J. Jpn. Bot. 90: 249–259 (2015)

Cytological Studies on in the Nepal Himalaya III. Chromosome Counts of 18 Species Collected from Api Himal and Adjacent Areas, Far Western Nepal

a, b c Okihito Yano *, Hiroshi Ikeda , Colin Alistair Pendry d and Keshab Raj Rajbhandari

aFaculty of Biosphere-Geosphere Science, Okayama University of Science, 1-1, Ridai-cho, Kita-ku, Okayama, 700-0005 JAPAN; bThe University Museum, The University of Tokyo, 7-3-1, Hongo, Bunkyo-ku, Tokyo, 113-0033 JAPAN; cRoyal Botanic Garden Edinburgh, 20a, Inverleith Row, Edinburgh EH3 5LR, Scotland, UK; dG.P.O. Box 9446, Kathmandu, NEPAL *Corresponding author: [email protected]

(Accepted on February 28, 2015)

Chromosome numbers are presented for 18 species from seven genera of Cyperaceae collected from Api Himal and adjacent areas in Far Western Nepal. The chromosome numbers for cruenta Nees (2n = 44), C. kumaonensis Kük. (2n = 50), C. obscura Nees (2n = 56), C. stracheyi Boott ex C. B. Clarke (2n = 44), Kobresia esbirajbhandarii Rajbh. & H. Ohba (2n = 62), and Schoenoplectiella fuscorubens (T. Koyama) Hayasaka (2n = 42) are reported for the first time. Our finding of 2n = 46 for compressus (L.) Panz. ex Link and 2n = 62 for Carex foliosa D. Don differ from previous reports. The chromosome numbers of Carex filicina Nees (2n = 46), C. inanis Kunth (2n = 42), C. longipes D. Don (2n = 46), C. microglochin Wahlenb. (2n = 50), C. myosurus Nees (2n = 44), C. rufulistolon T. Koyama (2n = 46), uniglumis (Link) Schult. (2n = 42), Fimbristylis complanata (Retz.) Link (2n = 10), F. falcata (Vahl) Kunth (2n = 22), and setacea (L.) R. Br. (2n = 28 = 14II) agree with earlier studies.

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

Cytological studies have proven to be useful additional cytological studies are needed. for understanding the diversification of various In 2012, a Japan-UK-Nepal botanical Himalayan taxa (Wakabayashi and Ohba collecting team visited Api Himal and adjacent 1988, Akiyama et al. 1992, Ikeda and Ohba areas in Darchula District, Far Western Nepal, 1999). Chromosome numbers of Cyperaceae in and collected materials of Cyperaceae for the Nepal Himalaya have been also reported by cytological examination. This is the third report Dietrich (1972), Hoshino et al. (2000) and Yano of a series of papers on cytological characteristics et al. (2010a, 2010b, 2011). They reported the of Cyperaceae in the Nepal Himalaya, with the chromosome numbers for 28 taxa in nine genera. previous studies having reported chromosome However, in order to understand the evolution numbers in material collected from Manaslu and diversification of Cyperaceae in Nepal, Himal (Yano et al. 2010b) and Bajhang District

