Botanical Studies (2007) 48: 349-355
Corresponding author: E-mail:
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.
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). ˪Lsx 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
Gentiana bambuseti (Gentianaceae), a new species from
, Tien-Chuan HSU
, Yoshiko KONO
, Shin-Ming KU
, and Ching-I PENG
Department of Horticulture, National Chung-Hsing University, 250 Kuo Kuang Road, Taichung 402, Taiwan
Institute of Ecology and Evolutionary Biology, National Taiwan University, 1 Roosevelt Road, Sec. 4, Taipei 106, Taiwan
Herbarium (HAST), Research Center for Biodiversity, Academia Sinica, Nangang, Taipei 115, Taiwan
(Received January 10, 2007; Accepted May 30, 2007)
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.
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
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.
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
+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
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
for G. alsinoides, G. anisostemon,
G. asterocalyx, G. prattii, G. spathulifolia; 2n = 20 = 16
m+2 m
+2 sm for G. pseudoaquatica; 2n = 20 = 12 m+2
+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 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
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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Chen, C.H., J.C. Wang, and Y.C. Chang. 2006. Tripterospermum
lilungshanensis (Gentianaceae), a new species in Taiwan.
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above, it is also possible that it represents a hexaploid
derived from x = 10. After 2n = 60 was established by
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+2sm, chromosomal changes in the complement
o f 2 n = 60 may have been caused by translocation via
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cytological studies are necessary to elucidate the chromo-
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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.
Botanical Studies, Vol. 48, 2007
HSIEH et al. X
Gentiana bambuseti
, a new species from Taiwan
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