Murata | F-37 | Bot. Bull. Acad. Sin. 37(1)

Bot.
Acad. Sin. (1996)
37: 61-87
WangBull.
— Systematics
of Taiwanese
Arisaema (Araceae)
61
The Systematic Study of Taiwanese Arisaema (Araceae)
Jenn-Che Wang
Department of Biology, National Taiwan Normal University,Wenshan, Taipei, Taiwan 11718, Republic of China
(Received August 11, 1995; Accepted November 14, 1995)
Abstract. The genus Arisaema (Araceae) in Taiwan is revised based on comparative morphological, palynological,
and cytotaxonomical studies. Nine species and one variety, including two new taxa (A. ilanense J. C. Wang sp.
nov. and A. thunbergii ssp. autumnale J. C. Wang, J. Murata & Ohashi ssp. nov.), are recognized. By the recollection of fresh materials and a detailed comparison, A. matsudai Hayata, which was previously reduced to
synonymy under A. grapsospadix Hayata, is now treated as distinct. Rhizome shape, arrangement of leaflets, ratio
of pseudostem to petiole length, and spadix-appendage morphology are all important characters in separating the
Taiwanese taxa. Pollen morphology also displays interspecific variation in size, spinule length, and spinule density.
Cytologically, all examined species have 28 chromosomes (diploid), but three of them are also found to have 56
chromosomes (tetraploid). The chromosome numbers 2n=28 of A. ilanense and A. thunbergii ssp. autumnale, and
2n=56 of A. formosanum and A. heterophyllum are determined for the first time. A key to the taxa along with
descriptions, illustrations, distribution maps, and taxonomic notes are provided.
Keywords: Araceae; Arisaema; Chromosome number; Pollen; Systematics; Taiwan.
Introduction
The genus Arisaema consists of about 150 species
(Murata, 1990a) with the center of distribution and differentiation in warm to cool temperate regions of Asia
(Li, 1980; Murata, 1984). The genus was first established
by Martius in 1831 based on the Himalayan species
(Murata, 1990a). The most important early monograph
was by Engler (1920), who divided the genus into 15 sections based predominantly on the floral morphology. His
system was later revised by Hara (1971), who emphasized
the arrangement of leaflets rather than floral morphology and rearranged the Himalayan species into 13
sections. Hara’s system was adopted by Li (1979) in her
study of the Chinese species. Recently, Murata (1984) reexamined the characters which were used in the previous
studies, and with special reference to fresh materials, he
found many new taxonomic characters. Based on extensive studies— including gross morphology, chromosome
numbers, and pollen morphology— he proposed a revised
system (Murata, 1984, emend 1991) in which the genus
was divided into 11 sections.
The Taiwanese Arisaema have been studied by few
taxonomists. Most species hitherto known were described
by Hayata (1915, 1916, 1920), Hosokawa (1936), and
Kitamura (1941). The first revision of the Taiwanese species was by Huang (1960), who recognized five species
and two forms. His treatment was adopted with little modification by Liu & Huang (1978) in the Flora of Taiwan,
first edition. Subsequently, four taxa, viz. A. formosanum
var. bicolorifolium Huang, A. taiwanense J. Murata, A.
taiwanense var. brevipedunculatum J. Murata and A.
thunbergii ssp. urashima (Hara) Ohashi & J. Murata, have
been reported from Taiwan (Huang, 1982; Murata, 1985;
Wang, 1992).
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In the present paper an intensive revision is made based
on comparative morphological, palynological, and cytological studies. As a result, nine species and one variety,
including two new taxa, are recognized in Taiwan. All of
these taxa are classified into five sections based on the
system proposed by Murata (1984, 1991).
Materials and Methods
Materials used in the present study were collected from
the field throughout Taiwan. Most material was pressed
and dried for voucher specimens deposited in the
Herbarium, Department of Biology, National Taiwan Normal University (TNU). Voucher specimens for pollen observation and chromosome counts were also deposited in
TNU. Living material for the study was cultivated in the
shade house of the Department of Biology, National Taiwan Normal University. In addition, specimens preserved
in the following herbaria were examined: HAST:
Herbarium, Institute of Botany, Academia Sinica, Taipei;
PPI: Herbarium, Department of Forest Resource Management and Technology, National Pingtung Polytechnic
Institute; TAI: Herbarium, Department of Botany, National Taiwan University; TAIF: Herbarium, Taiwan Forestry Research Institute; TNU: Herbarium, Department of
Biology, National Taiwan Normal University.