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Table 1. Species, localities, voucher specimens and chromosome numbers of 18 species of Cyperaceae collected from Far Western Nepal Chromosome Species, locality and voucher specimen Previous report (2n) number (2n) (L.) Panz. ex Link Dhaulo Odar to Joge Tal, 3399 m (Ikeda & al. 1215040) 46 40 (Yano et al. 2010b) 44 (Håkansson 1928, Tischler 1934, Rohweder 1937, Mehra and Sachdeva 1975b, Pogan and Izmailow 1983, Stoeva 1992a) 79 (Stoeva 1992b) Carex cruenta Nees Joge Tal, 4030 m (Ikeda & al. 1215052) 44 C. filicina Nees Lithi to Khayekot, 2200 m (Ikeda & al. 1215022) 46 42 (Mehra and Sachdeva 1971, 1975b) 44 (Nijalingappa and Leela 1990) 46 (Yano et al. 2010b) 48 (Sachdeva 1977, Mehra and Sachdeva 1979) C. foliosa D. Don Lithi to Khayekot, 2200 m (Ikeda & al. 1215019) 62 58 (Mehra and Sachdeva 1971, 1975b) C. inanis Kunth Khayekot to Simar Kharka, 2060 m (Ikeda & al. 1215024) 42 42 (Yano et al. 2010b) C. kumaonensis Kük. Simar Kharka to Khayekot, 2480 m (Ikeda & al. 1215062) 50 C. longipes D. Don Khayekot to Simar Kharka, 2060 m (Ikeda & al. 1215026) 46 42 (Sachdeva 1977, Mehra and Sachdeva 1979) 44 (Nijalingappa and Leela 1990, Hoshino et al. 2000) 46 (Yano et al. 2010a) C. microglochin Wahlenb. Simar Kharka to Dhaulo Odar, 3399 m (Ikeda & al. 1215038) 50 48 (Dietrich 1972, Yurtsev and Zhukova 1982) Joge Tal, 4100 m (Ikeda & al. 1215050) 50 50 (Löve and Löve 1981) 58 (Löve and Löve 1956, Moore and Chater 1971, Moore 1981) C. myosurus Nees Khayekot to Lithi, 2200 m (Ikeda & al. 1215064) 44 44 (Hoshino et al. 2000) C. obscura Nees Simar Kharka to Dhaulo Odar, 3150 m (Ikeda & al. 1215037) 56 C. rufulistolon T. Koyama Simar Kharka to Dhaulo Odar, 3010 m (Ikeda & al. 1215033) 46 46 (Yano et al. 2010b) C. stracheyi Boott ex C. B. Clarke Simar Kharka to Dhaulo Odar, 2930 m (Ikeda & al. 1215029) 44 Eleocharis uniglumis (Link) Schult. Joge Tal to Simar Kharka, 3399 m (Ikeda & al. 1215059) 42 42–50 (Bureš 1998) Fimbristylis complanata (Retz.) Link Lithi to Khayekot, 2200 m (Ikeda & al. 1215021) 10 10 (Kral 1971, Nijalingappa 1972, 1975, Rath and Patnak 1974) 16 (Tanaka 1939, Yano and Hoshino 2006) 20 (Rath and Patnaik 1978, Mehra and Sachdeva 1979) F. falcata (Vahl) Kunth Bitale to Pari Bagar, 920 m (Ikeda & al. 1215011) 22 22 (Sharma and Sarkar 1967–1968, Sanyal and Sharma 1972, Mehra and Sachdeva 1975a, 1976, Bir et al. 1986, 1988, 1992, Paramjeet et al. 1992) 44 (Nijalingappa 1975, 1977) (L.) R. Br. Joge Tal to Simar Kharka, 3399 m (Ikeda & al. 1215060) 28 = 14II 26 (Davies 1956) 28 (Mičieta 1986, Hoshino et al. 2000) 32 (Yano et al. 2010b) Kobresia esbirajbhandarii Rajbh. & H. Ohba Joge Tal, 4100 m (Ikeda & al. 1215049) 62 Schoenoplectiella fuscorubens (T. Koyama) Hayasaka Okhal to Bitale, 1100 m (Ikeda & al. 1215066) 42 August 2015 The Journal of Japanese Botany Vol. 90 No. 4 251

