Botanical Studies (2006) 47: 199-205.
*
Corresponding author: E-mail: biofv017@ntnu.edu.tw;
Fax: 886-2-29312904.
Tripterospermum lilungshanensis (Gentianaceae), a new
species in Taiwan
Chih-Hsiung CHEN
1
, Jenn-Che WANG
2,
*, and Yung-Chan CHANG
3
1
Department of Botany, National Museum of Natural Science, 1, Guancian Road, Taichung, Taiwan
2
Department of Life Science, National Taiwan Normal University, 88 Ting-Chow Road, Sec 4, Taipei, Taiwan
3
Department of Life Science, National Chung-Hsing University, 250, Kuo Kuang Road, Taichung, Taiwan
(Received April 12, 2005; Accepted November 10, 2005)
ABSTRACT.
A new species Tripterospermum lilungshanensis C. H. Chen & J. C. Wang from Taiwan is
described. Line drawing, color photos, and SEM micrographs of pollen and seed are also provided to aid
in the identification. This species was found at the altitude of ca. 600-1,000 m from Mt. Lilungshan in the
southern part of the Central Mountain Range. The new species is most similar to T. alutaceifolium (T. S. Liu
& C. C. Kuo) J. Murata, which is mainly restricted to northern Taiwan, but differs from the latter by having
shorter leaf blades (2-5 cm vs. 4-9 cm), lanceolate to ovate-lanceolate and slightly spreading calyx-lobes (vs.
ensiform and straight), and an elongated gynophore which results in fruit exerting from the calyx-tube when
mature.
Keywords: Gentianaceae; New species; Taxonomy; Taiwan; Tripterospermum; Tripterospermum
lilungshanensis.
INTRODUCTION
Tripterospermum Blume (Gentianaceae) is a genus of
climbing perennials restricted to Asia (Murata, 1989).
Twenty-five species have been recognized by Murata
(1989) in his worldwide revision. Recently, Hul (2002)
described three new species from Vietnam. The genus is
the sister group of Crawfurdia and Gentiana based on
recent studies from morphology and molecular phylogeny
(Ho et al., 1996; Yuan and Kupfer, 1995). Most members
of the Taiwanese Tripterospermum have been published
by early Japanese taxonomists (Hayata, 1911; Yamamoto,
1929; Masamune, 1938). The first revision was made
by Satake (1951), who recognized four species. Liu and
Kuo (1970) treated Taiwanese Tripterospermum as three
species. Later, they added two infraspecific taxa (Liu and
Kuo, 1974) and subsequently gave it a similar treatment
in the Flora of Taiwan, First Edition (Liu and Kuo, 1978).
Murata (1989) made a comprehensive revision according
to the morphological survey in which six Taiwan species
were treated. Basically, later treatments of Taiwanese
Tripterospermum, e.g. Flora of China (Ho and Pringle,
1995) and Flora of Taiwan, Second Edition (Wang and
Chen, 1998), were the same as Murata¡¦s.
Recently, in our botanical exploration in southern
Taiwan, an unknown species of Tripterospermum was
found. After further field observations and comparisons
with the herbarium specimens in the HAST, NMNS, TAIF,
TAI, and TNU herbaria, we concluded that the taxon is a
new species.
MATERIALS AND METHODS
Materials used in the present studies were collected
from field, pressed and dried for voucher specimens, and
deposited in the herbaria NMNS and TNU. Seeds and
pollen grains for scanning electron microscope (SEM)
study were collected from fresh capsules and flowers of
holotype (Y. C. Chang 21, NMNS).
Pollen grains were treated by the acetolysis method
(Erdtman, 1952) before being dried to the critical point.
Seed and pollen were coated with gold and examined
under Hitachi S3000N scanning electron microscope.
For the comparison of calyx-lobes, eight individuals
o f T. lilungshanensis in Lilungshan and twelve
individuals from four populations of T. alutaceifolium
in northern Taiwan were measured and plotted. The
voucher specimens with geographic information are as
follows: T. lilungshanensis: Pingtung, Lilungshan, Y. C.
