyo, 7-3-1, Hongo, Bunkyo-ku, Tokyo, 113-0033 JAPAN; bRoyal Botanic Garden Edinburgh, 20a, Inverleith Row, Edinburgh EH3 5LR, Scotland, U. K.; cG. P. O. Box 9446, Kathmandu, NEPAL *Corresponding author e-mail: [email protected]

(Received on January 7, 2010)

Chromosome numbers of 14 species of Cyperaceae recently collected from the Manaslu Himalaya in Central Nepal are presented. Chromosome numbers for Carex inanis Kunth (2n = 42), C. rufulistolon T. Koyama (2n = 46), and C. speciosa Kunth (2n = 44) are reported here for the first time. Our findings of 2n = 40 for compressus (L.) Panz. ex Link, 2n = 46 for C. filicina Nees, and 2n = 32 for Isolepis setacea (L.) R. Br. differ from previously published reports. Chromosome numbers of C. atrofusca Schkuhr subsp. minor (Boott) T. Koyama (2n = 48), Cyperus haspan L. (2n = 26), Eleocharis congesta D. Don (2n = 20), Fimbristylis dichotoma (L.) Vahl (2n = 20), F. littoralis Gaudich (2n = 10), F. schoenoides (Retz.) Vahl (2n = 10), Lipocarpha chinensis (Osbeck) Kern (2n = 26), and Schoenoplectus juncoides (Roxb.) Palla (2n = 74) coincide with past reports.

Key words: Aneuploidy, chromosome number, Cyperaceae, Himalaya, Nepal.

The Nepal Himalaya is known for high genera. In particular, chromosome numbers of levels of diversification of many vascular Cyperaceae from the Himalaya and adjacent (Ohba 1988). With over 5000 species regions have been reported by several authors, the Cyperaceae is one of the largest monocot such as Mehra and Sachdeva (1971, 1975a, families (Goetghebeur 1998), with about 180 1975b, 1976, 1979), Dietrich (1972), Sachdeva taxa in 19 genera recorded from Nepal (Koyama (1977), Bir et al. (1986, 1988, 1991, 1992), 1978). Cytological analyses have proven useful Cheema et al. (1992, 1993), and Hoshino et for understanding the diversification of several al. (2000). Most of these cytological works taxa in the Nepal Himalaya (Wakabayashi and used material collected from India, except Ohba 1988, Akiyama et al. 1992, Ikeda and Dietrich (1972) and Hoshino et al. (2000) from Ohba 1999), and the present study contributes Nepal. For understanding the evolution and to assessing its value with respect to Nepalese diversification of Cyperaceae in the Nepal Cyperaceae. Himalaya, more cytological studies are necessary Roalson (2008) summarized the many past from the region. This is the first of a series of cytological studies on the Cyperaceae including papers reporting cytological characteristics of chromosome counts for 851 species of 31 Cyperaceae in the Nepal Himalaya and adjacent

—157— 158 植物研究雑誌 第 85 巻 第 3 号 2010 年 6 月

Table 1. Species, localities, voucher specimens and chromosome numbers of fourteen species of Cyperaceae collected from the Manaslu Himalaya and adjust regions in Central Nepal Chromosome Species, locality and voucher specimen Previous report (2n) number (2n) Blysmus compressus (L.) Panz. ex Link Lho – Samagaon, 3210 m (Ikeda & al. 20814068) 40 44 (Håkansson 1928, Tischler 1934, Rohweder 1937, Mehra and Sachdeva 1975b, Pogan and Izmailow 1983, Stoeva 1992a) 79 (Stoeva 1992b) Carex atrofusca Schkuhr subsp. minor (Boott) T. Koyama Samagaon, on way to Manaslu Base Camp, 4130 m 48 48 (Dietrich 1972) (Ikeda & al. 20814087) C. filicina Nees Bimthang – Gho, 2990 m (Ikeda & al. 20814112) 46 42 (Mehra and Sachdeva 1971, 1975b) 44 (Nijalingappa and Leela 1990) 48 (Sachdeva 1977, Mehra and Sachdeva 1979) C. inanis Kunth Ghap – Namrung, 2130 m (Ikeda & al. 20814044) 42 Ghap – Namrung, 2390 m (Ikeda & al. 20814050) 42 C. rufulistolon T. Koyama Samagaon, on way to Manaslu Base Camp, 4220 m 46 (Ikeda & al. 20814089) C. speciosa Kunth Ghap – Namrung, 2330 m (Ikeda & al. 20814049) 44 Gho – Tal, 2490 m (Ikeda & al. 20814115) 44 Cyperus haspan L. Khanchok – Arughat, 960 m (Ikeda & al. 20814011) 26 16 (Sharma and Sarkar 1967–1968, Sanyal 1972) 26 (Nijalingappa & al. 1978, Rath and Patnaik 1978, Tejavathi and Nijalingappa 1990, Matoba and Uchiyama 2009) 30 (Baquar 1978) 32 (Tanaka 1948) 36 (Mehra and Sachdeva 1971, 1975b) Eleocharis congesta D. Don Syange – Bhulbhule, 1130 m (Ikeda & al. 20814120) 20 20 (Tanaka 1948, Nijalingappa 1972, Yano et al. 2004) Fimbristylis dichotoma (L.) Vahl Gorkha town, 1270 m (Ikeda & al. 20814002) 20 20 (Mehra and Sachdeva 1971 ) F. littoralis Gaudich. Khanchok – Arughat, 960 m (Ikeda & al. 20814009) 10 10 (Nijalingappa 1975) F. schoenoides (Retz.) Vahl Khanchok – Arughat, 960 m (Ikeda & al. 20814013) 10 10 (Nijalingappa 1972, 1975, Sanyal and Sharma 1972, Rath and Patnaik 1974, Mehra and Sachdeva 1975a, 1976) Isolepis setacea (L.) R. Br. Ghap – Namrung, 2240 m (Ikeda & al. 20814046) 32 26 (Davies 1956) 28 (Mičieta 1986, Hoshino et al. 2000) Lipocarpha chinensis (Osbeck) Kern Khanchok – Arughat, 960 m (Ikeda & al. 20814010) 26 26 (Sanyal and Sharma 1972, Rath and Patnaik 1974, Tejavathi and Nijalingappa 1990) Schoenoplectus juncoides (Roxb.) Palla Khanchok – Arughat, 960 m (Ikeda & al. 20814014) 74 74 (Iwasaki and Ueki 1979, Yano and Hoshino 2005) Syange – Bhulbhule, 1130 m (Ikeda & al. 20814121) 74 June 2010 Journal of Japanese Botany Vol. 85 No.3 159 regions. This part focuses on the chromosome Kashmir Himalaya. In this study, a new number, numbers of Cyperaceae collected around 2n = 40, was observed. Somatic metaphase Manaslu Himalaya, Central Nepal. chromosomes were less than 1.2 µm in length.

Materials and Methods 2. Carex atrofusca Schkuhr subsp. minor Karyomorphological observations were (Boott) T. Koyama (2n = 48, Fig. 1B) conducted on 14 species of eight genera in Carex atrofusca subsp. minor is distributed Cyperaceae mostly collected from Manaslu in Pamir, the Himalayas (Nepal to Bhutan), Himalaya and adjacent areas, Central Nepal and Eastern Tibet (Koyama 1975, 1978). (Table 1). Somatic chromosomes were Chromosome number of the species from the observed in the meristematic cells of root tips. Manaslu Himalaya was 2n = 48, confirming Root tips were harvested and prepared from the previous report from the Mahalangur Himal material whilst in the field, pretreated in 2 mM in eastern Nepal (Dietrich 1972). Somatic 8-hydroxyquinoline solution for 3–6 hours at metaphase chromosomes were less than 1.3 µm 10–20°C, and then fixed with Newcomer’s fluid in length. (see Sharma and Sharma 1980). In the laboratory the fixed roots were hydrated with an alcohol 3. Carex filicina Nees (2n = 46, Fig. 1C) series (70%, 30%, 15%) and distilled water, then Carex filicina is distributed in the Himalaya, hydrolyzed with 1N HCl for 10 minutes at 60°C, Eastern India, East China, Taiwan, and south to and stained with leuco-basic fuchsin for 1 hour Malaysia (Koyama 1978). In this study, a new at 23°C. The samples were then macerated in a number, 2n = 46, was observed from the material mixture of 2% pectinase and 2% cellulase for 1 from the Manaslu Himalaya. Somatic metaphase hour at 37°C, restained with 1% aceto-orcein, chromosomes were less than 1.3 µm in length. and squashed. Voucher herbarium specimens are Several chromosome numbers of C. filicina were kept in the Herbarium of the University of Tokyo reported in the previous reports. Chromosome (TI), with duplicates available in the National numbers of 2n = 42 were reported by Mehra and Herbarium in Nepal (KATH) and Royal Botanic Sachdeva (1971, 1975b) from Nainital in the Garden, Edinburgh (E). West Himalaya, 2n = 48 by Sachdeva (1977) and Mehra and Sachdeva (1979) from Darjeeling in Results and Discussion the East Himalaya, and 2n = 44 by Nijalingappa Chromosome numbers determined in this and Leela (1990) from Tamilnadu and Karnataka study are shown in Table 1. Primary constriction in South India. It is thought that C. filicina has a was not observed in any species. series of aneuploids in the species.