(Yano et al. 2010a). consists of a series of aneuploids (Yano et al. 2010b), a fairly common phenomenon in Materials and Methods Cyperaceae species which possess diffuse Chromosome counts were made for 18 centromeric chromosomes. species of seven genera in Cyperaceae (Table 1). The methods for somatic chromosome 2. Carex cruenta Nees (2n = 44, Fig. 1B) observation followed Yano et al. (2010b), Carex cruenta is distributed in the Himalaya with meiotic chromosomes examined in (Kashmir to Sikkim) and Tibet (Koyama 1978). pollen mother cells. Anthers were fixed with Carex cruenta had the chromosome number Newcomer’s fluid (see Sharma and Sharma of 2n = 44, and this is the first chromosome 1980) in the field. In the laboratory the fixed count to be determined for this species. Somatic anthers were stained with 1% aceto-orcein and metaphase chromosomes were less than 1.4 µm gently squashed before being examined under in length. This species has often been included the microscope. Voucher specimens are kept within the C. atrofusca complex (Noltie 1994), in the Herbarium of the University of Tokyo and the chromosome number for C. atrofusca s.l. (TI), with duplicates available in the National has variously been reported as 2n = 36, 38, 40, Herbarium of Nepal (KATH), the Herbarium of ca. 42, 48 or 56 (Heilborn 1924, Jørgensen et al. the Royal Botanic Garden Edinburgh (E) and 1958, Löve and Löve 1965, Dietrich 1967, 1972, the Herbarium of the Okayama University of Hedberg 1967, Hoshino et al. 1993, Yano et al. Science (OKAY). 2010b). These results indicate that this complex consists of a series of aneuploids. However, the Results and Discussion relationships between chromosome numbers Chromosome numbers determined in this and taxonomic treatments of the C. atrofusca study are shown in Table 1. None of the species complex are still unresolved. Further, large scale had a primary constriction in their chromosomes. studies with material from the whole distribution range of this species complex are needed to 1. Blysmus compressus (L.) Panz. ex Link (2n = fully understand the relationships between 46, Fig. 1A) chromosome numbers and their taxonomic Blysmus compressus is widely distributed treatments. in Europe and Asia (Iran, Pakistan, Nepal, N India, and SW China) (Koyama 1978). 3. Carex filicina Nees (2n = 46, Fig. 1C) Although the chromosome number of European Carex filicina is distributed in the Himalaya, B. compressus has previously been reported Eastern India, east to China and Taiwan, and as 2n = 44 (Håkansson 1928, Tischler 1934, south to Malaysia (Koyama 1978). Chromosome Rohweder 1937, Pogan and Izmailow 1983, numbers of C. filicina have previously been Stoeva 1992a) and 79 (Stoeva 1992b), that of reported as 2n = 42, 44, 46, 48 (Mehra and Himalayan B. compressus has previously been Sachdeva 1971, 1975b, 1979, Sachdeva 1977, reported as 2n = 40 (Yano et al. 2010b; Manaslu Nijalingappa and Leela 1990, Yano et al. 2010b) Himalaya, Central Nepal) and 2n = 44 (Mehra (Table 1). The chromosome number, 2n = 46, and Sachdeva 1975b; Kashimir Himalaya, India, in this study, was congruent with one of the as compressus (L.) Pers.). In the present numbers in the aneuploid series reported by study, a new number, 2n = 46, was observed. Yano et al. (2010b) from Manaslu Himalaya. Somatic metaphase chromosomes were less Somatic metaphase chromosomes were less than than 1.1 µm in length. Our present result also 1.4 µm in length. supported the observation that B. compressus 252 植物研究雑誌 第 90 巻 第 4 号 2015 年 8 月

