INTRODUCTION
Tricyrtis (Liliaceae) is a genus of about 20 species
that are classified in four sections in eastern Asia,
ranging from the Himalayas to Japan, Taiwan, and the
Philippines (Gutierrez, 1974; Jessop, 1979; Takahashi,
1987; Mabberley, 1997; Chen and Takahashi, 2000).
The taxonomy of the genus Tricyrtis in Taiwan has been
difficult (Shimizu, 1962; Takahashi, 1976). Five species
with one additional variety were recognized in the 1st
edition of Flora of Taiwan (Liu and Ying, 1978) while
two species with four additional varieties were recognized
in the 2nd edition of Flora of Taiwan (Ying, 2000).
Almost concurrently, four species (with no indication
of infraspecific taxa) were treated by Yang et al. (2001).
Abundant field examination and a thorough biosystematic
study of the genus Tricyrtis in Taiwan led us to recognize
four distinct species, namely T. formosana Baker, T.
lasiocarpa Matsumura, T. ravenii C.-I Peng & C. L. Tiang,
sp. nov., and T. suzukii Masamune on this island. Tricyrtis
Botanical Studies (2007) 48: 357-364.
*
Corresponding author: Email: bopeng@sinica.edu.tw.
ravenii resembles T. formosana in general aspects.
However, a detailed comparison reveals significant
differences in habit, altitudinal distribution, phenology,
trichome types and distribution pattern of purplish-red
speckles on perianth, sizes of stamen and seed, spur size
and characteristics, which support their recognition as a
new species.
The four Taiwanese species of Tricyrtis belong to
two sections, namely sect. Hirtae (T. formosana, T.
lasiocarpa, T. ravenii) and sect. Tricyrtis (T. suzukii).
All but T. formosana are endemic to the island. Tricyrtis
formosana was recorded also from Iriomote Island of the
Ryukyu Archipelago, which is adjacent to the east coast of
Taiwan (Shimabuku, 1997). Gutierrez (1974) reported a
remarkable finding of a new species, Tricyrtis imeldae H.
G. Gutierrez, from Midanao, Philippines, well over 1,500
km south of the genus¡¦ main distribution in East Asia.
Based on the original description, line drawing and the
image of the type specimen of Tricyrtis imeldae, we agree
with Takahashi (1980) and Flores (1997) that T. imeldae
is hardly distinguishable from T. formosana and probably
should be reduced to its synonymy. A key to the species
of Tricyrtis sect. Hirtae is given below.
TaxONOmy
Tricyrtis ravenii (Liliaceae), a new species from Taiwan
Ching-I PENG
1,
*, Choon-Lin TIANG
1
, and Tsai-Wen HSU
2
1
Herbarium (HAST), Research Center for Biodiversity, Academia Sinica, Nangang, Taipei 115, Taiwan
2
Endemic Species Research Institute, 1 Ming-Sheng E. Road, Chi-Chi, Nantou 552, Taiwan
(Received May 22, 2006; Accepted December 29, 2006)
aBSTRaCT.
Tricyrtis ravenii (Liliaceae) is reported here as a new species endemic to Taiwan. It belongs
to sect. Hirtae, which comprises three species heretofore, namely T. hirta (Japan mainland, mainly on the
Pacific side), T. formosana (Taiwan and Iriomote, Japan) and T. lasiocarpa (Taiwan). Bearing a superficial
resemblance to Tricyrtis formosana, specimens of T. ravenii in most herbaria were nearly always erroneously
annotated as the former or its synonyms such as T. stolonifera. Careful morphological studies of these plants
both from the field and in the experimental greenhouse reveal that they differ from T. formosana significantly
in floral details, habit, occur at higher elevations and deserve recognition of a new species. A taxonomic
treatment, line drawings, field and micro-photographs of epidermal characters, karyotype analysis, and a map
showing their distribution on this island is provided. A key is prepared to aid in the identification of all four
species of Tricyrtis sect. Hirtae.
Keywords: Chromosome number; Endemic species; Karyotype; Liliaceae; New species; sect. Hirtae; Taiwan;
Taxonomy; Tricyrtis hirta; Tricyrtis formosana; Tricyrtis imeldae; Tricyrtis ravenii; Tricyrtis lasiocarpa.
pg_0002
358
Botanical Studies, Vol. 48, 2007
KEy TO SPECIES OF TRICYRTIS SECT. HIRTAE
1a. Stem pendent; flowers axillary throughout most of the stem; seeds black purple; plant of Japan ......................... T. hirta
1b. Stem erect or ascending; flowers in bifurcate racemes on summit and upper axils of stem; seeds brown; plant of
Taiwan.