1. Blysmus compressus (L.) Panz. ex Link (2n 4. Carex inanis Kunth (2n = 42, Fig. 1D) = 40, Fig. 1A) Carex inanis is distributed from the Blysmus compressus is widely distributed highlands of Central Asia, and Kashmir to the in Europe and Asia (Iran, Pakistan, Nepal, North Sikkim Himalayas (Koyama 1978). We have India, and Southwest China) (Koyama 1978). not found any earlier chromosome number for The chromosome numbers of B. compressus C. inanis and so our count of 2n = 42 is the were reported as 2n = 44 (Håkansson 1928, first to be determined for this species. Somatic Tischler 1934, Rohweder 1937, Pogan and metaphase chromosomes were less than 1.3 µm Izmailow 1983, Stoeva 1992a) and 2n = 79 in length. (Stoeva 1992b) from Europe. Mehra and Sachdeva (1975b) observed 2n = 44 from the 160 植物研究雑誌 第 85 巻 第 3 号 2010 年 6 月

Fig. 1. Photomicrographs of somatic metaphase chromosomes of fourteen species of Cyperaceae from the Manaslu Himalaya. A. Blysmus compressus (2n = 40). B. Carex atrofusca subsp. minor (2n = 48). C. C. filicina (2n = 46). D. C. inanis (2n = 42). E. C. rufulistolon (2n = 46). F. C. speciosa (2n = 44). G. Cyperus haspan (2n = 26). H. Eleocharis congesta (2n = 20). I. Fimbristylis dichotoma (2n = 20). J. F. littoralis (2n = 10). K. F. schoenoides (2n = 10). L. Isolepis setacea (2n = 32). M. Lipocarpha chinensis (2n = 26). N. Schoenoplectus juncoides (2n = 74). Bar = 5 µm.

5. Carex rufulistolon T. Koyama (2n = 46, Fig. 1E) rufulistolon was found to be 2n = 46, the first Carex rufulistolon is endemic to the alpine record for this species. Somatic metaphase regions in Nepal (Koyama 1978, Katsuyama chromosomes were less than 1.1 µm in length. 2008). The chromosome number of C. June 2010 Journal of Japanese Botany Vol. 85 No.3 161

6. Carex speciosa Kunth (2n = 44, Fig. 1F) 9. Fimbristylis dichotoma (L.) Vahl (2n = 20, Carex speciosa is distributed from India, Fig. 1I) Nepal, China to Southeast Asia (Liang et al. Fimbristylis dichotoma is widely distributed 2000). Carex speciosa had the chromosome from the tropical to the temperate zones number of 2n = 44, the first chromosome (Koyama 1978). Fimbristylis dichotoma from count be determined for this species. Somatic the Manaslu Himalaya had the chromosome metaphase chromosomes were less than 1.3 µm number of 2n = 20, confirming a previous report in length. from Nainital in the West Himalaya (Mehra and Sachdeva 1971). Somatic metaphase 7. Cyperus haspan L. (2n = 26, Fig. 1G) chromosomes ranged between 1.4 and 2.0 µm Cyperus haspan is widely distributed in in length. Intraspecific polyploidy (2n = 10 and the tropical, subtropical and warm-temperate 20) has been reported previously in this species regions of both hemispheres (Koyama 1978). In from India by Mehra and Sachdeva (1975b, this study, we record a chromosome number of 1979) and Bir et al. (1986, 1988, 1992). Our 2n = 26 for this species, and somatic metaphase chromosome number of 2n = 20 is considered to chromosomes of less than 1.3 µm in length. be tetraploid with the basic chromosome number The chromosome numbers of C. haspan have x = 5. Although many intraspecific taxa have been variously reported as 2n = 16 (Sharma been described in F. dichotoma (Ohwi 1944, and Sarkar 1967–1968, Sanyal 