Although most type specimens were unavailable, photographs of type specimens preserved in the Herbarium
of Tokyo University (TI) (Ohashi, 1981) were examined
by the author.
Pollen grains for scanning electron microscopic (SEM)
study were collected from fresh anthers and prepared by
the method proposed by Erdtman (1952). The acetolyzed
grains were dehydrated through ethanol series and
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acetone, critical point dried, coated with gold, and examined under an ABT DS-180S scanning electron microscope in the Department of Biology, National Taiwan
Normal University.
Root tips for chromosome counts were cut from living
plants. After being pretreated with 0.05% colchicine for
3-5 hours, the material was fixed in a mixture of absolute
alcohol and glacial acetic acid (3:1/v:v), then macerated
with 0.5% pectinase, stained with acetocarmine, squashed,
and observed under a Leitz Aristoplan microscope.
Taxonomic Characters
Previous studies (Engler, 1920; Hara, 1971; Li, 1979)
have shown that the arrangement of leaflets, shape of the
spathe and spadix-appendage, sterile flowers, and dehiscence pattern of anthers were important characters for the
taxonomy of this genus. Moreover, Murata (1984) intensively examined living plants and proposed some new
taxonomic characters such as the growth pattern of the
stem, position of the foliage leaves, phyllotaxy, etc.. Huang
(1960) critically revised the characters used for the taxonomy of the Taiwan species. He also described in detail
the infraspecific variation of characters. The taxonomic
characters used for the present study are concisely described in the following categories:
Stem
The stem of Arisaema is can be either an abbreviated
subterranean tuber or rhizome. The former occurs in all
the Taiwanese taxa except A. grapsospadix. These tubers
are characterized by a more or less depressed globose
shape with a whorl of roots growing at the top around
the pseudostem. As a rule, this kind of stem forms an abscission layer at the base of the new stem and separates
the old stem of the previous year (Murata, 1984). Plants
with tubers often are deciduous and mainly found in the
temperate regions. Taxa with a rhizome have a stem that
is more or less cylindrical in shape with somewhat scattered roots growing on the upper part. Plants with rhizomes are evergreen and mainly found in the tropics. In
Taiwan, the rhizome type stem is uniquely found in A.
grapsospadix and can be used to easily distinguish it from
a similar species, A. matsudai Hayata, which has the tuber type stem.
Leaves
The plants of Arisaema bear two kinds of leaves on
the top of the stem, the outer cataphyll (a sheath-like leaf)
and the inner foliage leaf. The basal part of the petiole
forms a sheath-like pseudostem encircled the peduncle.
The foliage leaves display a wide variation in the number,
arrangement, and shape of leaflets. Most Taiwanese species produce one or two foliage leaves except A.
grapsospadix, which may produce up to three leaves. In
some species the number of foliage leaves is rather stable.
For example, A. thunbergii ssp. autumnale and A.
heterophyllum always produce a single foliage leaf while
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A. ilanense and A. ringens always produce two. However,
in species belonging to Arisaema sections Fimbriata and
Sinarisaema, the number of foliage leaves is unstable. In
these species, as Huang (1960) pointed out, female plants
usually have two leaves and males only one.
The arrangement of leaflets is an important character
in identifying species or species groups. Among Taiwanese species, three patterns of leaflets arrangement can be
recognized, namely trifoliolate, radiate, and pedate compound (Figure 1). These patterns are distinguished not
only by the number of leaflets but also the presence or
absence of petiolules.
Trifoliolate (Figure 1A): Leaves comprise three leaflets,
but sometimes pedate five leaflets in A. grapsospadix. The
leaflets in A. ringens are sessile (Figure 1A) but those of
A. matsudai and A. grapsospadix are petiolulate.
Radiate (Figure 1B): Each leaf comprises more than
seven leaflets which are sessile or subsessile. Also, the
rachis between leaflets is undeveloped so that the leaflets
are palmately arranged (Figure 1B). Species of this type
all belongs to Arisaema section Sinarisaema.