4. Carex foliosa D. Don (2n = 62, Fig. 1D) al. 2000, Yano et al. 2010a) (Table 1). In the Carex foliosa is distributed in Bhutan, India, present study the chromosome number, 2n = Nepal and Pakistan (Koyama 1978, Kukkonen 46, corresponds to one of the numbers in the 1987, Noltie 1994). In the present study we aneuploid series reported by Yano et al. (2010a) observed 62 somatic chromosomes, and they from Bajhang district, Far Western Nepal. were less than 1.1 µm in length. This is a new Somatic metaphase chromosomes were less than chromosome number for C. foliosa as it has 1.2 µm in length. previously been reported as 2n = 58 by Mehra and Sachdeva (1971, 1975b: as C. muricata 8. Carex microglochin Wahlenb. (2n = 50, Fig. L. var. foliosa (D. Don) C. B. Clarke) from the 1H) Nainital, Kumaon in India, western Himalaya Carex microglochin is distributed in (Table 1). It seems that C. foliosa may also Kashmir, Nepal and arctic-alpine regions in consist of a series of aneuploids. the Northern Hemisphere (Koyama 1978). The chromosome number of C. microglochin from 5. Carex inanis Kunth (2n = 42, Fig. 1E) Api Himal was 2n = 50, confirming the previous Carex inanis is distributed from the highlands report from Canada (Löve and Löve 1981). of Central Asia, and Kashmir to the Sikkim Somatic metaphase chromosomes were less Himalayas (Koyama 1978). The chromosome than 1.2 µm in length. Chromosome numbers number of C. inanis from Api Himal and for C. microglochin have been reported as 2n = adjacent areas was 2n = 42, confirming our 48 (Dietrich 1972, Yurtsev and Zhukova 1982), previous report from the Manaslu Himalaya 2n = 50 (present study, Löve and Löve 1981) (Yano et al. 2010b). Somatic metaphase and 2n = 58 (Löve and Löve 1956, Moore and chromosomes were less than 1.4 µm in length. Chater 1971, Moore 1981) (Table 1). It seems that C. microglochin may also consist of a series 6. Carex kumaonensis Kük. (2n = 50, Fig. 1F) of aneuploids. Carex kumaonensis is distributed in the Himalaya (Kumaon to Nepal) and China 9. Carex myosurus Nees (2n = 44, Fig. 1I) (Koyama 1978). Carex kumaonensis had the Carex myosurus is distributed in the chromosome number of 2n = 50, and this is the Himalaya and India (Koyama 1978). The first chromosome count to be determined for this chromosome number of C. myosurus from species. Somatic metaphase chromosomes were Api Himal was 2n = 44, confirming a previous less than 1.5 µm in length. Kükenthal (1909) report from the Langtang Himal, Central Nepal assigned C. kumaonensis to subsect. Decorae (Hoshino et al. 2000). Somatic metaphase Kük. together with C. stracheyi Boott ex C. chromosomes were less than 1.3 µm in length. B. Clarke (for further discussion, see 12. C. stracheyi). 10. Carex obscura Nees (2n = 56, Fig. 2A) Carex obscura is distributed in the Himalaya 7. Carex longipes D. Don (2n = 46, Fig. 1G) (Kashmir to Sikkim) and Tibet (Koyama Carex longipes is distributed in the 1978). Carex obscura was found to have the Himalayas (Kashmir to Bhutan), Nilgiri Mts., chromosome number 2n = 56, and this is the Indo-China, and Central China (Koyama 1978). first chromosome count to be determined for The chromosome number of C. longipes has this species. Somatic metaphase chromosomes previously been reported as 2n = 42, 44, 46 were less than 1.0 µm in length. Carex obscura (Sachdeva 1977, Mehra and Sachdeva 1979, belongs to sect. Atratae Kunth and is closely Nijalingappa and Leela 1990, Hoshino et related to C. augustinowiczii Meinsh, C. hallii August 2015 The Journal of Japanese Botany Vol. 90 No. 4 253

Fig. 1. Photomicrograph of somatic metaphase chromosomes of Cyperaceae from Far Western Nepal. A. Blysmus compressus (2n = 46). B. Carex cruenta (2n = 44). C. C. filicina (2n = 46). D. C. foliosa (2n = 62). E. C. inanis (2n = 42). F. C. kumaonensis (2n = 50). G. C. longipes (2n = 46). H. C. microglochin (2n = 50). I. C. myosurus (2n = 44). Bar = 5 µm.

