Botanical Studies (2007) 48: 349-355

*

Corresponding author: E-mail: bopeng@sinica.edu.tw.

INTRODUCTION

Gentiana, the largest genus of Gentianaceae, contains

about 360 species, mainly in temperate Asia but also

common in Europe and North America (Ho and Liu,

2001). Gentians are traditionally used as herbal medicines

in Taiwan and mainland China. Many of them bear

attractive flowers of great horticultural value. Eleven

species and two additional varieties of Gentiana are

known in Taiwan (Wang and Chen, 1998; Chen and

Wang, 1999), mainly in grasslands on high mountains

(to 3,900 m). All but one variety are endemic. Shortly

after the treatment of Gentianaceae was published in the

2nd edition of Flora of Taiwan (Wang and Chen, 1998),

Lomatogonium chilaiensis, representing a newly recorded

genus and a new species (Chen and Wang, 2000), and

Tripterospermum lilungensis, a new species (Chen et al.,

2006), were reported from this island.

During our botanical surveys in 2006, we collected a

unique Gentiana that was not identifiable to any species

known in Taiwan. After consulting relevant floras

and literature (Hayata, 1908, 1917; Yamamoto, 1936;

Masamune, 1943; Liu and Kuo, 1970, 1978; Ho, 1988;

Ying, 1989; Chen and Wang, 1999; Ho and Pringle, 1995;

Ho and Liu, 2001; Li et al., 2001) and abundant herbarium

specimens, we are convinced that it is a new species,

which we describe and illustrate here.

NEW SPECIES

Gentiana bambuseti T. Y. Hsieh, T. C Hsu, S. M. Ku &

C.-I Peng, sp. nov.¡XTYPE: TAIWAN. Chiayi County,

Chuchi Hsiang, Fenchihu, in bamboo forests, ca. 1,500

m, 5 Feb 2006, Tung-Yu Hsieh 101 (holotype: HAST;

isotypes: A, MO). ¦ËªLÀsÁx Figures 1, 2

Haec species affinis Gentianae zollingerii Fawcett,

sed differt herba bienni, foliis basi haud cordatis, margine

scabris et cartilaginosis (vs. anguste membranaceis),

floribus minoribus, ca. 9-12 mm (vs. 14-17 mm) longis,

caule a medio (vs. basi) ramoso, inferne aphyllo (vs.

folioso), calicis lobis apice aristato-acuminatis (vs. acutis).

Biennial herb. Roots fibrous, dark brown. Stem solitary;

lower part covered by leaf litter on bamboo forest floor,

simple, leafless, glabrous, whitish, 3-14 cm long, 0.8-1.5

mm across; upper stem erect, 2.5-5 cm long, glabrous,

simple to much branched from middle, internodes

short, distally with many leaves. Leaves fringed,

margins white cartilaginous, apex aristate-acuminate,

cartilaginous, scabrous; lower leaves larger, suborbiculate

to elliptic, 1.5-2.3 cm long, 1-1.3 cm wide; upper leaves

gradually smaller, narrowly elliptic, lanceolate or linear-

oblanceolate, to 2 cm long. Inflorescences terminal and

axillary, dense, capitulum-like or sometimes cyme-like.

Flowers (1-)3-9(-13), subsessile; calyx tube tubular-

campanulate, ca. 6 mm long, costae 4 or 5, scabrescent;

calyx lobes 4 or 5, ca. 2.5 mm long, 1 mm wide,

lanceolate, margins slightly scabrous, cartilaginous, apex

TaxONOmy

Gentiana bambuseti (Gentianaceae), a new species from

Taiwan

Tung-Yu HSIEH

1

, Tien-Chuan HSU

2

, Yoshiko KONO

3

, Shin-Ming KU

3

, and Ching-I PENG

3,

*

1

Department of Horticulture, National Chung-Hsing University, 250 Kuo Kuang Road, Taichung 402, Taiwan

2

Institute of Ecology and Evolutionary Biology, National Taiwan University, 1 Roosevelt Road, Sec. 4, Taipei 106, Taiwan

3

Herbarium (HAST), Research Center for Biodiversity, Academia Sinica, Nangang, Taipei 115, Taiwan

(Received January 10, 2007; Accepted May 30, 2007)

ABSTRACT.

Gentiana bambuseti T. Y. Hsieh, T. C Hsu, S. M. Ku & C.-I Peng, a new species from Taiwan,

is described and illustrated. This new species resembles G. zollingerii Fawcett, differing in the radical leaves

not cordate, flowers shorter (ca. 9-12 mm long); branches from middle or above; lower half of stem simple

and leafless; leaf margins cartilaginous and scabrescent; and apex of calyx lobes aristate-acuminate. This

species occur spradically in the Alishan Range at 1,400-1,700 m altitude in central Taiwan. It is remarkably

distinct from all congeners in Taiwan in the long, simple, leafless stem base. A somatic chromosome number

of 2n = 60 was determined and the karyotype was analyzed for G. bambuseti. A color plate, line drawings,

distribution map, and SEM microphotographs of seeds and pollen are documented for this new species.