1972), 2n = 26 Koyama 1961, 1974, Kern 1974), Yano and (Nijalingappa et al. 1978, Rath and Patnaik Hoshino (2006a) demonstrated the polyphyly of 1978, Tejavathi and Nijalingappa 1990, Matoba F. dichotoma using DNA sequence data. Further and Uchiyama 2009), 2n = 30 (Baquar 1978), studies, comparing morphological variation 2n = 32 (Tanaka 1948), and 2n = 36 (Mehra and cytological variation with DNA sequence and Sachdeva 1971, 1975b). It is thought that data, will be needed to clarify the relationships C. haspan has series of aneuploids, and the between and intraspecific polyploidy chromosome count from the Manaslu Himalaya for this widespread and variable species. coincides with several specimens from India (Nijalingappa et al. 1978, Rath and Patnaik 10. Fimbristylis littoralis Gaudich. (2n = 10, 1978, Tejavathi and Nijalingappa 1990). Fig. 1J) Fimbristylis littoralis is widely distributed 8. Eleocharis congesta D. Don (2n = 20, Fig. in pantropical regions (Koyama et al. 2000). 1H) We recorded a chromosome number of 2n Eleocharis congesta is distributed in India, = 10 for this species and somatic metaphase Nepal, Indo-China, China, and Japan (Koyama chromosomes ranged between 1.4 and 1.9 µm in 1978). Eleocharis congesta from the Manaslu length. This chromosome numbers agrees with a Himalaya had the chromosome number of previous record from India (Nijalingappa 1975). 2n = 20, confirming previous reports from India (Nijalingappa 1972) and Japan (Tanaka 11. Fimbristylis schoenoides (Retz.) Vahl (2n = 1948, Yano et al. 2004). Somatic metaphase 10, Fig. 1K) chromosomes ranged between 1.9 and 2.2 µm Fimbristylis schoenoides is distributed in length. Although Koyama (1961) noted in Nepal, India, Indo-China, Southern China, that Japanese E. congesta had longer perianth Malaysia, and North Australia (Koyama 1978). bristles than those from the Himalaya, there Fimbristylis schoenoides had the chromosome was no karyomorphological difference between number of 2n = 10, and somatic metaphase Japanese and the Himalayan samples. chromosomes ranged from 1.4 to 1.9 µm 162 植物研究雑誌 第 85 巻 第 3 号 2010 年 6 月 in length. Our chromosome count agrees In this study, chromosome numbers of with previous reports from Indian material Carex inanis (2n = 42), C. rufulistolon (2n = 46), (Nijalingappa 1972, 1975, Sanyal and Sharma and C. speciosa (2n = 44) were determined for 1972, Rath and Patnaik 1974, Mehra and the first time. Carex inanis and C. rufulistolon Sachdeva 1975a, 1976). belong to section Trachychlaenae (Koyama 1954, Liang et al. 2000). Heilborn (1924) 12. Isolepis setacea (L.) R. Br. (2n = 32, Fig. 1L) pointed out that closely related species in Carex Isolepis setacea is widely distributed shared similar chromosome numbers. Yano et in the Himalayas, Asia, Europe, and Africa al. (2008) also reported that Japanese Carex (Katsuyama 2008). The chromosome number of within the same section possessed similar I. setacea was reported to be 2n = 26 by Davies chromosome numbers. Hipp (2007) suggested (1956), and 2n = 28 by Mičieta (1986) and that chromosome numbers in Carex section Hoshino et al. (2000). Among them, Hoshino Ovales indicated an evolutionary clade-specific et al. (2000) reported 2n = 28 (n = 14II) from equilibrium, based