Bull. Natn. Sci. Mus., Tokyo, Ser. B, 24(2), pp. 53- 59, June 23 , 1998

Morphology and Taxonomic Status ofTricyrtis viridula (Liliaceae)
Hiroshi Takahashil, Xiang-Kun Qin 2 , Si-jun Hao 2and Fumihiro Konta 3
1

Department of Biology, Faculty of Education, Gifu University,Gifu, 501-1 193 Japan
2
Botanical Department, Shanghai Museum of Natura l History,Shanghai 20023 1, PR China
3
Department of Botany, National Science Museum,
Tsukuba, 305- 0005 Japan

Abstract Taxonomic status of Tricyrtis viridula, a new species recently added tothe chinese flora , was discussed based on morphology of the flower, the inflorescence and the rhizome, coloring of the flower including nectar guide, pigmentationin the roots, and the karyotype. Tricyrtis viridula has many featu res common to
those ofT a./finis, T setouchiensis and T pilosa, and it is ass igned to sect. Tricyrtis .
The di stribution map of T viridula and a key to the species of the sect. Tricyrtis aregiven.
Key words : C hinese flora, di stribution, inflorescence, karyotype, nectar guide,taxonomy, Tricyrtis viridula.

Tricyrtis viridula Hir. Takah. is a new species recently described from south andsoutheast China (Takahashi, 1997). It is similar to T pilosa and T macropoda in habitand morphology of flower and/or inflorescence, and has been misidentified very often
as either species in the chinese flora (e.g., Tsi, 1980; Takahashi , 1980; Lin, 1993). Itis also similar to two Japanese species, T a./finis and T setouchiensis, in morphologyof flower and/or inflorescence (Takahashi, 1997). Therefore, it is suggested that T
viridula is closely related to these four species. These species are comprised in thesect. Tricyrtis, together with more two species (Takahashi, 1980; Table I).
We examined morphological characters of Tricyrtis viridula more in detail incomparison with six species in the sect. Tricyrtis. As a result of the present comparative study, a key to the species of the sect. Tricyrtis and a di stribution map of thi s
spec ies are also given.
Materials and Methods
Materials for the morphological observation were collected in the provinces ofZhejiang, Jiangxi and Guizhou in China.

54

Hiroshi Takahashi, Xiang-Kun Qin, Si-jun Hao and Fumihiro KontaPresence of anthraquinoid pigments was tested by the same method used byTakahashi ( 1974) in the roots of some herbarium specimens in SHM as well as ourcollections.
Seeds of Tricyrtis viridula were collected at Feng Yang Mountain in Zhejiang
Province, and seedlings were used for karyotype analysis. Following Takahashi( 1980), after the root tips were pretreated in 0.05% colchicine at about 11 oc for 2.5hours and fixed in acetic acid-ethanol mixture (I : 3) for 5 minutes, they were washedand macerated in 1 N-HCI at 60°C for 20 seconds, and then the root meristem wassquashed in 1% aceto-orcein.
The specimens deposited in KUN, SHM and Tl as well as the plants collected inour expeditions to south and southeast China were used for the distribution analysis.

Results and Discussion
Flower:
The tip of the outer tepals of Tricyrtis viridula is stick-like, being quite different
from those in sects. Brachycyrtis and Flavae (cf. Takahashi, 1980).
The filaments of Tricyrtis viridula are papillate on the basal part, and it is acommon character in sect. Tricyrtis, sect. Brachycyrtis and some plants ofT hirta insect. Hirtae (Takahashi, 1980).
The perianth of Tricyrtis viridula is very similar in shape to that of T affins andT setouchiensis in sect. Tricyrtis; it is patent at the level of about 1/3 from the base(Takahashi , 1997). However, some traits differ from each other. The ground color of
the perianth is white in T affinis and T setouchiensis, while it is greenish white in T
viridula (Figs. 1- 3). Tricyrtis setouchiensis has an orange nectar guide just below thepatent point in the adaxial side of the tepal (Takahashi, 1989; Fig. 1). Tricyrtis affinis
has a nectar guide of a large purple spot in the same area (Takahashi, 1992; Fig. 2).
This spot is clearly different in size from other spots on the tepal. Tricyrtis viridula
has some small purple spots in that area, and mostly has also a large pale-orange spot
there (Fig. 3). These spots must be nectar guides, because bumblebees grope with
their proboscis around them (Fig. 4). The orange spots as nectar guides are the mostcommon in the genus Tricyrtis and may be a plesiomorphic character. The purple
spot as the nectar guide in T affinis is probably a derivative character, and T macropoda probably lost nectar guides (Takahashi, 1994 ). The plants with orange spots in Tviridula may barely retain such a plesiomorphic character.
Flowers of Tricyrtis pilosa are very similar to that of T viridula in shape andcolor (Bot. Mag. t. 4955). However, information about nectar guides is still insufficient.
Inflorescence:
Tricyrtis viridula has the inflorescence composed of some cymose-branchingsMorphology and Taxonomic Status of Tricyrtis viridula
Table I.