Gentiana bambuseti is highly vulnerable to heavy disturbance during harvesting in the bamboo forests.

Keywords: Chromosome cytology; Gentiana bambuseti; Gentiana zollingerii; Karyotype analysis; New

species; Rare species; Taiwan; Taxonomy.

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Botanical Studies, Vol. 48, 2007

aristate-acuminate; corolla urceolate-tubular, 9-12 mm

long, 4- or 5-lobed, lobes triangular, ca. 1.5 mm long,

apex of lobes white tinged blue or bluish, apex aristate-

acuminate; margins of plicae entire, apically 2-cleft or

erose. Stamens 4 or 5, anthers lanceolate, ca. 1 mm long,

light yellow, filaments 5-6 mm long, attached slightly

above base of corolla tube. Ovary shortly stipitate,

obovoid-ellipsoid, ca. 5-5.5 mm long, 1.3-3 mm wide;

style ca. 2 mm long; stigma bifurcate and spirally recurved

at anthesis. Capsules obovoid to ellipsoid, 10-11 mm long,

2-valved, with obscure serrulate wings along sutures,

exerted beyond persistent corolla. Seeds numerous, ovoid,

Figure 1. Gentiana bambuseti. A, Habit; B, Leaf; C, Pistil; D, Corolla and stamen; E, Calyx; F, Dehiscent capsule; G, Seeds.

HSIEH et al. ¡X

Gentiana bambuseti

, a new species from Taiwan

351

minutely longitudinally reticulate, 0.4-0.45 mm long,

0.2-0.25 mm wide, dark brown.

Additional specimens examined. TAIWAN. CHIAYI

HSIEN: Meishan Hsiang, Reili, in bamboo forest, ca.

1,450 m, 5 Feb 2006, Tung-Yu Hsieh 118 (HAST); Fen-

Rui Ancient Trail, in bamboo forest, ca. 1,600 m alt., 5

Feb 2006, Tien-Chuan Hsu 394 (TAIF); Meishan Hsiang,

Changshuhu, in bamboo forest, ca. 1,650 m alt., 9 Apr

2006, Tung-Yu Hsieh 541 (HAST); in bamboo forest, ca.

1,600 m alt., 9 Apr 2006, Tien-Chuan Hsu 875 (TAI);

Chuchi Hsiang, Chicho, in bamboo forest, ca. 1,450 m alt.,

25 Dec 2006, Tung-Yu Hsieh 932 (HAST); same loc., Tien-

Figure 2. Gentiana bambuseti. A, Habit and habitat; B, Whole plants; C, Leaf, showing the scaberulous margin with white-colored

fringe, and aristate-acuminate apex; D, Upper portion of a flowering plant; E, Flower, viewed from above. F, G, Seed, SEM micropho-

tograph; H-J, Pollen, SEM microphotograph.

352

Botanical Studies, Vol. 48, 2007

and the rest have median centromeres. Satellites were

observed in the distal regions of the short arms of four

metacentric chromosomes. The karyotype formula is 2n

= 60 = 52m+4m

SAT

+4sm (Figure 3). The chromosome

complement with 2n = 60 contains a set of four longer

chromosomes in addition to sets of four submetacentric

and metacentric chromosomes with satellites, which

suggests that G. bambuseti may be a tetraploid derived

from a diploid with 2n = 30 = 26m+2m

S AT

+2sm.

In Gentiana section Chondrophyllae (163 species: Ho

and Liu, 2001) to which this new species belongs (see

below), somatic chromosomes numbers of 2n = 12, 14, 18,

20, 24, 26, 32, 36, 38, 40, 44, 48, 60, 72, 76, 96-98 have

been reported (Chuang et al., 1962; Love and Love, 1975;

Shigenobu, 1982; Kupfer and Yuan, 1996; Ho et al., 1997;

Yuan et al., 1998; Ho and Liu, 2001). Species with 2n =

30, however, has not been reported so far for any member

of this section. A somatic chromosome number of 2n = 60

was known only for one species, G. micantiformis (Yuan

et al., 1998), but its karyotype was not reported. In sect.

Chondrophyllae, karyotypes of species with 2n = 20 have

been reported as 2n = 20 = 18m+2sm for G. haynaldii;

2n = 20 = 18m+2m

S AT

for G. alsinoides, G. anisostemon,

G. asterocalyx, G. prattii, G. spathulifolia; 2n = 20 = 16

m+2 m

S AT

+2 sm for G. pseudoaquatica; 2n = 20 = 12 m+2

sm

SAT

+6 sm for G. pseudosquarrosa (Kupfer and Yuan,

1996; Yuan et al., 1998). Thus, although G. bambuseti (2n

= 60) may be a tetraploid derived from x = 15 as stated

Chuan Hsu 876 (TAIF); Alishan Hsiang, Huti, in bamboo

forest, ca. 1,550 m alt., 25 Dec 2006, Tung-Yu Hsieh 939

(HAST); Alishan Hsiang, Dinhu, in bamboo forest, ca.