2a. Plant evergreen; ovary/capsule entirely glabrous or rarely with a few glandular hairs basally; perianths and
pedicels covered with both glandular and hirtellous hairs ....................................................................T. formosana
2b. Plant deciduous; ovary/capsule hirtellous or covered with glandular hairs throughout; perianths and pedicels
covered with either glandular or hirtellous hairs.
3a. Lower leaves usually obovate-spatulate; ovary/capsule hirtellous along septicidal furrows; capsules dehiscent
by tranverse slits; spurs 6-6.4 mm long, in divergent pair; stamens 2.4-2.6 mm long; perianths and pedicels
hirtellous
.......................................................................................................................................... T. lasiocarpa
3b. Lower leaves elliptic; ovary/capsule covered with glandular hairs on bulging locular surfaces; capsules
septicidal dehiscent; spurs 2-2.9 mm long, in coalescent pairs; stamens 3.2-4.9 mm long; perianths and
pedicels covered with glandular hairs .................................................................................................. T. ravenii
mm wide; inner tepals lanceolate, 1.75-2.65 cm long,
3-5.5 mm wide, without basal spurs. Filaments white with
reddish spots, 1.57-2.43 cm long, with minute glandular
hairs at base; anthers yellowish, elliptic, 2.2-4.2 mm long.
Ovaries 0.9-1.33 cm long, covered with glandular hairs.
Capsules septicidal-dehiscent. Seeds many, brownish,
elliptic and flat, 1.5-2.6 mm long, (0.7-) 1-2.2 mm wide.
Somatic chromosome number, 2n = 26. Flowering and
fruiting Jul-Dec.
Additional specimens examined. TAIWAN. ILAN
HSIEN: Nanao Hsiang, Taipingshan Forest Recreation
Area, along path side near Chan-An temple, ca. 2,000 m,
Leong 2384 (HAST); Taroko National Park, vicinity of
710 truck rd., ca. 2,400 m, Wagner 6559 (HAST); Tatung
Hsiang, Nanshan Ssuyuan, 1,480 m, Wang & Lin 2302
(TAIF); Su-yuan along highway, Huang 7311, 7329 (TAI);
Tatung Hsiang, Mingchih to Chilan, on moist soil wall
beside road, ca. 1,100 m, Wang 10791 (TAIF); Northen
Cross-island Highway, at entrance of Forest Road 100,
ca. 1,100 m, Yang & Chiou 5141 (TAIF); at Kefa Bridge,
branched road of Central Cross-island Highway, 1,300 m,
Chen 7545 (TAIF); Taipingshan, ca. 1,850-1,900 m, Liou
et al. 530 (TAIF); Chilanshan, Forest Road (¡¥Highway¡¦)
100, starting point, ca. 1,190 m, Clarke & Gardner et
al. 306 (TAIF). HSINCHU HSIEN: Chienshih Hsiang,
Yuanyang Lake Nature Preserve, at road marker ca. 1.5
km on Forest Trail 160, ca. 1,880 m, Leong 2344 (HAST).
TAICHUNG HSIEN: Hoping Hsiang, abundant along
the Forest Road #710, 121¢X21¡¦21¡¦¡¦ E, 24¢X23¡¦13¡¦¡¦ N,
ca. 2,000 m alt. On gentle slope in mixed coniferous-
broadleaf forest. Herb at peak anthesis, flowers pink. At a
semishaded, wet trailside by a creek. 15 Aug 2001, Leong
2403 (HAST); Lishan ("Mt. Leeshan"), hillside wet place,
Feung & Kao 5014 (TAI); Ssuyuan, Lu 12803 (TAIF);
Ssuyuan, ca. 1,900 m, Chiang 1504 (TAIF); Chingshan,
Chiang 734 (TAIF). NANTOU HSIEN: Jenai Hsiang,
Juiyenchi Forests Nature Protected area, ca. 2,300 m,
Leong 1341 (HAST); along trail to Paiyun lodge in Yushan
National Park, 0.5-1.5 km above Tatachia Saddle, ca.
