Chromosome Botany (2015)10: 61-66 ©Copyright 2015 by the International Society of Chromosome Botany

Karyomorphological study of Taiwanese Tricyrtis () and the taxonomic implication

Yoshiko Kono1, Ching-I Peng2, Chien-Ti Chao3 and Kazuo Oginuma1,4

1The Community Center for the Advancement of Education and Research at the University of Kochi, 5-15 Eikokuji-cho, Kochi 780-8515, ; 2Herbarium (HAST), Biodiversity Research Center, Academia Sinica, Nangang, Taipei 115, ; 3Department of Forestry, National Chung Hsing University, Taichung 402, Taiwan

4Author for correspondence: ([email protected]) Received June 15, 2015; accepted June 30, 2015

ABSTRACT: Taiwanese Tricyrtis consists of two sections and four species, namely sect. Hirtae: T. formosana, T. lasiocarpa, T. ravenii, and sect. Tricyrtis: T. suzukii. The taxonomic treatments of the species in Taiwanese Tricyrtis have been complicated heretofore. To elucidate the inter- and intra-sectional relationships of the two sections, the detailed karyomorphological analyses of all four species were applied in the present study. The chromosome number and the karyotype of T. suzukii were reported here for the first time. Although all species showed the basic chromosome number, x=13, and the diploid chromosome number, 2n=26, the two sections were distinguishable by their karyotypes. However, the intra-sectional karyotypes showed the similarities, i.e., the three species (T. formosana, T. lasiocarpa, T. ravenii) in sect. Hirtae exhibited 2n=26=4Lst+14m+6sm+2st and T. suzukii in sect. Tricyrtis, 2n=26=4Lst+14m+5sm+1st+2t.

KEYWORDS: Karyomorphology, sect. Hirtae, sect. Tricyrtis, Taiwan, , , Tricyrtis lasiocarpa, Tricyrtis ravenii, Tricyrtis suzukii

The Tricyrtis (Liliaceae), ornamentally known as Takahashi (1980) also analyzed the karyotype of T. ‘Toad lily’, is perennial herbaceous endemic to formosana [as ‘T. formosana var. amethystina’ in eastern , including the , , Japan, Nakamura (1968)]. However, the main emphasis of their Taiwan and the (Takahashi 1987; Chen and studies were the existence and position of satellites, thus, Takahashi 2000). A taxonomic revision of the genus was they did not refer on the detailed karyomorphological reported by Takahashi (1980) who classified four sections: features of respective species. Consequently, in the present Brachycyrtis, Flavae, Hirtae, Tricyrtis, and about 20 study we analyzed the karyomorphology of the four species (Takahashi 1980, 1997; Takahashi et al. 2001; species in detail and elucidated the inter- and Takahashi and Koyama 2006). intra-sectional relationships in Taiwanese Tricyrtis. Since Baker (1879) described the first Taiwanese Tricyrtis, T. formosana, the taxonomic treatments of MATERIALS AND METHODS Taiwanese species have been complicated (Table 1): three Plant materials The four species of Tricyrtis species and three forms were recognized in Shimizu investigated here were collected from natural populations (1962); three species and two forms in Takahashi (1976); in Taiwan: T. formosana (Ching-I Peng et al. 23343; four species in Takahashi (1980); two species and five Taichung Shi, Wufeng Hsiang, Niucheng); T. lasiocarpa varieties in Ying (2000); four species in Yang et al. (2001). (Chien-Ti Chao s.n.; Nantou Hsien, Zhushan Zhen, Most recently, Peng et al. (2007) described a new species, Shanlinxi); T. ravenii (Ching-I Peng et al. 22975; Ilan T. ravenii, and categorized four distinct species in Taiwan. Hsien, Nanao Hsiang, Taipingshan Forest Recreation The four Tricyrtis species were composed of two sections, Area); T. suzukii (Chien-Ti Chao, s.n.; Hualien Hsien, i.e., sect. Hirtae: T. formosana, T. lasipcarpa, T. ravenii Xiulin Hsiang, Shakadang Forestry Road). All were and sect. Tricyrtis: T. suzukii (Peng et al. 2007). All cultivated in the experimental greenhouse of Academia species are endemic to Taiwan with the exception of T. Sinica, Taiwan for cytological studies. Voucher specimens formosana. of all plants studied were deposited at HAST Herbarium. Several cytological studies have been reported in the genus (e.g., Sinoto and Kikkawa 1932; Sato 1937, 1939; Chromosome observations Somatic chromosomes were Nakamura 1968; Takahashi 1980; Peng et al. 2007). examined using root tips from plants of the type collection. According to previous reports, the basic chromosome Root tips were pretreated in 2 mM 8-hydroxyquinoline at number of the genus is x=13 and most species are diploids 15-18 ℃ for 6-8 h, then fixed overnight in a 3:1 (2n=26). Concerning the karyotypic analyses on the ethanol-acetic acid solution below 4℃. Chromosomes Taiwanese species, Sato (1939) first reported those of T. were macerated and stained in 2% acetic orcein with 1N formosana and T. lasiocarpa (as ‘T. formosana var. hydrochloric acid (10:1) and observed. Classification of lasiocarpa’), subsequently, Nakamura (1968) and chromosome morphology is based on the centromere 62 KONO ET AL.