Chang 21 (NMNS), C. H. Chen 6267 (NMNS), Lu & Li
1663 (TNU); T. alutaceifolium: Taipei, Chihsingshan,
TAXONOMY
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Botanical Studies, Vol. 47, 2006
Figure 1. Tripterospermum lilungshanensis C. H. Chen & J. C. Wang. 1, Habit; 2, Calyx; 3, Corolla and stamens; 4, Stamens; 5,
Pistil; 6, Berry with gynophore and dissected calyx; 7, Cross section of berry; 8, Seeds. (C. H. Chen 6267).
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Tripterospermum lilungshanensis
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Figure 2. A-C, Tripterospermum lilungshanensis. A, Outside view of flower, showing white corolla tinged with purple near apex; B,
Calyx remains fresh after corolla withered, compare the slightly spreading calyx-lobes with T. alutaceifolium (Figure D); C, Fruit and
partially dissected calyx. Notice the gynophore of fruit is longer than calyx-tube; D, Mature fruit of T. alutaceifolium, showing the
base of fruit included in calyx-tube. Scale bars 1 cm.
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Figure 3. SEM-micrographs of Tripterospermum lilungshanensis. A: seed; B: polar view of pollen grain; C, D, E: equatorial view of
pollen grain. Scale bars 1 mm (A), 10 £gm (B, C, D), 5 £gm (E).
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name "Crawfurdia") using LM and also reported
little interspecific difference. SEM study of pollen
morphology for the Taiwan taxa has been known for
T. lanceolatum only (Chen and Wang, 1999b). Our
palynological observation revealed some similarities with
T. lanceolatum. However, the latter is characterized by the
subprolate to prolate pollen grains with constricted colpi.
Sexine sculpture seems also to display minute differences
between them when compared with the photographs of
Chen and Wang (1999b). However, their description
indicated a wide range of sexine sculpture. Therefore, its
implications for the systematics of this genus call for an
extensively comparative study.
Notes. Based on Murata¡¦s system (Murata, 1989),
the new species belongs to the Tripterospermum sect.
Tripterospermum because of the baccate fruits. The
characters of elongated gynophore and fruit exerted from
calyx-tube occurred also in T. championii Gardner of
SE Asia (Hul, 2003) and T. japonicum (Sieb. & Zucc.)
Maxim. of NE Asia (Murata, 1989), suggesting that they
are allied taxa.
Morphologically, T. lilungshanensis is most similar
to T. alutaceifolium (T. S. Liu & C. C. Kuo) J. Murata.
Y. C. Chang 19 (NMNS); Taipei, Shihting, Ergeshan,
C. C. Chen 892 (TNU); Taipei, Hsiaoyi, Z. W. Lee 38
(TNU); Taipei, Wulai, Y. K. Chen, 280 (HAST); Taipei,
Yangmingshan National Park, C. H. Liu 201 (HAST).
SYSTEMATIC TREATMENT
Tripterospermum lilungshanensis C. H. Chen & J. C.
Wang sp. nov.¡ÐTYPE: TAIWAN. Pingtung Hsien: Shi-
htzu Hsiang, Lilungshan Y. C. Chang 21, 20 Nov 2004
(holotype: NMNS; isotype: NMNS, TNU).
¨½Às¤sªÍ§Î¯ó (Figures 1, 2)
Species T. alutaceifolio affinis sed foliis lamina
ovatolanceolata vel cordata minus quam 5 cm longa et
calycibus lobis lanceolatis vel ovatolanceolatis circa 6-10
mm longis differens.
Perennial herbs. Stems usually spirally twining. Petiole
2-3 cm; leaf blade ovate-lanceolate, ovate to cordate, 2-5
cm long, 1-3 cm wide, base rounded to cordate, margin
entire, apex acute to acuminate. Inflorescences 1-flowered
or cymous; bracts 1-3 pairs. Calyx campanulate; tube
7-9 mm, winged; lobes lanceolate to ovate-lanceolate,
6-10 mm long, 2-3 mm wide; apex acute to acuminate.