on DNA sequence data. the Langtang Himal in Central Nepal. In our In the present study our cytological results, in study a new number, 2n = 32, was counted from particular the similarity of chromosome number, the Manaslu Himalaya. It is thought that this support a close relationship between C. inanis species has intraspecific aneuploids. The somatic and C. rufulistolon. metaphase chromosomes were less than 1.1 µm The chromosome numbers of Blysmus in length. compressus, Carex filicina, and Isolepis setacea differ from those of previous reports. 13. Lipocarpha chinensis (Osbeck) Kern (2n = These three species are widely distributed and 26, Fig. 1M) they may all have intraspecific aneuploids. Lipocarpha chinensis is distributed in the Cytogeographically, two types of intraspecific tropical and subtropical regions of Africa, Asia, aneuploidy have been reported in Carex and south to Malaysia, and North Australia (Koyama Eleocharis: one is populational variation 1978). Lipocarpha chinensis from the Manaslu (Hoshino et al. 1993, Hoshino and Okamura Himalaya was found to have a chromosome 1994, Hoshino and Onimatsu 1994, Yano and number of 2n = 26, confirming previous reports Hoshino 2006b), and another is geographic from India (Sanyal and Sharma 1972, Rath variation (Hoshino 1992, Hoshino and and Patnaik 1974, Tejavathi and Nijalingappa Waterway 1994, Ohkawa et al. 2000). Future 1990). Somatic metaphase chromosomes ranged studies including further sampling are needed to between 1.1 and 1.4 µm in length. understand the basis for intraspecific aneuploidy in these three species. 14. Schoenoplectus juncoides (Roxb.) Palla (2n = 74, Fig. 1N) The authors extend their appreciation to Schoenoplectus juncoides is distributed in Dr. L. R. Sharma, Director of the Department the Himalayas, India, Indo-China, China, Japan, of Resources, Ministry of Forests and and Malaysia (Koyama 1978). Schoenoplectus Soil Conservation, Government of Nepal, for juncoides from the Manaslu Himalaya had the his guidance and assistance in conducting field chromosome number of 2n = 74, confirming research in Nepal. We also thank the members previous reports from Japan (Iwasaki and of the field trip in Nepal in 2008 for their help Ueki 1979, Yano and Hoshino 2005). Somatic for collecting plant materials. This study was metaphase chromosomes were less than 0.9 µm partly supported by a Grant-in-Aid for Scientific in length. Research (A), No. 18255004 in 2008 (to H. O.) June 2010 Journal of Japanese Botany Vol. 85 No.3 163 from the Ministry of Education, Culture, Sports, Hoshino T. and Onimatsu A. 1994. Cytological studies of Science and Technology, Government of Japan. Carex duvaliana (Cyperaceae) with special references to meiotic configurations of intraspecific aneuploids. J. References Jpn. Bot. 69: 37–41. Hoshino T. and Waterway M. J. 1994. Cytogeography and Akiyama S, Wakabayashi M. and Ohba H. 1992. meiotic chromosome configurations of six intraspecific Chromosome evolution in Himalayan Impatiens aneuploids of Carex conica Boott (Cyperaceae) in (Balsaminaceae). Bot. J. Linn. Soc. 109: 247–257. Japan. J. Plant Res. 107: 131–138. Baquar S. R. 1978. Cytological studies of some southern Hoshino T., Aosaki K. and Onimatsu A. 1993. Cytological Nigerian Cyperaceae. La Kromosomo II 9: 263–270. studies of Carex stenostachys (Cyperaceae) with Bir S. S., Cheema P. and Sidhu M. K. 1991. Cytological special references to meiotic configurations of observations on Scirpus Linn. from North India. intraspecific aneuploids. La Kromosomo II 71–72: Cytologia 56: 645–651. 