Table 1. Taxonomic treatments of Taiwanese Tricyrtis from Shimizu (1962) to Peng et al. (2007) Shimizu Takahashi Ying Yang et al. Peng et al. Takahashi (1976) (1962) (1980) (2000) (2001) (2007) 3 spp., 3 forms 3 spp., 2 forms 4 spp. 2 spp., 5 varieties 4 spp. 4 spp. Sect. Hirtae Sect. Hirtae Sect. Hirtae Sect. Hirtae Sect. Hirtae Sect. Hirtae T. formosana T. formosana T. formosana

f. formosana var. formosana

T. formosana var. grandiflora T. formosana T. formosana f. formosana f. glandulosa var. stolonifera

f. amethystiana f. amethystiana T. lasiocarpa var. lasiocarpa T. lasiocarpa T. lasiocarpa T. lasiocarpa T. lasiocarpa

T. ovatifolia var. ovatifolia T. ovatifolia T. ravenii

Sect. Tricyrtis Sect. Tricyrtis Sect. Tricyrtis Sect. Tricyrtis Sect. Tricyrtis Sect. Tricyrtis

T. suzukii T. suzukii T. suzukii T. suzukii T. suzukii T. suzukii

position, following Levan et al. (1964). metacentric chromosomes (m) in the respective chromosome complements of the three species. Satellites RESULTS AND DISCUSSION were not observed. Therefore, the three species of sect. Currently, the Taiwanese Tricyrtis comprised two sections Hirtae shared the single karyotype of 2n=26=4Lst+14m+ and four species (Takahashi 1980; Peng et al. 2007). All 6sm+2st. four species showed the same chromosome number, 2n=26 (Figs. 1 and 2). Their detailed chromosome features 2. Section Tricyrtis: Tricyrtis suzukii were as follows: Somatic chromosomes at metaphase of T. suzukii was determined here to be 2n=26 (Figs. 1D and 2D) for 1. Section Hirtae: Tricyrtis formosana, T. lasiocarpa, T. the first time. A measurement of somatic chromosomes ravenii in T. suzukii was shown in Table 5. Chromosomes at All three species in sect. Hirtae analyzed in this study mitotic metaphase also showed a bimodal variation in were 2n=26. Karyomorphological features were common chromosome length as same as the three species in sect. to all three species (Figs. 1A-C and 2A-C). Hirtae. Among the 26 chromosomes, the first four Measurements of somatic chromosomes in the three st-chromosomes were longer than the others (ca. species (e.g., chromosome length, relative length and arm 3.7-4.4 µm) and the remaining 22 chromosomes ratio) were summarized in Tables 2-4. Chromosomes at gradually varied from ca. 1.9-2.8 µm long. Except for mitotic metaphase showed a bimodal variation in them, five (arrows in Fig. 1D), five (small arrowheads chromosome length. Among the 26 chromosomes, the first in Fig. 1D) and two (stars in Fig. 1D) were sm-, st- and four subtelocentric chromosomes (st) were longer than the telocentric chromosomes (t), respectively, and the rest others (ca. 4.1-4.4 µm in T. formosana; ca. 3.4-4.1 µm in T. of 14 were m-chromosomes. Satellites were not lasiocarpa; ca. 2.8-3.3 µm in T. ravenii) and the remaining observed. Therefore, the karyotype of T. suzukii was 22 chromosomes gradually varied (ca. 1.6-3.1 µm in T. determined to be 2n=26=4Lst+14m+5sm+1st+2t. The formosana; ca. 1.5-2.5 µm in T. lasiocarpa; ca. 1.4-2.4 µm chromosome morphology of the Nos. 5 and 6 in T. ravenii). Apart from them, they have six (arrows in chromosomes showed to be heteromorphic (Fig. 2D). Figs. 1A-C) and two chromosomes (small arrowheads in Figs. 1A-C) were submetacentric (sm) and st- In this study, the defined chromosomal features of the chromosomes, respectively, and the rest of 14 were four species in Taiwanese Tricyrtis were revealed. Based KARYOMORPHOLOGICAL STUDY OF TAIWANESE TRICYRTIS AND TAXONOMIC IMPLICATION 63

Fig 1. Chromosomes at mitotic metaphase of the four Tricyrtis species in Taiwan (2n=26). A, T. formosana; B, T. lasiocarpa; C, T. ravenii; D, T. suzukii. Large and small arrowheads indicate longer st- and small st-chromosomes, respectively. Arrows indicate sm-chromosomes; stars show t-chromosomes. Scale bar = 5µm.

