Corresponding author: Tseng Yen-Hsueh (
Academic editor: L. Peruzzi
A taxonomic revision of
Chien-Ti C, Yen-Hsueh T (2019) Taxonomic revision of
Taiwanese species have been studied by several authors from a morphological and karyological standpoint (
Recently, in studying
The study materials were collected from the field and from herbarium specimens. Living plants were cultivated in the greenhouse of the Department of Forestry,
Pollen was collected from fresh anthers of flowers in anthesis. Voucher specimens (Table
Pollen materials of
Taxa | Coll. location | Coll. no. |
---|---|---|
|
Hualien: Yenhai forest track | Chao 1376 |
|
Hsinchu: Yuanyang lake | Chao 1400 |
|
Keelung: Tawulun fort | Chao 1285 |
The distribution of each taxon was determined from the data on herbarium sheets and from our own field records. Only recognised specimens were marked on the map. The conservation rank evaluation followed the protocols of The Red List of Vascular Plants of Taiwan, 2017 (
The habit of
The roots of
The leaves of
The inflorescences of
The pollen grains of
Pollen morphology of
Pollen morphology of
|
|
|
|
polar axis (µm) | 29.12±0.97 | 26.90±0.20 | 27.41±1.32 |
equatorial axis (µm) | 43.31±2.02 | 37.45±3.30 | 39.77±2.21 |
P/E | 0.67±0.02 | 0.78±0.08 | 0.69±0.03 |
shape | oblate | oblate | oblate |
size | medium | medium | medium |
aperture | 1-sulcate | 1-sulcate | 1-sulcate |
sculpture | foveolate | areolate | rugulate |
The chromosome number of
Previous chromosome count of
Taxon | Chromosome number | Reference | |
---|---|---|---|
n | 2n | ||
|
8 |
|
|
16 | |||
32 |
|
||
|
16 | ||
|
14 | ||
16 |
|
Distribution of
Distribution of
Distribution of
Comparisons of
|
|
|
|
---|---|---|---|
habit | evergreen | deciduous | deciduous |
stolon | absent | present | present |
leaf shape | elliptic to oblong | linear-lanceolate | linear-lanceolate |
color of tepal | pale-yellow with reddish apex | white with green base and apex | yellow |
distribution | low to medium altitude mountain area | medium altitudes ca. 1500−2900m | north-eastern part, seashore to low altitude mountains |
According to a study by the
Another taxon, related to
According to the results, the following taxonomic treatments are made:
1 | Plant deciduous, with stolon; leaves lanceolate to linear-lanceolate |
|
– | Plant evergreen, without stolon; leaves elliptic to oblong |
|
2 | Tepals yellow, without spots |
|
– | Tepals white, often with purplish spots, apex green |
|
Perennial herbs, often with short rhizome, sometimes with long creeping stolon, often glabrous, sometimes scabrous. Roots fleshy, glabrous. Stem erect, simple or branched at the upper part, with scale leaves at the lower nodes, persistent. Leaves evergreen or deciduous, simple, alternate,
About 20 species, from Himalaya region, India, Myanmar, Bhutan, Sikkim, Thailand, Vietnam, China, Taiwan, Japan to Korea.
Stem erect, up to 1.3 m, branched at the upper part, lower nodes covered with persistent scale leaves. Leaves evergreen, simple, alternate, elliptic to oblong, 6−8 cm long, 3−6 cm wide, 3−5 nerved, apex acuminate, base attenuate, petiole short, 3−5 mm long, green, sometimes reddish, glabrous, estipulate. Inflorescence pseudoterminal, solitary to 3−5 flowers fascicled, peduncle short, 3−5 mm long, bract absent. Tepals 6, arranged into 2-whorls, spathulate, 1.5−2 cm long, 5−8 mm wide, light yellow to yellow, often with reddish spots near apex, papillate at the base of the inner surface and margin, 3-nerved, base with a short spur, 1−2 mm long, nectary inside. Stamens 6, inserted at the base of tepals, filaments ca. 1 cm long, expansion at the lower part, slightly papillose at proximal part, anthers 2-loculed, basifixed, ca. 3 mm long, longitudinally dehiscent. Ovary superior, ovate to orbicular, ca. 3 mm long, 3-loculed, style slender, 7−9 mm long, stigma 3-fid, pubescent. Fruits berry, globose or compress globose, 6−7 mm long, 4−6 mm in diam., purplish-black. Seeds numerous. 2n = 16.
Distributed in low to medium altitude mountains up to 2,000 m.