2,700-2,850 m, Lammers 8436 (HAST); Yushan National
Tricyrtis ravenii C.-I Peng & C. L. Tiang, sp. nov.¡X
TYPE: TAIWAN. Nantou Hsien: Yushan National Park,
along trail from Tatachia Saddle to Paiyun Lodge, 120¢X
57¡¦ E, 23¢X28¡¦ N, ca. 2,800 m alt., 6 Sep 1991, Ching-I
Peng 14337 (with B. Bartholomew, T. G. Lammers and
T. K. Lowrey). Holotype: HAST; isotypes: A, CAS, E,
HAST, K, KYO, MO, PE, TAIF, TI, TNM, TUS, US.
°ª¤sªoÂI¯ó Figures 1, 2
Haec species quoad aspectum T. formosanae similis,
sed ab ea habitu deciduo (vs. sempervirente), perianthio
pedicellisque trichomatibus glandularibus tantum
(vs. glandularibus hirtellisque) vestitis, perianthio
maculis rubropurpureis confertis fascias horizontales
parallelas saepe formantibus (vs. fortuito dispersis)
ornato, staminibus longioribus (3.2-4.9 vs. 2.1-3.3 mm),
seminibus majoribus (14-25.6 ¡Ñ 7-21.5 vs. 11.5-21 ¡Ñ
5.5-13 mm), calcaribus brevioribus (2-2.9 vs. 4-5.5
mm) in paria coalescentia (vs. divergentia) dispositis
atque ovario trichomatibus glandularibus ad superficiem
locularem aequaliter dispersis (vs. omnino glabro vel raro
trichomatibus glandularibus ad basem) ornato distinguitur;
etiam altitudines altiores (1,100-2,800 vs. 100-1,500 m)
plerumque habitat.
Erect perennial herb with stolons; stem (35-) 55-115
(-147) cm tall, leafy and usually unbranched, glabrous.
Stolons slender, several from stem base, to 23-40 cm long,
0.2-0.35 cm thick, glabrous. Leaves alternate, 7-16 (-18)
cm long, 1-5 cm wide, glabrous except along venations
beneath. Lower leaves narrowly elliptic-oblanceolate,
cuneate at base; upper leaves broadly lanceolate to
elliptic, cordate at base. Inflorescence terminal and lateral,
bifurcate racemes each (3.5-) 8-13 (-18.9) cm long,
pedicels glandular. Flowers erect; corolla infundibuliform,
glabrous inside, with glandular hairs outside. Tepals 6, in
2 whorls, with purplish-red speckles that often clustered,
forming horizontal bands against a white to pinkish
background on upper surfaces, with a yellowish-orangish
patch toward the base; outer tepals narrowly elliptic,
1.7-2.65 cm long, 5-8.5 mm wide, with a pair of saccate
spurs at base, spurs subglobose, ca. 2-2.9 mm long, 2.2-4
pg_0003
PENG et al. ¡X
Tricyrtis ravenii
, a new species from Taiwan
359
Figure 1. Tricyrtis ravenii. A, Habit; B, Flower; C, Inner perianth; D, Outer perianth; E, Gynoecium; F, Fully mature and dehiscent
capsule; G. Seed; H, Stem base, showing stolons and thick roots (fibrous roots not shown).
pg_0004
360
Botanical Studies, Vol. 48, 2007
Distribution. Tricyrtis ravenii is endemic to Taiwan.
Plants of T. ravenii occur commonly at 1,100 to 2,800
m elevation in Machilus-Castanopsis and Tsuga-Picea
vegetation zones in Taiwan (cf. Su, 1992) in the Central
Mountain Range, but are found also in some limestone
mountains of eastern Taiwan (Figure 3). They usually
grow on soil slope at forest margin or along semi-exposed
road cut or trails in forest, often near small mountain
creeks.
Etymology. The specific epithet commemorates Prof.
Peter H. Raven, mentor of the senior author, for his
enduring support, guidance, and friendship.
Phenology. Tricyrtis ravenii is a deciduous perennial.
Plants set flowers and fruits from July to early December.