Fig. 2. Somatic chromosomes of Taiwanese Tricyrtis serially arranged by their chromosome length and centromere position in the chromosome complement of the respective species. A. T. formosana, B. T. lasiocarpa, C. T.ravenii, D. T. susukii. Scale bar = 5µm. 64 KONO ET AL.

Table 2. Measurement of somatic chromosomes in Table 3. Measurement of somatic chromosomes in Tricyrtis formosana at metaphase Tricyrtis lasiocarpa at metaphase Chromo Relative Arm Chromo Relative Arm Length (µm) Form Length (µm) Form -some length ratio -some length ratio 1 0.7 + 3.7 = 4.4 6.7 5.3 st 1 0.9 + 3.2 = 4.1 6.9 3.6 st 2 0.8 + 3.6 = 4.4 6.7 4.5 st 2 0.7 + 3.4 = 4.1 6.9 4.9 st 3 1.0 + 3.4 = 4.4 6.7 3.4 st 3 0.6 + 3.1 = 3.7 6.2 5.2 st 4 0.8 + 3.3 = 4.1 6.2 4.1 st 4 0.5 + 2.9 = 3.4 5.7 5.8 st 5 1.3 + 1.8 = 3.1 4.7 1.4 m 5 1.1 + 1.4 = 2.5 4.2 1.3 m 6 1.3 + 1.7 = 3.0 4.6 1.3 m 6 1.1 + 1.4 = 2.5 4.2 1.3 m 7 1.3 + 1.6 = 2.9 4.4 1.2 m 7 0.9 + 1.4 = 2.3 3.9 1.6 m 8 1.2 + 1.4 = 2.6 3.9 1.2 m 8 1.0 + 1.3 = 2.3 3.9 1.3 m 9 0.8 + 1.8 = 2.6 3.9 2.3 sm 9 0.6 + 1.6 = 2.2 3.7 2.7 sm 10 0.7 + 1.8 = 2.5 3.8 2.6 sm 10 0.6 + 1.6 = 2.2 3.7 2.7 sm 11 1.0 + 1.4 = 2.4 3.6 1.4 m 11 0.6 + 1.6 = 2.2 3.7 2.7 sm 12 1.0 + 1.4 = 2.4 3.6 1.4 m 12 0.6 + 1.6 = 2.2 3.7 2.7 sm 13 0.8 + 1.6 = 2.4 3.6 2.0 sm 13 1.1 + 1.1 = 2.2 3.7 1.0 m 14 0.8 + 1.4 = 2.2 3.3 1.8 sm 14 1.0 + 1.1 = 2.1 3.5 1.1 m 15 0.9 + 1.3 = 2.2 3.3 1.4 m 15 0.9 + 1.1 = 2.0 3.4 1.2 m 16 1.0 + 1.2 = 2.2 3.3 1.2 m 16 0.9 + 1.1 = 2.0 3.4 1.2 m 17 0.8 + 1.3 = 2.1 3.2 1.6 m 17 0.9 + 1.1 = 2.0 3.4 1.2 m 18 0.9 + 1.1 = 2.0 3.0 1.2 m 18 0.9 + 1.1 = 2.0 3.4 1.2 m 19 0.6 + 1.4 = 2.0 3.0 2.3 sm 19 0.9 + 1.1 = 2.0 3.4 1.2 m 20 0.7 + 1.3 = 2.0 3.0 1.9 sm 20 0.9 + 1.0 = 1.9 3.2 1.1 m 21 0.8 + 1.0 = 1.8 2.7 1.3 m 21 0.5 + 1.2 = 1.7 2.8 2.4 sm 22 0.8 + 1.0 = 1.8 2.7 1.3 m 22 0.5 + 1.1 = 1.6 2.7 2.2 sm 23 0.3 + 1.5 = 1.8 2.7 5.0 st 23 0.3 + 1.3 = 1.6 2.7 4.3 st 24 0.3 + 1.5 = 1.8 2.7 5.0 st 24 0.3 + 1.3 = 1.6 2.7 4.3 st 25 0.7 + 0.9 = 1.6 2.4 1.3 m 25 0.6 + 0.9 = 1.5 2.5 1.5 m 26 0.7 + 0.9 = 1.6 2.4 1.3 m 26 0.6 + 0.9 = 1.5 2.5 1.5 m