Pedate: Leaves have more than seven leaflets, except
for A. grapsospadix which is 5-foliolate but is characterized by having well-developed rachis. A. grapsospadix,
A. ilanense and Arisaema section Tortuosa (A. thunbergii
ssp. autumnale and A. heterophyllum) belong to this type.
The leaves of A. grapsospadix are characterized by 5foliolate and having a long petiolule on the middle leaflet (Figure 1C). The other three species have more than 7
leaflets in which the middle one is sessile or nearly so.
Among the three taxa, the size of middle leaflet is an important character for identification. In some species (A.
ilanense and A. thunbergii ssp. autumnale, Figure 1E) the
middle leaflet is larger or as large as the adjacent lateral
ones, or much shorter than the adjacent lateral ones in
other species (A. heterophyllum, Figure 1D).
Inflorescence
The inflorescence of Arisaema is a spadix enveloped
by a foliaceous spathe. The upper part of the inflorescenceaxis, called the spadix-appendage, is often covered with
modified sterile flowers (projections). The morphology of
the inflorescence, including spathe, spadix, and spadixappendage, has the most important characters in the taxonomy of the genus and has been used to discriminate
between infrageneric groups (Engler, 1920; Murata,
1984).
Spathe—The shape and coloration of the spathe varies
greatly between species. Basically, the lower part of spathe
forms a cylindrical tube with overlapping spathe-margins
called the spathe-tube which usually gradually opens to
the mouth. The mouth of the spathe-tube often widely
spreads outward with an auricle on each side. In some
species, namely A. grapsospadix, A. matsudai, and A.
heterophyllum, the mouth of the spathe-tube spreads very
narrowly, rendering the auricles inconspicuous. By
contrast, in A. ringens the auricles are particularly
obvious.
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 1. Leaflets arrangement of Arisaema in Taiwan. A, A.
ringens; B, A. formosanum; C, A. grapsospadix; D, A.
heterophyllum; E, A. thunbergii ssp. autumnale.
The upper part of the spathe is laminar and is called
the spathe-blade, which often arches over the spadixappendage. In general, the spathe-blade is triangular or
triangular-ovate with an aristate tip, but some species have
a distinctive feature. For example, the spathe-blade tip of
A. taiwanense is thread-like and up to 40 cm long; the
spathe-blade base of A. heterophyllum enlarges into an
auricle on each side. The spathe-blade of A. ringens is
firstly concave then convex and forms a galeate structure
which is unique to the Taiwanese species.
Coloration of the spathe provides an obvious character,
although it is variable in some taxa especially those of
Arisaema section Sinarisaema. For example, in A.
consanguineum and A. formosanum variously colored
spathes, from green to red, are found in different
individuals. The most conspicuous feature is the white
semilunar mottling on the spathe throat of A.
grapsospadix and A. matsudai. This feature is sometimes
found in A. formosanum, but it is not so obvious and is
somewhat irregular (Figure 3B). In A. grapsospadix, A.
matsudai, and A. heterophyllum both surfaces of the
spathe are green and concolorous. In A. ilanense, the outer
surface is pale to greenish yellow while the inner is reddish brown to dark purple. In A. thunbergii ssp. autumnale
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(Figure 2C), A. ringens (Figure 3D), and the species of
Arisaema section Sinarisaema (Figures 3A-C), the inner
surface of spathe is longitudinally striped, though those
of the Arisaema section Sinarisaema are not always stable
as mentioned above.
Pistillate flowers—The pistillate flowers of the genus
consist of an unilocular ovary which contains one to several orthotropous ovules. The structure is so simple that
previous taxonomists have not considered it important.
Hosokawa (1936) proposed A. taihokense which is separated from A. ringens by having more ovules per ovary.