Chiayi county: Alishan-Fengshan, 8 Sep 1993, C. M. Wang W00265 (TNM); Yuntan Rest Station-Shuisheliao, 10 Sep 2004, C. M. Wang & Y. M. Hsu 7730 (TNM); Tatienyupupu, 28 Oct 1997, M. Y. Shen 1980 (TAIE); en route from Chiehtung “Villa” to Tienyunshan, 850 m alt., 1 Nov 1985, C. I Peng 8873 (HAST); Juili, 860 m alt., 26 Jan 1997, T. Y. A. Yang et al. 7838 (PPI); Hsinchu county: Kenliaoping, 26 Apr 1994, C. M. Wang W00733 (TNM); on the way from Shihlu to Chingchuan, 27 Jul 1994, K. Y. Wang & T. Y. Liu 75 (TNM); Niaotsui, 21 Oct 2001, C. M. Wang & C. Y. Li 5300 (TNM); Shakaro ancient trail, 1500−1700 m alt., 10 May 2016, P. H. Chen 1096 (PPI); Luoshan forest track, 1000 m alt. 20 Dec 2002, Y. Y. Huang 1309 (HAST); Hualien county: Sanchan, 7 Sep 1996, C. K. Liou et al. 539 (TNM); Yenhai-lindao, 3 Apr 1991, T. Y. Yang et al. 5506 (TNM); Mt. Paliwanshan, 28 Jul 2007, C. H. Chen et al. 8339 (TNM); Shakadang forest road 2 km, 30 Mar 2008, T. Y. A. Yang et al. 20106 (TNM); Huitouwan, 9 Dec 1983, T. C. Huang & S. F. Huang 10200 (TAI); Lanshan, 1900 m alt. 16 Aug 1967, S. C. Hsu 3539 (TAI); Yushan National Park Nanan Recreation Center to Walami, 365 m alt., 16 Apr 1995, H. Y. Shen 683 (HAST); Mukuashan, 700−1000 m alt., 16 Apr 2003, P. J. Lin et al. 159 (HAST), Taorko to Tali, 15 Jun 2007, P. F. Lu 13967 (HAST); Ilan county: Mingchih-Chilan, 15 Apr 2002, C. M. Wang 5468 (TNM); Taipingshan, 5 Apr 2005, C. H. Wu et al. 343 (TNM); Chunchien, 1500–1600 m alt., 30 Jul 1996, Y. C. Chen et al. 17 (HAST); Ssuchi forest road, 1360 m alt., 18 Jul 1999, C. I Huang 399 (HAST). Nanao, 28 Apr 2001, W. F. Ho 1215 (TNM); Mt. Tuli, 450 m alt., 31 Mar 1985, S. Y. Lu 15610 (PPI); Kaohsiung city: Tenchih, 28 Jun 2000, C. M. Wang 4321 (TNM); same loc., 21 May 2016, S. Z. Yang 77621 (PPI); Nanfengshan, 18 Dec 1961, J. M. Liao 788 (TAI); Meilan forest road 8.6 km, 1300 m alt., 10 May 1994, T. Y. Liu et al. 419 (HAST); Southern cross highway, 26 Dec 2000, S. Z. Yang 29567 (PPI); Miaoli county: Shihmen, 20 Mar 1997, M. Y. Shen 1557 (TAIE); Nanchuang shenmiku, 24 Mar 2001, J. H. Lii 428 (TAIF); Taian hot-spring, 300–400 m alt., 30 Jun 1992, J. C. Wang et al. 7707 (HAST, PPI); Tahu, 200−400 m alt., 27 Jan 2007, H. Y. hsieh 138 (PPI); Nantou county: Bihhwu, 2 Sep 1993, C. M. Wang W00160 (TNM); Kunglingshan, 1 May 1991, H. Y. Chen et al. 1118 (TNM); Chitou, 15 May 1988, S. L. Shern & H. M. Su 42 (TNM); same loc., 5 Feb 2010, C. T. Chao 1323, 1324 (
Stem erect, 15−45 cm, lower nodes covered with scale leaf, persistent, branched at the upper part, with creeping stolon. Leaf deciduous, simple, alternate, lanceolate to linear-lanceolate, 3-nerved, 5−6 cm long, 2−3 cm wide, apex acuminate, base obtuse, petiole short, 3−5 mm long, green, sometimes reddish, estipule, glabrous. Inflorescences pseudoterminal, solitary to 2−3 flowers, peduncle 3−5 mm, bract absent. Pedicels 1−1.5 cm long, tepals 6, arranged into 2-whorls, spathulate, white, apex green with violet spots, 1.5−2 cm long, 6−8 mm wide, margin papillose, 3-nerved, base spurred, with nectary. Stamens 6, inserted at base of tepals, filaments 1−1.5 cm long, anthers 3 mm long, 2-loculed, longitudinally dehiscent. Ovary superior, 3-loculed, style slender, subglabrous, 1−1.5 cm long, stigma 3-fid, pubescent. Fruits berry, globose, purplish-black when mature. Seeds numerous. 2n = 16
Distributed in medium altitudes, ca. 1,500 m to 2,900 m.