on previous cytological studies (cf. Takahashi 1980; Peng Intra-specific Robertsonian fusion is considered an et al. 2007), the basic chromosome number of Tricyrtis is important event in mammalian chromosomal evolution generally accepted to be x=13, and almost all species (e.g., Piálek et al. 2005), however, relatively rare showed the diploid chromosome numbers (2n=26). All phenomenon in plants [e.g. radiata (Kurita 1987); four plants studied here were diploids with 2n=26 in Lysimachia mauritiana (Oginuma et al. 2004; Kono et al. agreement with 13 species of Japanese and Himalayan 2011)]. However, the exceptional chromosome number species summarized by Takahashi (1980), and three noted above was few, hence, the basic chromosome Taiwanese species previously reported: T. formosana and number of x=13 and the diploids with 2n=26 are prevalent its synonyms T. formosana var. amethystina; T. formosana in the genus Tricyrtis. var. stolonifera and T. formosana var. kotoensis (Sinoto Although cytological studies of the Taiwanese and Kikkawa 1932; Sato 1937, 1939; Nakamura 1968; Tricyrtis without T. suzukii have been reported (e.g., Sato Takahashi 1980); T. lasiocarpa formerly recognized as T. 1939; Nakamura 1968; Takahashi 1980), they mainly formosana var. lasiocarpa (Sinoto and Kikkawa 1932; examined the locations of satellites, thus, the precise Sato 1937, 1939); T. ravenii (Peng et al. 2007). As an karyotypic analyses of the respective species were exception, Sato (1939) found a different chromosome insufficiently described with the exception of T. ravenii number with 2n=25 in T. formosana var. stolonifera reported by Peng et al. (2007). [considering now to be synonym of T. formosana (Ying Comparing the karyotypes of the four Taiwanese 2000)]. The species with 2n=25 had a longer Tricyrtis species, inter-sectional karyomorphological m-chromosome, therefore, Sato (1939) suggested the differentiations (e.g., the number of sm- and st- possible occurrence of the centric (Robertsonian) fusion of chromosomes, existence of t-chromosomes in the two chromosomes of the individual with 2n=26. chromosome complement) were found between two KARYOMORPHOLOGICAL STUDY OF TAIWANESE TRICYRTIS AND TAXONOMIC IMPLICATION 65