Olney, C. parryana Dewey, C. soyaeensis Kük., 2010b). Our present cytological results also or C. alpina Swartz complex (Kükenthal 1909). support a close relationship among C. obscura, Among these species chromosome numbers C. augustinowiczii, C. hallii, C. parryana, or C. have been reported as 2n = 54, 56, and 66 for alpina complex within sect. Atratae. C. augustinowiczii (Okuno 1939, 1940, Tanaka 1948), 2n = 54 for C. hallii and C. parryana 11. Carex rufulistolon T. Koyama (2n = 46, Fig. (Löve and Löve 1981), and 2n = 52, 54, 56, 60, 2B) 66 for the Carex alpina complex (see Yano et al. Carex rufulistolon is endemic to the alpine 2010a). In Carex, closely related species within regions of Nepal (Koyama 1978, Katsuyama the same section possess similar number of 2008). The chromosome number of C. chromosomes (Heilborn 1924, Yano et al. 2008, rufulistolon from Api Himal was 2n = 46, 254 植物研究雑誌 第 90 巻 第 4 号 2015 年 8 月 confirming our previous report from Manaslu ranged from 2.2 to 3.6 µm. Himal (Yano et al. 2010b). Somatic metaphase Eleocharis uniglumis belongs to subser. chromosomes were less than 1.2 µm in length. Eleocharis (González-Elizondo and Peterson 1997). Several taxa of subser. Eleocharis have 12. Carex stracheyi Boott ex C. B. Clarke (2n = a bimodal karyotype, e.g. 2n = 16 = 4L + 12S 44, Fig. 2C) for E. palustris (L.) Roem & Schult. (Yano et Carex stracheyi is distributed in the Himalaya al 2010a). Yano and Hoshino (2006b) reported (Kumaon and Nepal) (Koyama 1978). Carex 2n = 42 = 8L + 34S for E. kamtschatica (C. A. stracheyi had the chromosome number of 2n = Mey.) Kom. and 8L chromosomes caused by 44, and this is the first chromosome count to be the duplication of 4L chromosomes of taxa with determined for this species. Somatic metaphase 2n = 16. In the present study the chromosome chromosomes were less than 1.4 µm in length. number, 2n = 42 = 8L + 34S for E. uniglumis Although Kükenthal (1909) recognized 18 might have arisen by a similar process. species in subsect. Decorae, the chromosome Although chromosome numbers of 2n = 42– number had been previously reported for only 50 have been reported for E. uniglumis, the most one species, C. insignis Boott, with 2n = 48 frequent and stable chromosome number was 2n (Sachdeva 1977, Mehra and Sachdeva 1979). = 46 (Bureš 1998). Strandhede (1966) reported This study has provided chromosome numbers chromosome numbers of 2n = (41–)42(–43) for for two species in subsect. Decorae, and we now the interspecific hybrid between E. uniglumis have chromosome numbers for three species subsp. uniglumis and E. palustris subsp. vulgaris in the subsection C. stracheyi (2n = 44), C. Walters from Europe. However, it is not known insignis (2n = 48) and C. kumaonensis (2n = 50). whether the chromosome number of 2n = 42 Our cytological results also support the close for E. uniglumis in Nepal has arisen as a result relationship among C. stracheyi, C. insignis, of hybridization, because we did not find E. and C. kumaonensis because of their similar palustris or other species growing sympatrically chromosome numbers. Further cytological in the field. Further studies are needed to studies are needed for other species in subsect. understand the cytological diversification for this Decorae, to understand the relationships among species group in Nepal. members of the subsection. 14. Fimbristylis complanata (Retz.) Link (2n = 13. Eleocharis uniglumis (Link) Schult. (2n = 10, Fig. 2E) 42, Fig. 2D) Fimbristylis complanata is widely distributed Eleocharis uniglumis is widely distributed in pantropical areas, extending northwards to in Europe, Asia (Turkey, Iraq, Iran, Afghanistan, South China and temperate Japan (Koyama Pakistan, India, China, Kazakhstan, Kyrgyzstan, 1978). The chromosome number of F. Mongolia, Uzbekistan), Russia, North America, complanata has previously been reported as 2n and North West Africa (Kukkonen 1998, = 10 (Kral 1971, Nijalingappa 1972, 1975, Rath Smith et al. 2002, Dai and Strong 2010), and and Patnaik 1974), 2n = 16 (Tanaka 1939 as F. was recently recorded from Nepal by Yano complanata var. kraussiana C. B. Clarke, Yano et al. (2014). Eleocharis uniglumis had the and Hoshino 2006a), and 2n = 20 (Sachdeva chromosome number 2n = 42, and the somatic 1977, Rath and Patnaik 1978, Mehra and metaphase chromosomes showed a bimodal Sachdeva 1979) (Table 1). In the present study karyotype with 8 large (L) and 34 small (S) the chromosome number, 2n = 10, confirms the chromosomes. L-chromosomes ranged from previous reports by Kral (1971) from Mexico, 4.4 to 5.8 µm in length while S-chromosomes and Nijalingappa (1972, 1975), Rath and August 2015 The Journal of Japanese Botany Vol. 90 No. 4 255