2451–2455. Bir S. S., Cheema P. and Sidhu M. K. 1992. Chromosomal Hoshino T., Rajbhandari K. R. and Ohba H. 2000. analysis of Fimbristylis Vahl in Punjab, North West Cytological studies of eleven species of Cyperaceae India. Proc. Indian Natl. Sci. Acad., B. Biol. Sci. 58: collected from Central Nepal. Cytologia 65: 219–224. 63–70. Ikeda H. and Ohba H. 1999. A systematic revision of Bir S. S., Sidhu M. K. and Kamra S. 1986. Karyotypic genus Potentilla L. section Leptostylae (Rosaceae) in studies on certain members of Cyperaceae from the Himalaya and adjacent regions. In: Ohba H. (ed.), Punjab, North West India. Cytologia 51: 95–106. The Himalayan Plants 3: 31–117. University of Tokyo Bir S. S., Kamra S., Sidhu M. K. and Cheema P. 1988. Press, Tokyo. Cytomorphological studies on some members of Iwasaki K. and Ueki K. 1979. Chromosome numbers of Cyperaceae from North India. J. Cytol. Genet. 23: three bulrushes (Scirpus taxa). J. Weed Sci. Technol. 14–37. 24: 240–242 (in Japanese with English abstract). Cheema P., Bir S. S. and Sidhu M. K. 1993. Chromosomal Katsuyama T. 2008. Cyperaceae. In: Ohba H., Iokawa Y. variabilities in Cyperus Linn. II. Section Pycreus and Sharma L. R. (eds.), Flora of Mustang, Nepal. pp. (Beauv.) Griseb. from NW India. Cytologia 58: 345– 431–471. Kodansha Scientific, Tokyo. 349. Kern J. H. 1974. Cyperaceae. In: van Steenis C. G. G. J. Cheema P., Kumari S., Sidhu M. and Bir S. S. 1992. (ed.), Flora Malesiana Ser. I, 7: 540–592. Noordhoff Karyomorphology of Fimbristylis from Punjab. J. International Publishers, Leiden. Cytol. Genet. 27: 163–173. Koyama T. 1954. New carices from Nepal collected by Davies E. W. 1956. Some new chromosome numbers in the Sasuke Nakao. Acta Phytotax. Geobot. 15: 111–114. Cyperaceae. Watsonia 3: 242–243. Koyama T. 1961. Classification of the familyCyperaceae I. Dietrich W. 1972. In IOPB chromosome number reports. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 8: 84–99. XXXVI. Taxon 21: 333–346. Koyama T. 1974. The genus Fimbristylis (Cyperaceae) in Goetghebeur P. 1998. Cyperaceae. In: Kubitzki K. Ceylon. Bot. Mag. (Tokyo) 87: 301–331. (ed.), The Families and Genera of Vascular Plants 4. Koyama T. 1975. Cyperaceae. In: Ohashi H. (ed.), Flora of Flowering Plants, : Alismatanae and Eastern Himalaya, 3rd report. pp. 122–129. University Commelinanae (except Gramineae). pp. 141–190. of Tokyo Press, Tokyo. Springer, Berlin. Koyama T. 1978. Cyperaceae. In: Hara H., Stearn W. Håkansson A. 1928. Die chromosomen einiger Scirpoideen. T. and Williams H. J. (eds.), An Enumeration of the Hereditas 10: 277–292. Flowering Plants of Nepal 1: 96–120. British Museum Heilborn O. 1924. Chromosome numbers and dimensions, (Natural History), London. species-formation and phylogeny in the genus Carex. Koyama T., Kuoh C.-S. and Leong W.-C. 2000. Hereditas 5: 129–216. Cyperaceae. In: Hsieh C.-F., Huang T.-C., Kuoh C.-S., Hipp A. L. 2007. Non-uniform processes of chromosome Ohashi H. and Su H.