Table 4. Measurement of somatic chromosomes in Table 5. Measurement of somatic chromosomes in Tricyrtis ravenii at metaphase Tricyrtis suzukii at metaphase Chromo Relative Arm Chromo Relative Arm Length (µm) Form Length (µm) Form -some length ratio -some length ratio 1 0.6 + 2.7 = 3.3 6.5 4.5 st 1 0.8 + 3.6 = 4.4 6.8 4.5 st 2 0.7 + 2.4 = 3.1 6.1 3.4 st 2 0.6 + 3.4 = 4.0 6.2 5.7 st 3 0.4 + 2.5 = 2.9 5.7 6.3 st 3 0.8 + 2.9 = 3.7 5.7 3.6 st 4 0.4 + 2.4 = 2.8 5.5 6.0 st 4 0.8 + 2.9 = 3.7 5.7 3.6 st 5 1.1 + 1.3 = 2.4 4.7 1.2 m 5 0.6 + 2.2 = 2.8 4.3 3.7 st 6 1.0 + 1.3 = 2.3 4.5 1.3 m 6 0.9 + 1.9 = 2.8 4.3 2.1 sm 7 1.1 + 1.2 = 2.3 4.5 1.1 m 7 1.2 + 1.4 = 2.6 4.0 1.2 m 8 1.1 + 1.1 = 2.2 4.3 1.0 m 8 1.1 + 1.3 = 2.4 3.7 1.2 m 9 0.5 + 1.5 = 2.0 3.9 3.0 sm 9 1.1 + 1.3 = 2.4 3.7 1.2 m 10 0.5 + 1.5 = 2.0 3.9 3.0 sm 10 1.1 + 1.3 = 2.4 3.7 1.2 m 11 0.9 + 1.1 = 2.0 3.9 1.2 m 11 1.1 + 1.2 = 2.3 3.6 1.1 m 12 1.0 + 1.0 = 2.0 3.9 1.0 m 12 1.1 + 1.1 = 2.2 3.4 1.0 m 13 0.8 + 1.0 = 1.8 3.5 1.3 m 13 0.6 + 1.6 = 2.2 3.4 2.7 sm 14 0.7 + 1.0 = 1.7 3.4 1.4 m 14 0.6 + 1.5 = 2.1 3.3 2.5 sm 15 0.5 + 1.2 = 1.7 3.4 2.4 sm 15 1.0 + 1.1 = 2.1 3.3 1.1 m 16 0.5 + 1.2 = 1.7 3.4 2.4 sm 16 1.0 + 1.1 = 2.1 3.3 1.1 m 17 0.8 + 0.9 = 1.7 3.4 1.1 m 17 0.6 + 1.5 = 2.1 3.3 2.5 sm 18 0.7 + 0.9 = 1.6 3.1 1.3 m 18 0.6 + 1.5 = 2.1 3.3 2.5 sm 19 0.6 + 0.8 = 1.4 2.8 1.3 m 19 1.0 + 1.1 = 2.1 3.3 1.1 m 20 0.6 + 0.8 = 1.4 2.8 1.3 m 20 1.0 + 1.1 = 2.1 3.3 1.1 m 21 0.4 + 1.0 = 1.4 2.8 2.5 sm 21 1.0 + 1.0 = 2.0 3.1 1.0 m 22 0.4 + 1.0 = 1.4 2.8 2.5 sm 22 0.9 + 1.1 = 2.0 3.1 1.2 m 23 0.3 + 1.1 = 1.4 2.8 3.7 st 23 0.2 + 1.8 = 2.0 3.1 9.0 t 24 0.3 + 1.1 = 1.4 2.8 3.7 st 24 0.2 + 1.8 = 2.0 3.1 9.0 t 25 0.7 + 0.7 = 1.4 2.8 1.0 m 25 0.9 + 1.0 = 1.9 3.0 1.1 m 26 0.7 + 0.7 = 1.4 2.8 1.0 m 26 0.9 + 1.0 = 1.9 3.0 1.1 m

sections of sect. Hirtae and Tricyrtis, however, not our present cytological findings conformed closely to the intra-sectional. Therefore, the karyotypes of the three classification of Tricyrtis sections proposed by Takahashi species in sect. Hirtae: T. formosana and T. lasiocarpa (1980) and Peng et al. (2007) . (present study), and T. ravenii (Peng et al. 2007) uniformly showed the same karyotype of 2n=26=4Lst+ ACKNOWLEDGEMENTS. We greatly thank Ms. Yumi Kondo 14m+6sm+2st. On the other hand, T. suzukii in sect. for improving the data. This study was supported in part by research Tricyrtis exhibited the specific karyotype, 2n=26=4Lst+ grant to Ching-I Peng from Academia Sinica, Taiwan. 14m+5sm+1st+2t. In particular, the unique karyomorpho- logical feature of the species is the heteromorphic LITERATURE CITED chromosome morphology with sm- and st-chromosomes Baker, J. G. 1879. A synopsis of Colchicaceae and the aberrant (Nos. 5 and 6 in Figs. 2D and 3D), concerning the position tribes of Liliaceae. J. Linn. Soc. Bot. 17: 405-510. of centromeres. Oginuma et al. (unpub. res.) found its Chen, X. and H. Takahashi. 2000. Tricyrtis. In Z.Y. Wu and P.H. chromosomal heteromorphism in other three Japanese Raven (eds.), Flora of China 24: 151-153. species belonging to sect. Tricyrtis. Consequently, it is Kono, Y., Y. Hoshi, H. Setoguchi, M. Yokota, and K. Oginuma. more likely that T. suzukii belongs to sect. Tricyrtis and the 2011. Distribution patterns rDNAs and telomeres and characteristic chromosomal heteromorphism is one of chromosomal rearrangement between two cytotypes of landmark features in the section. Lysimachia mauritiana L. (Primulaceae). Caryologia 64: In summary, the detailed karyomorphological features 91-98. of all four species in Taiwanese Tricyrtis revealed here Kurita, S. 1987. Variation and evolution in the karyotype of stayed constant within the respective sections. Moreover, Lycoris, IV. Intraspecific variation in the 66 KONO ET AL.

Fig. 3. Idiograms of somatic chromosomes at metaphase following Fig. 2 in the four species of Taiwanese Tricyrtis. A, T. formosana; B, T. lasiocarpa; C, T. ravenii; D, T. suzukii. [m- (grey), sm- (red), st- (blue) and t- chromosomes (green).]

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