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.

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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

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).

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.

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.

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).

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.

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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

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chromosome complement seven pairs are metacentric and

three pairs are submetacentric. Based on the nomenclature

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+ 14m + 6sm + 2Sst (Figure 7).

Acknowledgements. We are grateful to Simon J. Owens

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