Corolla narrowly campanulate, 2.5-3.5 cm long; lobes
triangular, 3-4 mm long, apex acuminate; plicae obliquely
triangular, 1-2 mm long. Filaments linear, 1-2 cm long;
anthers ellipsoid, ca. 1 mm long. Disc 1-1.5 mm long;
ovary narrowly ellipsoid, 7-10 mm long; gynophore 2-3
mm long; style linear, ca. 2 cm long. Berries red-purple,
subglobose to long-ovoid, 8-12 mm long, 4-6 mm across;
fruit gynophore 8-12 mm long (including disc). Seeds dark
purple, ellipsoid to ovoid, triquetrous, ca. 1.5 mm long,
wingless or somewhat winged (Figure 3: A).
Flowering and fruiting Sep.-Feb.
Paratypes. PINGTUNG HSIEN: Shihtzu Hsiang,
Lilungshan, 600-1,000 m, Lu & Li 1663 (TNU, TAI);
same loc., 1,062 m, Huang et al. 16247 (TAI); same loc.,
650-950 m, C. H. Chen 6267 (NMNS).
Distribution. Endemic to Taiwan, presently known
from 700 to 1,000 m in the southern part of the Central
Mountain Range. Found on forest floor or semi-shady
grasslands.
Palynology. Pollen grain (Figure 3: B-E) tricolporate,
isopolar, spheroidal to prolate spheroidal in equatorial
view, 23-28 ¡Ñ 21-28 £gm (P ¡Ñ E); semiangular in polar
view. Colpi long, crassimarginate, ends acuminate, colpus
membranes finely granulate. Ora circular. Sexine striate;
striae meridinally distributed, forking or anastomosing,
0.5-0.7 £gm wide.
The pollen morphology of Gentianaceae has been
studied extensively by Nilsson (1967), in which only
one species of Tripterospermum, T. microphylla, w as
from Taiwan. He concisely pointed out the interspecific
differences in the sexine sculpture of this genus. Huang
(1972)
observed three Taiwan species (under the genus
Figure 4. Leaf-blades of Tripterospermum lilungshanensis
(A-D) and T. alutaceifolium (E-H). A-D and E-H are traced from
mature leaves of different specimens and show the infraspecific
variation of both species respectively. Scale bars 5 cm.
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Botanical Studies, Vol. 47, 2006
The new species has smaller and shorter leaf blades than
the latter (2-5¡Ñ1-3 cm vs. 4-9 ¡Ñ 1.5-3.5 cm) (Figure 4).
The flowers look very much alike. Both species have
white corollas sometimes tinged with purple abaxially
near the apex (Figure 2A). However, they are easily
distinguishable in calyx morphology. Calyx morphology
is an important character in discriminating species within
the genus Tripterospermum (Murata, 1989) and also in the
closely related genus Gentiana (Chen and Wang, 1999a).
Tripterospermum lilungshanensis and T. alutaceifolium
share relatively broad calyx-lobes that retain their fresh
condition even when the corolla is withered or the fruits
have matured and dropped (Figure 2, B-D). However, the
slightly spreading calyx-lobes in the new species (Figure
2B) differ from the erect form in T. alutaceifolium (Figure
2D). The scatter diagram of calyx lobes length and width
(Figure 5) also clearly shows that T. lilungshanensis has
shorter and wider calyx-lobes than T. alutaceifolium. In
addition, the gynophore of T. lilungshanensis is usually
elongated and becomes longer than the calyx-tube.
Consequently, the fruit is exerted from the calyx-tube
when mature (Figure 2C), in contrast to T. alutaceifolium,
in which the gynophore is slightly shorter than the calyx-
tube, and the base of fruit is included within the calyx-tube
(Figure 2D). Their distribution is allopatric, respectively
being restricted to the southern and northern parts of
Taiwan.