Chiayi county: Alishan, near Tziyun temple, 14 Apr 1991, T. Y. Yang et al. 5548 (TNM); Alishan, 9 Jul 1981, C. E. Chang 18237 (PPI); on the road side of Mienyueh railroad, 16 Jul 1986, T. Y. A. Yang 3073 (PPI); Tungpu lodge to Chichung, 2400 m alt., 19 Oct 1987, S. Z. Yang 3767 (PPI); Hsinchu county: Yuanyanghu, 1670 m alt., 28 Apr 1995, K. Y. Wang et al. 1059 (HAST); Hualien county: Mt. Muh-kwa, 24 Jul 1961, M. T. Kao K4146 (TAI); Lanshan, 1 Apr 1994, Y. C. Sun 236 (TAIE); Hoping forest track, 1700−2000 m alt., 24 May 1993, S. F. Huang et al. 5128 (TAI); near Shihtung suspension bridge of Patungkuan ancient trail; 1500−1800 m alt., 7 Apr 2009, J. C. Wu 328 (PPI); Taitung county: Hsiangyang in forest, 2350 m alt., 24 Jul 1988, C. I Peng et al. 11876 (HAST); Ilan county: Mt. Taipingshan, 1970 m alt., 3 Apr 1986, T. C. Huang 10813 (TAI); Nan-shan to Chi-li-ting, 1200−1400 m alt., 20 Aug 1969, C. C. Hsu 5834 (TAI); Mt. Nanakotaizan, 14 Jul 1937, SH1330 (PPI); Kaohsiung city: Kuaiku, 5 May 2006, C. M. Wang et al. 8857 (TNM); Chungtzukuan, 20 Dec 2000, S. Z. Yang 25492 (TAIF); Tienchi to Chungchikuang trail, 2000−2400 m alt., 14 Jul 2000, C. C. Hsu 56 (PPI); Nantou county: Tienchi, 16 Jun 1996, S. T. Chiu et al. 3363 (TNM); Tatachiaanbu to entrance of Yushan, 23 Jul 2009, C. T. Chao 838 (
Stem erect, 15−45 cm, covered with persistent scale leaf at lower nodes, branch at the upper part. Leaf deciduous, simple, alternate, lanceolate, 4−10 cm long, 2−4 cm wide, apex acuminate, base obtuse, margin entire, 3-nerved at the base, petiole short, 3−5 mm long, glabrous, estipulate. Inflorescences pseudoterminal, solitary to 3−5 flowers fascicled, peduncle short, 3−5 mm long, bract absent. Tepals 6, arranged into 2-whorls, spathulate, 2−3 cm long, 5−8 mm wide, the inner surface of basal part and margin papillose, base with a short spur, 1−2 mm long, nectary inside. Stamens 6, inserted at base of tepals, filament often expansion at proximal part, 10−15 mm long, glabrous, anthers 2-loculed, basifixed, 3 mm long, longitudinally dehiscent. Ovary superior, 3-loculed, obovate, style 1−1.5 cm long, stigma 3-lobed, pubescent. Fruits berry, purplish-black, seed numerous. 2n = 14.
Distributed in north-eastern part, from seashore to low altitude mountains.
Ilan county: Tali, 20 Apr 1985, W. S. Tang 1324 (TAI); Kueishan island, 31 May 2003, S. T. Chiu et al. 7542 (TNM); Keelung city: Chyngrenhu, 16 May 2007, T. Y. A. Yang et al. 19220 (TNM), Masu, 6 Mar. 1908, T. Kawakami & S. Simada 4311 (Isotype! TAI); Tawulun Fort, 3 Nov 2009, C. T. Chao 1108, 1111 (
We are grateful to Dr. Jenn-Che Wang, Dr. Chiu-Mei Wang, Dr. Kuo-Fang Chung and Mr. Chih-Yi Chang for improving the manuscript, Dr. Wu-Chun Tu and Dr. Hao-You Tzeng for assistance with SEM photography, Dr. Hiroshi Ikeda and Dr. Hidetoshi Nagamasu for the specimen examination of the herbarium of Tokyo University (TI) and Kyoto University (KYO), Ms. Yu-Ting He for her awesome line drawing pictures and Dr. Gary Bentley, an editor with Peerwith, for his English editing of the manuscript. This study was supported by the project grants no. MOST103-2313-B-005-009-MY3 of Ministry of Science and Technology to Yen-Hsueh Tseng.