The erect main stem produces one to several stolons at
ground level (often covered by grasses) or underground
during the growth season. From the node on the stolon
Park, by Tungpu Shanchuang, ca. 2,600 m alt., Peng
14413 (HAST); swampy roadside, ca. 1,950-2,200 m,
Huang & Hsieh 4863 (TAI); Tungpu to Yushanchienshan,
2,500-2,800 m, Hsu et Hsu 3960 (TAI); Yuanfeng,
2,200-2,400 m, Lin 581 (TAIF); Nanshan to Szuyuan,
1,200-1,800 m, Wu et al. 1464 (TAIF); Hsinlunkang, Lin
322 (TAIF); Tanta, Tanta forest trail, 1,800-2,400 m, Li
157 (TAIF). HUALIEN HSIEN: en route from summit
of Chingshuishan to Taroko National park Headquarters,
ca. 1,540-2,300 m, Ho 1295 (HAST); Hohuanshan open
grassland, ca. 2,000 m, Hsieh 1003 (TAI); Hsiulin Hsiang,
Taroko Natl. Park, en route from Tatung to (hiking)
entrance of Chingshuishan, broadleaf forest, 1,200-1,350
m, Chen & Hung 1135 (TAIF); Hsiulin Hsiang, Hoping
logging trail roadside under the broadleaf forest,
1,200-1,600 m, Lin et al. L1218 (TAIF); Hoping logging
tract between 41 & 46 km, ca. 1,900-2,000 m, Wang et al.
8523 (TAIF).
Figure 2. Tricyrtis ravenii. A, Habitat (Huang 326); B, Habit (Peng 14413); C, Freshly dug up plant (Huang 326); D, Inflorescence,
in part (Peng 5561); E, Flowers and developing fruits (Peng 14413); F-I, Flowers; F, Leong 2403; G, Peng 14504; H, Peng 14337; I,
Huang 100.
pg_0005
PENG et al. ¡X
Tricyrtis ravenii
, a new species from Taiwan
361
produces a young bud with thick roots (3-6 mm in diam.)
beneath (Figures 1-H; 2-C). The young bud develops into
an aerial shoot that are sterile the first year and may flower
the next year in an experimental shade house. Similarly,
plants from seeds begin to flower in the second year under
cultivated condition. On each shoot small prop roots occur
at the node near ground. Both aerial stems and stolons
wither during the winter, leaving several underground
dormant buds with thick storage roots. Such buds may
be physically detached from the mother plant and grow
up into an independent plant in the next season. The
deciduous habit is shared by another endemic species, T.
lasiocarpa Matsumura, in Taiwan (Peng and Hsu, 2003).
Tricyrtis ravenii is restricted to 1,100 to 2,800 m
elevation and flowers only from summer through late
autumn. By contrast, Tricyrtis formosana, with which it is
often confused, is widely distributed in Taiwan, occurring
from as low as 100 to 1,500 m above sea level (Figure 3).
Tricyrtis formosana is an evergreen perennial that flowers
nearly all year round in southern Taiwan and Lanyu,
a small islet off the southeastern coast of Taiwan. We
observed plants of both T. ravenii and T. formosana along
a mountain trail, separated by 5-6 km, in Chingshuishan,
a metamorphosed limestone mountain in eastern Taiwan.
Natural hybrids, however, were not found.
Morphological notes. Specimens we examined of
Tricyrtis ravenii in nearly all herbaria were erroneously
annotated as T. formosana or its synonym, T. stolonifera.
Tricyrtis ravenii resembles T. formosana in general aspect,
but is distinct in that the perianths and pedicels are covered
only with glandular (vs. both glandular and hirtellous)
hairs; purple-red speckles on perianth are clustered, often
forming parallel, horizontal bands (vs. speckles scatteredly
and randomly distributed); longer stamens (3.2-4.9 vs.
2.1-3.3 mm); larger seeds (14-26.5 ¡Ñ 7-21.5 vs. 11.5-21
¡Ñ 5.5-13 mm); shorter spurs (2-2.9 vs. 4-5.5 mm) that
are coalescent (vs. divergent); glandular hairs evenly
distributed on bulging locular surface of the ovary (vs.
glabrous throughout or rarely with a few glandular hairs at
ovary base) (Figures 4, 5).
Chromosome Cytology. The basic chromosome number
of the genus Tricyrtis is X = 13 with diploids being
prevalent (Takahashi, 1980). Diploid chromosome number
of 2n = 26 was previously reported for two species of
Tricyrtis in Taiwan, namely T. formosana (Nawa, 1928;
Sinoto and Kikkawa, 1932; Sato, 1939 [as ¡¥T. formosana
var. amethystina¡¦ and ¡¥T. formosana var. kotoensis¡¦];
Takahashi, 1980) and T. lasiocarpa (as ¡¥T. formosana var.