-J. (eds.), Flora of Taiwan 2nd. ed. evolution in sedges (Carex: Cyperaceae). Evolution 5: 194–237. National Taiwan University, Taipei. 61: 2175–2194. Liang S.-Y., Dai L.-K, Tang Y.-C. and Li P.-C. 2000. Hoshino T. 1992. Cytogeographical study of four Cyperaceae (2) Caricoideae. In: Dai L.-K. and Liang aneuploids of Carex oxyandra Kudo in Japan. Bot. S.-Y. (eds.), Flora Reipublicae Popularis Sinicae 12: Mag. (Tokyo) 105: 639–648. 1–582. Science Press, Beijing (in Chinese). Hoshino T. and Okamura K. 1994. Cytological studies on Matoba H. and Uchiyama H. 2009. Chromosome counts meiotic configurations of intraspecific aneuploids of of aquatic and wetland plants growing in acid sulfate Carex blepharicarpa (Cyperaceae) in Japan. J. Plant soil from the Central Plain, Thailand. J. Jpn. Bot. 84: Res. 107: 1–8. 313–320. 164 植物研究雑誌 第 85 巻 第 3 号 2010 年 6 月

Mehra P. N. and Sachdeva S. K. 1971. In: IOPB Sachdeva S. K. 1977. In IOPB chromosome number chromosome reports. XXXIII. Taxon 20: 609–614. reports. LVI. Taxon 26: 257–274. Mehra P. N. and Sachdeva S. K. 1975a. In: IOPB Sanyal B. 1972. Cytological studies in Indian Cyperaceae chromosome reports. XLIX. Taxon 24: 501–516. II. Tribe Cypereae. Cytologia 37: 33–42. Mehra P. N. and Sachdeva S. K. 1975b. Cytology of some Sanyal B. and Sharma A. 1972. Cytological studies in west Himalayan Cyperaceae. Cytologia 40: 497–515. Indian Cyperaceae. Tribe Scirpeae. Cytologia 37: Mehra P. N. and Sachdeva S. K. 1976. Cytology of some 13–32. sedges from N.W. India. Cytologia 41: 585–590. Sharma A. K. and Sarkar A. K. 1967–1968. Chromosome Mehra P. N. and Sachdeva S. K. 1979. Cytological number reports of plants in Annual Report, observations on some east Himalayan monocots. Cytogenetics Laboratory, Department of Botany, Cytologia 44: 233–240. University of Calcutta. The Research Bulletin 2: 38–48. Mičieta, K. 1986. Karyological study of the Slovak flora Sharma A. K. and Sharma A. 1980. Chromosome XI. Acta Fac. Rerum Nat. Univ. Comenianae, Botanica Techniques. Theory and Practice, 3rd. ed. p. 55. 33: 51–55. Butterworths, London. Nijalingappa B. H. M. 1972. In: IOPB chromosome Stoeva M. P. 1992a. Karyological study of the genera reports. XXXVIII. Taxon 21: 679–684. Blysmus Panzer, Eriophorum L. and Scirpus L. Nijalingappa B. H. M. 1975. Cytological studies in (Cyperaceae) in Bulgaria. Dokl. Bulg. Akad. Nauk 45: Fimbristylis (Cyperaceae). Cytologia 40: 177–183. 113–115. Nijalingappa B. H. M. and Leela B. D. 1990. Cytological Stoeva M. P. 1992b. Chromosome numbers of Bulgarian studies in some south Indian species of Carex. Cyperaceae. Fitologiya (Sofia)43 : 77–78. Cytologia 55: 373–379. Tanaka N. 1948. The Problem of Aneuploidy. Biological Nijalingappa B. H. M., Nagaraj N. and Tejavathi D. H. Contribution in Japan, 4. Hokuryukan, Tokyo (in 1978. In: IOPB chromosome number reports. LXII. Japanese). Taxon 27: 519–535. Tejavathi D. H. and Nijalingappa B. H. M. 1990. Ohba H. 1988. The alpine flora of the Nepal Himalayas: an Cytological studies in some members of Cyperaceae. introductory note. In: Malla S. B. and Ohba H. (eds.), Cytologia 55: 363–372. The Himalayan Plants 1: 19–29. University of Tokyo Tischler G. 1934. Die Bedeutung der Polyploidie für die Press, Tokyo. Verbreitung der Angiospermen, erläuteret an den Ohkawa T., Yokota M. and Hoshino T. 2000. Aneuploidal Arten Schleswig-Holsteins, mit Ausblicken auf andere population differentiation in Carex sociata Boott Florengebiete. Bot. Jahrb. 