Acknowledgments. Thanks are given to three reviewers
for their critical review of this manuscript, and also to
the curators of HAST, NMNS, TAIF, and TAI for the
examination of specimens and kind help. This work was
supported by NSC93-2311-B-178-002 from the National
Science Council, ROC
LITERATURE CITED
Che n, C.H. and J .C. Wang. 1999a. Re visi on of t he genus
Gentiana L. (Gentianaceae) in Taiwan. Bot. Bull. Acad.
Sin. 40: 9-38.
Chen, S.H. and Y.F. Wang. 1999b. Pollen flora of Yuenyang
Lake Nature Preserve, Taiwan (I). Taiwania 44: 82-136.
Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy:
Angiosperma. Almgvist & Wiksell, Uppsala, pp. 50-51.
Hayata, B. 1911. Materials for a Flora of Formosa. J. Coll. Sci.
Imp. Univ. Tokyo 30: 200-205.
Ho, T.N. and J.S. Pringle 1995. Tripterospermum. In Z.-Y. Wu
& P. H. Raven (eds.), Flora of China, vol. 16, Gentianaceae
through Boraginaceae. Science Press (Beijing) and Missouri
Botanic Garden (St. Louis), pp. 6-11.
Ho, T.N., S.W. Liu, and X.F. Lu. 1996. A phylogenetic analysis
of Gentiana (Ge ntiana cea e). Acta Phyt otax. S in. 34:
505-530.
Huang, T.C. 1972. Pollen flora of Taiwan. Natl. Taiwan Univ.
Bot. Dept. Press, Taipei, pp. 117-118, pls. 71-72.
Hul, S. 2002. Nouvelles especes de Crawfurdia,
Tripterospermum et Gentiana (Gentianaceae) du Vietnam.
Adansonia, ser. 3, 24: 27-41.
Hul, S. 2003. Flore du Cambrodge, du Laos et du Vietnam 31,
Gentianaceae: 1-97. Museum National d¡¦histoire Naturelle,
Paris, France.
Liu, T.S. and C.C. Kuo. 1970. Studies on the Taiwan species of
Gentianoideae. Ann. Taiwan Mus. 13: 116-120.
Liu, T.S . and C.C. Kuo. 1974. A taxonomic revis ion of the
species of Gentianaceae in Taiwan. Bull. Exp. Forest Natl.
Taiwan Uniiv. 114: 192-200.
Liu, T.S. and C.C. Kuo. 1978. Tripterospermum. In H.-L. Li et
al. (eds.), Flora of Taiwan, vol. 4. Epoch Publ. Co. Taipei,
pp. 191-201.
Ma sam une, G. 1938. Mis ce llane ous note s on the fl ora of
the Eastern Asia XIII. Trans . Nat. Hist. Soc. F orm. 28:
138-144.
Murata, J. 1989. A synopsis of Tripterospermum (Gentianaceae).
J. Fac. Sci. Univ. Tokyo Sec. III. 16: 273-339.
N il s s on, S . 1 967 . P o ll en m or pho lo gi ca l s tu die s i n t he
Gentianaceae¡XGentianinae. Grana Palynol. 7: 46-145.
S atake, Y. 1951. On the genus Tripterospermum and some
Formosan species. J. Jap. Bot. 26: 103-108.
Wang, J.C. and Y.C. Chen. 1998. Tripterospermum. In T. C.
Huang et al. (eds.), Flora of Taiwan, 2nd Edit., vol. 4. Natl.
Sci. Council R.O.C. pp. 183-190.
Yamamoto, Y. 1929. Contributiones ad floram Formosanam.
Trans. Nat. Hist. Formosa. 19: 104-107.
Yuan, Y.-M. and P. Kupfer. 1995. Molecular phylogenetics of
the subtribe Gentianinae (Gentianaceae) inferred from the
sequences of internal transcribed spacers (ITS) of nuclear
ribosomal DNA. Pl. Syst. Evol. 196: 207-226.
Figure 5. Scatter plotting of calyx lobes length (X axis) and
width (Y axis) of Tripterospermum lilungshanensis (star) and T.
alutaceifolium (triangle).
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et al. ¡X
Tripterospermum lilungshanensis
new species in Taiwan
205