Fig. 2. Photomicrograph of somatic metaphase chromosomes (A–H) and meiotic metaphase chromosomes (I) of Cyperaceae from Far Western Nepal. A. Carex obscura (2n = 56). B. C. rufulistolon (2n = 46). C. C. stracheyi (2n = 44). D. Eleocharis uniglumis (2n = 42). E. Fimbristylis complanata (2n = 10). F. F. falcata (2n = 22). G. Kobresia esbirajbhandarii (2n = 62). H. Schoenoplectiella fuscorubens (2n = 42). I. Isolepis setacea (2n = 28 = 14II). Allows indicate eight large chromosomes. Bar = 5 µm.

Patnaik (1974) from India. Somatic metaphase Chromosome numbers of F. complanata var. chromosomes ranged from 1.5 to 2.3 µm in complanata f. complanata from India and Nepal length. are 2n = 10 and 20, while f. exalata from Japan Koyama (1959) recognized two forms in has 2n = 16. F. complanata in Asia: (1) f. complanata from India to Taiwan and Malaysia, through Indo- 15. Fimbristylis falcata (Vahl) Kunth (2n = 22, China, and (2) f. exalata T. Koyama (including Fig. 2F) var. kraussiana) from Japan and China. This Fimbristylis falcata is distributed in Bhutan, and previous studies suggest that there may be Ceylon, India, and Nepal (Koyama 1978, cytological differences between these two forms. Noltie 1994). The chromosome number of F. 256 植物研究雑誌 第 90 巻 第 4 号 2015 年 8 月 falcata had previously been reported as 2n = number of 2n = 62 for K. esbirajbhandarii 22 (Sharma and Sarkar 1967–1968, Sanyal which has a unispicate inflorescence fits within and Sharma 1972, Mehra and Sachdeva the polyploid series. 1975a, 1976: as F. junciformis Kunth, Bir et al. 1986, 1988, 1992, Paramjeet et al. 1992) 18. Schoenoplectiella fuscorubens (T. Koyama) and 2n = 44 (Nijalingappa 1975, 1977) from Hayasaka (2n = 42, Fig. 2H) India. The chromosome number of F. falcata Schoenoplectiella fuscorubens is distributed from Api Himal was 2n = 22, confirming the in Bhutan, China, and Nepal (Koyama diploid number of the previous reports. Somatic 1969, 1978, Noltie 1994: as Schoenoplectus metaphase chromosomes ranged from 1.4 to 2.2 fuscorubens (T. Koyama) T. Koyama). µm in length. Schoenoplectiella fuscorubens was found to have the chromosome number of 2n = 42, 16. Isolepis setacea (L.) R. Br. (2n = 28 = 14II, and this is the first chromosome count to be Fig. 2I) determined for this species. Somatic metaphase Isolepis setacea is widely distributed in chromosomes were less than 1.5 µm in length. the Himalayas, Asia, Europe, and Africa Koyama (1969, 2008) noted that this species (Katsuyama 2008). Chromosome number of I. resembled to S. juncoides (Roxb.) Lye or S. setacea has previously been reported as 2n = kandawlayensis (T. Koyama) Hayasaka. Yano et 26, 28, 32 (Davies 1956, Mičieta 1986, Hoshino al. (2010b) reported the chromosome number for et al. 2000, Yano et al. 2010b) (Table 1). In the S. juncoides from Nepal to be 2n = 74, which is present study the chromosome number, 2n = quite different from that of S. fuscorubens in this 28 = 14II, corresponds to one of the numbers in study. Further studies are needed to understand the aneuploid series reported by Mičieta (1986) the phylogenetic relationships among these from Slovakia and Hoshino et al. (2000) from species, using not only cytological but also by Langtang Himal, Central Nepal. Isolepis setacea the molecular analyses. had 14 normal bivalents pairing in meiotic division. The meiotic metaphase chromosomes The authors thank Dr. Annapurna Das and ranged from 0.8 to 2.1 µm in length. Dr. Sushim R. Baral, Department of Plant Resources, Nepal, for their help in conducting 17. Kobresia esbirajbhandarii Rajbh. & H. field research in Nepal. We thank the members Ohba (2n = 62, Fig. 2G) of the field trip in Far Western Nepal in 2012 Kobresia esbirajbhandarii is endemic to for their help for collecting plant materials. This the alpine regions in Nepal (Rajbhandari and study was partly supported by JSPS KAKENHI Ohba 1991). Kobresia esbirajbhandarii was Grant Numbers 23255005 (to H.I.) and found to have the chromosome number of 2n = 25840136 (to O.Y.). 62, and this is the first chromosome count to be determined for this species. Somatic metaphase References chromosomes were less than 0.9 μm in length. Akiyama S., Wakabayashi M. and Ohba H. 1992. Chromosome evolution in Himalayan Impatiens Starr and Ford (2009) reported the monophyly of (Balsaminaceae). Bot. J. Linn. Soc. 109: 247–257. unispicate Kobresia species in the tribe Cariceae, Bir S. S., Cheema P. and Sidhu M. K. 1992. Chromosomal based on DNA sequence data. Yano et al. (2011) analysis of Fimbristylis Vahl in Punjab, North West reported that the unispicate Kobresia species India. Proc. Indian Natl. Sci. Acad. B58: 63–70. showed a polyploid series, 2n = 32, ca. 36, Bir S. S., Kamra S., Sidhu M. K. and Cheema P. 1988. Cytomorphological studies on some members of 52–56, 58, 60–66, ca. 84, ca. 114, and ca. 122, Cyperaceae from North India. J. Cytol. Genet. 23: as well as some aneuploids. The chromosome 14–37. August 2015 The Journal of Japanese Botany Vol. 90 No. 4 257