1,660 m alt., 31 Dec 2006, Tung-Yu Hsieh 945 (HAST).

NANTOU COUNTY: Hsinyi Hsiang, Hoshe, ca. 3 km to

Shenmu village, in bamboo forest, ca. 1,650 m alt., 28 Dec

2006. Tung-Yu Hsieh 943 (HAST).

Pollen morphology. Pollen grains of Gentiana

bambuseti are prolate in equatorial view, trilobate-circular

in polar view, 36-38 ¡Ñ 23-26 £gm, 3-colporate, colpi long,

narrow, with obscurely thickened margins.

Etymology. The specific epithet ¡¥bambuseti¡¦ means

"dweller in bamboo forest".

Phenology. Seedlings appearing late July. Flowering

October to May; fruiting November to June.

Chromosome Cytology. For chromosome observations,

pretreatment, fixation and staining of chromosomes

followed Oginuma and Nakata (1988). Classification of

chromosome morphology on the basis of the position

of centromere followed Levan et al. (1964). Somatic

chromosomes at mitotic metaphase of Gentiana bambuseti

are 2n = 60, showing a bimodal karyotype (Figure 3). Four

of the 60 chromosomes have median centromeres and are

longer (ca. 3.3-3.8 £gm) than the rest of the chromosomes in

the complement. The remaining 56 chromosomes, ranging

from 1.3 to 2.7 £gm long, show a gradual change in length.

Of the 56 chromosomes, four have submedian centromeres

Figure 3. Somatic chromosomes at mitotic metaphase in Gentiana bambuseti (2n = 60). A,

Large arrowheads indicate longer metacen-

tric chromosomes. Small arrowheads indicate metacentric chromosomes with satellites. Arrows indicate submetacentric chromosomes.

Scale bar = 5 £gm. B, Serial arrangement of chromosomes at mitotic metaphase by chromosome length. Scale bar = 5 £gm.

HSIEH et al. ¡X

Gentiana bambuseti

, a new species from Taiwan

353

Acknowledgments. We thank David E. Boufford (GH)

for improving the manuscript; Qiner Yang (PE) for the

Latin diagnosis; Kongkanda Chayamarit (BKF) for

the loan of selected specimens of Gentiana hesseliana

Hosseus; and Yen-Lin Hsueh for her loan of a vehicle for

the field trips. This study was supported in part by grants

from the Research Center for Biodiversity, Academia

Sinica, Taiwan to Ching-I Peng (HAST).

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above, it is also possible that it represents a hexaploid

derived from x = 10. After 2n = 60 was established by

polyploidization from complements such as 2n = 20 = 16

m+2m

SAT

+2sm, chromosomal changes in the complement

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chromosome fission and subsequent fusion, resulting in

the 2n = 60 = 52m+4m

SAT

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cytological studies are necessary to elucidate the chromo-

somal evolution in members of Gentiana sect. Chondro-

phyllae.

Distribution. Endemic to central Taiwan in the Alishan

Range at 1,400-1,700 m (Figure 4). All populations were

found on the floor of bamboo forests, a habitat different

from the open alpine meadows where all other congeners

in Taiwan are frequently encountered. The oversight of

this new species over the years suggests the necessity of

more thorough inventory in bamboo forests in Taiwan,

which generally have poor biodiversity due to the alle-

lopathic potential of bamboo vegetation (Chou and Hou,

1981). Gentiana bambuseti is highly vulnerable to heavy

disturbance during harvesting in the bamboo forests.

Notes. According to the classification in Flora of China

(Ho and Pringle, 1995), Gentiana bambuseti belongs to

sect. Chondrophyllae, as do 10 of the other 11 species of

Gentiana in Taiwan. Within section Chondrophyllae, Gen-

tiana bambuseti resembles Gentiana zollingerii Fawcett,

differing from the latter in the radical leaves never cordate,

flowers smaller (9-12 mm vs. 14-17 mm long); branches

from middle (vs. from base), with (vs. without) leafless

lower stem, leaf margins scabrous cartilaginous (vs. nar-

rowly membranous) and apex of calyx lobe aristate-

acuminate (vs. acute). Gentiana bambuseti, characterized

by the simple, leafless lower stem and upper stem with

very short internodes and apically many-leaved, and with

flowering branches crowded into a dense capitulum, is

sharply distinct from all 11 known species of Gentiana in

Taiwan. A detailed morphological comparison among G.

bambuseti, G. zollingerii and related species is shown in

Table 1.

Figure 4. Distribution of Gentiana bambuseti (dots) in Taiwan.

354

Botanical Studies, Vol. 48, 2007

HSIEH et al. ¡X

Gentiana bambuseti

, a new species from Taiwan

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