lasiocarpa,¡¦ Sato, 1939, 1942). Sato (1942) reported a
tetraploid mitotic chromosome count of 2n = 52 for a plant
produced by experimentally treating seeds with colchicine
solution. Hsu (1971), while reporting "¡KThe material
(Tricyrtis formosana) collected at Kueihu (southern
Taiwan) is a diploid, 2n = 26 with large chromosomes,"
indicated in the figure legend of his plate I "Tricyrtis
formosana Bak., diakinesis with 26 bivalents" and listed
n = 26 for T. formosana in his Table 1. The discrepency
in this report confuses readers. In 1972, Hsu re-published
the chromosome count of Tricyrtis formosana (as n = 26),
based on the same voucher. Based on literature review and
our cytological sampling of T. formosana throughout its
range in Taiwan, we suspect that Hsu¡¦s report of n =26 is
erroneous.
Figure 3. Latitudinal and altitudinal distributions of Tircyrtis ravenii (open triangles) and T. formosana (black dots) in Taiwan.
pg_0006
362
Botanical Studies, Vol. 48, 2007
Figure 4. Compa ris on of flowers
of Tircyrtis ravenii (A-D) and T.
formosana ( E - H ). A ,E . F l o w e r
(side view). Note glandular hairs on
peri anths and pedic els in Tirc yr tis
ravenii ( A) a n d g l a n d u l a r a n d
hirtellous hairs on perianths in T.
formosana (E). B, C, F, G. Scanning
e l e c t r o n m i c ro g ra ph s o f a p a i r
of co al es c ent s pu rs co ver ed wit h
glandula r hairs in T. ravenii (B, C)
and a pair of divergent spurs covered
with bot h g landula r a nd hirtel lous
hairs in T. formosana (F, G). D, H.
Scanning electron m icrographs of a
part of ovary, showing glandular hairs
on locular s urface in T. r avenii (D)
and completely glabrous ovary in T.
formosana (H).
pg_0007
PENG et al. ¡X
Tricyrtis ravenii
, a new species from Taiwan
363
Takahashi and Goro Kokubugata for sending reprints on
Tricyrtis. Support of accommodation, research facilities
and travel expense in Japan to one of us (CLT) from
September 17 to 30, 2001 by Kochi Women¡¦s University is
appreciated. We are indebted to Roy Gereau for assistance
with the Latin diagnosis. We thank curators of HAST,
TAI, and TAIF for facilitating herbarium studies; Jr-Jen
Chen, Kuo-Fang Chung, Ya-Yi Huang, Chia-Hua Lin, and
Wai-Chao Leong for field assistance; and Hui-Jun Hsiao
for technical assistance. This study was supported in part
by research grants from National Science Council and
Academia Sinica, Taiwan to Ching-I Peng.
LITERATURE CITED
Chen, X. and H. Takahashi. 2000. Tricrytis. In Z. Y. Wu and P. H.
Raven (eds.), Flora of China 24: 151-153.
Flores , R.A.Q. 1997. Determination of the rank and pos ition
of a species of Tricyrtis. Undergraduate thesis, Institute of
Biology, College of Science, University of the Philippines
Diliman, Diliman, Quezon City.
Our examination of mitosis in T. ravenii revealed that
it is a diploid with 2n = 26 (Figure 6), which agrees with
most of cytological reports on species of Tricyrtis (cf.
Takahashi, 1980). Chromosomes of T. ravenii range from
1.5 to 4.6 £gm. The karyotype of T. ravenii is asymmetric.
The largest chromosomes (ca. 4.6 £gm, 2 pairs) in the
complement of T. ravenii are approximately 3 times as
long as the smallest ones (ca. 1.5 £gm, one pair). The two
pairs of the largest chromosomes and one pair of the
smallest chromosomes are subtelocentric. In the rest of the
chromosome complement seven pairs are metacentric and
three pairs are submetacentric. Based on the nomenclature
of chromosomes of Levin et al. (1964), T. ravenii has 4Lst
+ 14m + 6sm + 2Sst (Figure 7).
Acknowledgements. We are grateful to Simon J. Owens
and Paul Wilkin of the Royal Botanic Gardens, Kew for
granting permission and facilitating Shen-Horn Yen who
photographed the holotype and other historical specimens
of Tricyrtis formosana for us. We thank Kazuo Oginuma
for guidance in chromosome techniques; and Hiroshi
Figure 6. Microphotograph of somatic metaphase chromosomes
of Tricyrtis ravenii (2n = 26: Leong 2403, HAST).