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矢野興一 a，池田 博 b，C. A. Pendryc，K. R. Rajbhandarid: 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 III. アピ・ヒマラヤ周辺地域（ネパール極西部）より採 集した 18 種の染色体数 ネパール極西部のアピ・ヒマラヤ周辺地域より採集 C. longipes D. Don (2n = 46), C. microglochin Wahlenb. したカヤツリグサ科植物 7 属 18 種について染色体数を (2n = 50), C. myosurus Nees (2n = 44), C. rufulistolon T. 報告した．Carex cruenta Nees (2n = 44), C. kumaonensis Koyama (2n = 46), Eleocharis uniglumis (Link) Schult. Kük. (2n = 50), C. obscura Nees (2n = 56), C. stracheyi (2n = 42 = 8L + 34S), Fimbristylis complanata (Retz.) Boott ex C. B. Clarke (2n = 44), Kobresia esbirajbhandarii Link (2n = 10), F. falcata (Vahl) Kunth (2n = 22), Isolepis Rajbh. & H. Ohba (2n = 62), Schoenoplectiella setacea (L.) R. Br. (2n = 28 = 14II) については，これま fuscorubens (T. Koyama) Hayasaka (2n = 42) の染色体数 での報告と一致した． は今回が初めての報告である．Blysmus compressus (L.) （a岡山理科大学生物地球学部， Panz. ex Link (2n = 46) と C. foliosa D. Don (2n = 62) b東京大学総合研究博物館， は，これまでの報告と異なった染色体数が算定された． cイギリス・エディンバラ王立植物園， Carex filicina Nees (2n = 46), C. inanis Kunth (2n = 42), dネパール・カトマンズ G.P.O. Box 9446）