The species in the sect. Tricyrtis and their distribution area.

SpeciesDi stribution area

T suzukii
T setouchiensis
T macropoda
T latifolia
T a/finis
T pilosa

TaiwanJapanJapan, Korea, China
Japan , China
Japan
Himal aya , Chin aFigs. 1-4. Flowers of Tricyrtis affinis (I), T setouchiensis (2) and T viridula (3), and bumblebee
foraging on the flower ofT viridula (4). The arrows indicate nectar guides. Bar= I cm.

55

56

Hiroshi Takahashi , Xiang-Kun Qin, Si-jun Hao and Fumihiro Konta

on the stem terminal. The main shoot apex becomes the first flower of the firstbranching. This species usually produces many flowers, and, in that case, make threeor more cymose-branchings (up to five branchings) with up to eight flowers (Figs.
7- 9) . The same type of inflorescence is found in most species of sect. Tricyrtis (Takahash i, 1980). However, they usually have two branchings but no more than three(Figs. 5 and 6). Tricy rtis setouchiensis with many flowers makes some lateral flowering shoots in the axils of the foliage leaves (Fig. 5). Tricyrtis affinis makes sometimes
distinct cymose-branchings, although it mostly appear to hold flowers in the leaf
axils . The cymose-branchings ofT affinis, however, emerge from the upper leaf axils,
and the shoot apex of the main stem never becomes the first flower (Takahashi , 1980).
The same type of inflorescence as Tricyrtis viridula is made in most species ofthe sect. Hirtae (Takahashi, 1980). These inflorescences in which the main shootapex forms the first cymose-branching do not have more than three branchings on theterminal as well , though each branching holds many flowers .
Rhizome and root:
All species of sects. Brachycyrtis and Hirtae and most species of sect. Tricyrtis
produce distinct horizontal stoloniform rhizomes, terminal buds of which develop
into new plants in the following year. Since the internodes are decayed by the next
season, the new plants become independent. Tricy rtis macropoda in sect. Tricyrtis,
however, produces one or rarely two rhizomes which live for several years (Takahashi, 1974). The species of sect. Flavae, on the other hand, produce no horizontal
rhizomes, and the buds of the next generation are produced on the underground partof the stem, i.e., vertical rhizome (Takahashi, 1980). Distinct horizontal rhizomes areformed in T viridula. They are, however, fewer (usually only one) and shorter(0.5- 1.5 cm long) than those in T affinis and T setouchiensis (up to four and 1- 10 cmlong).
No anthraquinoid pigments were detected in any plants of Tricyrtis viridula. Although the pigments in roots are charcteristic of the sect. Tricyrtis , some species contain no pigments (Takahashi , 1974).
Karyotype :The karyotype of Tricyrtis viridula is very similar to that of T affinis and T
macropoda (Takahashi , 1980). The somatic chromosome number is 26 (Fig. 10). Twopairs of large chromosomes are acrocentric, one pair of middle-sized ones is subtelocentric, eight pairs of middle to small sized ones are submetacentric or metacentric
and two pairs of small ones are telocentrics. One pair of the large acrocentrics hassatellites on their short arm s, and one of the middle-sized submetacentrics also has asatellite on the short arm. This karyotype is also similar to that of T setouchiensis,
but satellites on the middle-si zed subtelocentrics, which were recogni zed in the latterspecies, were not observed in T viridula.