Figure 5. Comparison of seeds. A, Seeds of Tircyrtis ravenii; B, Seeds of Tircyrtis formosana.
Figure 7. Alignment of somatic metaphase chromosomes of
Tricyrtis ravenii (2n = 26: Leong 2403, HAST).
pg_0008
364
Botanical Studies, Vol. 48, 2007
Gutierrez, H.G. 1974. Tricyrtis imeldae, a new Philippine Lily.
Philipp. J. Sci. 103(3): 171-173.
Hsu, C.C . 19 71. Pre lim inary c hromo som e s tudi es on the
vascular plants of Taiwan (IV). Counts and systematic notes
on some monocotyledons. Taiwania 16(1): 123-136.
Hsu, C.C . 19 72. Pre lim inary c hromo som e s tudi es on the
vascular plants of Taiwan (V). Cytotaxonomy on some
monocotyledons. Taiwania 17: 48-65.
Jessop, J. P. 1979. Liliaceae. In C.G.G.J. van Steenis (ed.), Flora
of Malesiana 9 (pt. 1): 189-235.
Levan, A., K. Fredga, and A.A. Sandberg. 1964. Nomenclature
for centromeric position on chromosomes. Hereditas 52:
201-220.
Liu, T.S. and S.S. Ying. 1978. Liliaceae. In H.L. Li, T.S. Liu, T.C.
Huang, T. Koyama and C.E. DeVol (eds.), Flora of Taiwan,
Vol. 5. Epoch Publ. Co., Ltd., Taipei, pp. 40-84.
Mabberley, D.J. 1997. The Plant-Book, 2nd Edition. Cambridge
Univ. Press, Cambridge, New York, Melbourne, pp. 724.
Nawa, N. 1928. Some cytological observations in Tricyrtis,
Sagittaria and Lilium. Bot. Mag. (Tokyo) 42(493): 33-38.
Peng, J.J. and T.W. Hsu. 2003. Adaption of Tricyrtis lasiocarpa
Mats umura to arid environment. Nat. Cons erv. Quart.
(Taiwan) 41: 56-59.
Sato, D. 1939. Cyto-genetical studies on Tricyrtis, II. Karyotype
analysis in Tircyrtis and Brachycyrtis with special reference
to SAT- and nucleolar chromosomes. Cytologia 10:
127-157.
Sato, D. 1942. Karyotype alternation and phylogeny in Liliaceae
and allied families. Jap. J. Bot. 12: 57-161.
Shimabuku, K.I. 1997. Check List Vascular Flora of the Ryukyu
Islands, Revised Edition. Kyushu University Press, 855 pp.
Shimizu, T. 1962. A note on Tricyrtis of Taiwan. Bot. Bull. Acad.
Sin. 3: 35-37.
S inoto Y. and R. Kikkawa. 1932. Cyto-genetical studies on
Tricyrtis. I. Chromosomes in Tricyrtis. Jap. J. Genet. 7(4):
194-198.
Su, H.J. 1992. Vegetation of Taiwan: altitudinal vegetation zones
and geographical climatic regions. In C.-I Peng (ed.), The
Biological Resources of Taiwan: a Status Report. Inst. Bot.,
Acad. Sin. Monogr. Ser. 11: 39-53.
Takahashi, H. 1976. Studies in Tricyrtis (Liliaceae) 2. On the
Formosan species. Acta Phytotax. Geobot. 27: 169-173. (in
Japanese)
Takahashi, H. 1980. A taxonomic study on the genus Tricyrtis.
Sci. Rep. Fac. Educ., Gifu Univ. (Nat. Sci.) 6: 583-635.
Takahashi, H. 1987. Distribution of Tricyrtis and its
phytogeographical problems. Acta Phytotax. Geobot. 38:
123-132. (in Japanese)
Yang, Y.P., H.Y. Liu, and T.P. Lin (eds.) 2001. Manual of Taiwan
Vascular Plants, Vol. 5. Council of Agriculture, Executive
Yuan, Taipei, pp. 19-32. (in Chinese)
Ying, S.S. 2000. Liliaceae. In D.E. Boufford, C.F. Hsieh, T.C.
Huang, C.S. Kuoh, H. Ohashi, and H.J. Su (eds.), Flora of
Taiwan, 2nd Edition. Vol. 5. Editorial Committee of the
Flora of Taiwan, Department of Botany, National Taiwan
Univerisity, Taipei, pp. 35-71.