67: 1–36. (Cyperaceae) Ryukyu Islands, Japan. Bot. J. Linn. Soc. Wakabayashi M. and Ohba H. 1988. Cytotaxonomic study 132: 337–358. of the Himalayan Saxifraga. In: Ohba H. and Malla S. Ohwi J. 1944. Cyperaceae Japonicae, II. A synopsis of B. (eds.), The Himalayan Plants 1: 71–90. University the Rhynchosporoideae and Scirpoideae of Japan, of Tokyo Press, Tokyo. including the Kuriles, Saghalin, Korea, and Formosa. Yano O. and Hoshino T. 2005. Molecular phylogeny and Mem. Coll. Sci. Kyoto Imp. Univ. Ser. B, 18: 1–182. chromosomal evolution of Japanese Schoenoplectus Pogan E. and Izmailow R. 1983. Further studies in (Cyperaceae), based on ITS and ETS 1f sequences. chromosome numbers of Polish angiosperms. Part Acta Phytotax. Geobot. 56: 177–189. XVII. Acta Biol. Cracov., Ser. Bot. 25: 57–77. Yano O. and Hoshino T. 2006a. Phylogenetic relationships Rath S. P. and Patnaik S. N. 1974. Cytological studies in and chromosomal evolution of Japanese Fimbristylis Cyperaceae with special reference to its taxonomy. I. (Cyperaceae) using nrDNA ITS and ETS 1f sequence Cytologia 39: 341–352. data. Acta Phytotax. Geobot. 57: 205-217. Rath S. P. and Patnaik S. N. 1978. Cytologic studies in Yano O. and Hoshino T. 2006b. Cytological studies of Cyperaceae with special reference to its taxonomy II. aneuploids in Eleocharis kamtschatica (Cyperaceae). Cytologia 43: 643–653. Cytologia 71: 141–147. Roalson E. H. 2008. A synopsis of chromosome number Yano O., Katsuyama T. and Hoshino T. 2008. Cytological variation in the Cyperaceae. Bot. Rev. 74: 209–393. studies of Japanese Carex (Cyperaceae) I. J. Jpn. Rohweder H. 1937. Versuch zur Erfassung der Cyperol. (13): 9–21 (in Japanese with English abstract). mengenmässigen Bedeckung des Darss und Zingst mit Yano O., Katsuyama T., H. Tsubota and T. Hoshino. polyploiden Pflanzen. Ein Beitrag zur Bedeutung der 2004. Molecular phylogeny of Japanese Eleocharis Polyploidie bei der Eroberung neuer Lebensraume. (Cyperaceae) based on ITS sequence data, and Planta 27: 501–549. chromosomal evolution. J. Plant Res. 117: 409–419. June 2010 Journal of Japanese Botany Vol. 85 No.3 165

矢野興一 a, 池田 博 a, M. F. ワトソン b, K. R. ラジバンダ リ c, 大場秀章 a: ネパールヒマラヤ産カヤツリグサ科植物の 細胞学的研究 I. マナスルヒマラヤ ( 中央ネパール ) より採 (2n = 48), Cyperus haspan L. (2n = 26), Eleocharis 集した 14 種の染色体数 congesta D. Don (2n = 20), Fimbristylis dichotoma マナスルヒマラヤ ( 中央ネパール ) より採集したカヤツ (L.) Vahl (2n = 20), F. littoralis Gaudish (2n = 10), リグサ科植物 14 種について染色体数を報告した．Carex F. schoenoides (Retz.) Vahl (2n = 10), Lipocarpha inanis Kunth (2n = 42), C. rufulistolon T. Koyama (2n chinensis (Osbeck) Kern (2n = 26), Schoenoplectus = 46), C. speciosa Kunth (2n = 44) の 3 種については，今 juncoides (Roxb.) Palla (2n = 74) の 8 種については , こ 回が初めての報告である．また , Blysmus compressus (L.) れまでの報告と一致していた． Panz. ex Link (2n = 40), C. filicina Nees (2n = 46) , a ( 東京大学総合研究博物館 , Isolepis setacea (L.) R. Br. (2n = 32) の 3 種については , bエジンバラ王立植物園 , これまでの報告と異なった染色体数が算定された．Carex cネパール・カトマンズ 私書箱 9446 号 ) atrofusca Schkuhr subsp. minor (Boott) T. Koyama