Morphology and Taxonomic Status of Tricyrtis viridula

Figs. 5- 9. Inflorescences of Tricyrtis setouchiensis (5), T macropooda (6), and T viridula (7- 9).
Figs. 6 and 7: from Takahashi (1980).

~.-i.

1111 11 :: ;; ua aa ;: a••• aia ......
Fig. I0.

Somatic chromosomes of Tricyrtis viridula. Bar= 5 )1.

57

Hiroshi Takahashi , Xiang-Kun Qin, Si-jun Hao and Fumihiro Konta

58,_.... .....

r······. '.

.. - ~\..

r·····:~.--····.,': ........ ·... i. ;·~ ... · · ,.~·:-· ..··!

t.

J"Fig. 11 .

Distribution map of Tricy rtis viridula.

Although the karyotype of Tricyrtis viridula is similar to that of T formosana
and T hirta as well , the satellites are on the different type of chromosome from eachother and/or on the different arms of chromosomes of the same type (Takahashi,1980, 1991 ). The species of sects. Brachycyrtis and Flavae are different from thesethree species in having no telocentric chromosomes.
Distribution:
Tricyrtis viridula is distributed in south and southeast China; Zhejiang, Jiangxi ,Guizhou, Guangxi, and northeast Yunnan (Fig. 11 ). It grows at forest margins or onforest floors in the upper warm-temperate to the cool-temperate zone. Tricyrtis pilosa
is not found in East China, but in north-eastern Yunnan.

Conclusion and Key to the Species of the Section Tricyrtis
Our investigations of the above morphological characters show that it is reasonable to assigne Tricyrtis viridula to sect. Tricyrtis.
Key to the species of the sect. Tricyrtis is as follows:
I. Roots with pigments of anthraquinoid
2. Perianth spreading obliquely upward ................ .. ....... . .. ....... T suzukii
2. Perianth patent or recurved
3. Perianth patent .. ..... ..... . .. . . . ... . ... .. . .. . .... . .... .. ...... T setouchiensis

Morphology and Taxonomic Status of Tricyrtis viridula

59

3. Perianth recurved ......... ... . . ... . ... . . . ....
T macropoda
I. Roots without pigments of anthraquinoid
2. Perianth spreading obl ique ly upward yellow with red-purp le spot .... T latifolia
2. Perianth patent
3. Perianth white with purple spots .. .. . . . .. ... . . ....
T a./finis
3. Perianth greenish white with purple spots
4. Stem and leaves pubescent. Inflorescence with a few flowers . . .. T pilosa
4. Stem and leaves nearly glabrous. Inflorescence with many flowers ; perianth often with pale orange spots just below the patent point . . . .. .... . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T viridula0•••••••00•••••••••••••••••••••••••

••

•

•

Acknowledgement
This study is supported by Grants-in-Aid for Scientific Research, Ministry ofEducation, Science, Sports and Culture, Japan no. 05041068 (to Dr. Y. Doi) and no.
07041151 (to Dr. H. Koyama) . We would like to thank Dr. H. Koyama and Dr. Y. Doiof the National Science Museum in Japan, Mr. Z. Liu of the Shanghai Museum of
Natural History and other members of the expeditions to East China in 1994 to 1996
for their supports to our field studies. Thanks are also due to the curators ofKUN andTI for permission to examine specimens in the herbaria and to Professor T. Shimizuand Dr. H. Koyama for their kind advice on this manuscript.

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Takahashi, H., 1980. A taxonomic study on the ge nus Tricyrtis. Sci. Rep. Fac. Educ. Gifu Univ. (Nat.
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Takahashi, H. , 1987. Distribution of Tricyrtis and its phylogenetic problems. Acta Phyotax. Geobot., 38 :123- 132. (In Japanese .)Takahashi , H., 1994. Floral bio logy of Tricyrtis macropoda Miq. (Liliaceae). Acta Phyototax. Geobot. ,45 : 33-40.
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