However, Huang (1960) pointed out that the ovule number in this taxa ranged from two to six and was not a constant character. Therefore, he reduced A. taihokense to
synonymy under A. ringens. The ovary (i.e. pistillate
flower) is usually ovoid and perpendicular to the inflorescence-axis except that of A. grapsospadix which is
bottle-shaped and slanting.
Sterile flowers—The spadix (especially female) of most
Taiwanese Arisaema bears some projections above the fertile flowers (Figures 2, 3). These are considered to be
modified sterile flowers (Engler, 1920; Hara, 1971) because a rudimentary reproductive structure(e.g. ovule or
anther-like organ) is occasionally found (Murata, 1984).
I have also observed that A. heterophyllum bears an anther on the top of some projection. In Taiwan, the modified sterile flowers are found on the spadix of almost all
species except A. thunbergii ssp. autumnale and A. ringens
(Figure 3D). Among the species in Taiwan, two patterns
of modified sterile flowers are recognized. The first type,
found in A. grapsospadix and A. matsudai, bears baccate
modified sterile flowers at the top of spadix-appendage.
The second type, found in most other species, bears echinate sterile flowers just above the fertile flowers, i.e. at
the base of spadix-appendage. The former sometimes also
have linear sterile flowers at the base of the spadix-appendage (especially on the female spadix of A. matsudai)
just like the latter pattern (Figure 2A).
Spadix appendage—Morphology of the spadix-appendage is also greatly diversified (Figures 2, 3) and has been
considered to be important in discriminating species and
infrageneric groups (Li, 1979; Murata, 1984). The spadix-appendage can be divided into sessile and stipitate
types. In the most Taiwanese taxa, the basal part gradually narrows or enlarges, so the appendage is sessile
(Figures 2, 3A & 3B). In A. ringens and some individuals of A. taiwanense the basal part suddenly constricts and
forms a stalk, so the appendage is stipitate (Figures 3C &
3D).
For most species in Taiwan, the spadix-appendage is
erect, cylindrical, and included in the spathe; but in A.
heterophyllum and A. thunbergii ssp. autumnale, the spadix-appendage gradually narrows to form a flagellate tip
which is long-exerted from the spathe (Figures 2B, 2C).
In A. taiwanense, the appendage tip is distinguished by
its rugose surface, an important identifying character. A
key based on the characters of the spadix-appendage is
given below to distinguish each taxa in Taiwan.
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Key to
theand
Taiwanese
Taxa
Based
on the Characters
Figure 2. Inflorescences of Arisaema in Taiwan. A, A. matsudai, female
(left)
male (right);
B, A.
heterophyllum,
male; C, A.
thunbergii ssp. autumnale, male; D, A. ilanense, female.
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 3. Inflorescences of Arisaema in Taiwan (cont.). A, A. consanguineum, female; B, A. formosanum, female (left) and male
(right); C, A. taiwanense, female (left) and male (right); D, A. ringens, female.
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of Spadix-Appendage
Pollen
1. Spadix-appendage filiform, long-exerted from the
spathe.
2. Spadix-appendage black, upper part bent forward
and pendulous (Figure 2C) ....................................
A. thunbergii ssp. autumnale
2. Spadix-appendage green, upper part erect (Figure
2B) ................................................. A. heterophyllum
1. Spadix-appendage short-cylindrical or filiform, included in the spathe.
3. Spadix-appendage tip slender, less than 1.5 mm in
diameter
4. Spadix-appendage tip smooth, without modified
sterile flowers ...........A. formosanum
4. Spadix-appendage tip with baccate or linear sterile flowers.
5. Spadix-appendage tip usually with baccate sterile flowers, base with linear sterile flowers in
mature (female) plant ................. A. matsudai
5. Spadix-appendage tip usually with linear sterile flowers, base glabrous .............................
................................. A. grapsospadix
3. Spadix-appendage tip thick, more than 3 mm in diameter
6. Spadix-appendage tip rugose (Figure
3C)..............................................A. taiwanense
6. Spadix-appendage tip smooth
7. Spadix-appendage stipitate at the base (Figure
3D)...............................A. ringens
7. Spadix-appendage sessile at the base
8. Spadix-appendage erect, cylindrical (Figure
3A).....................A. consanguineum
8. Spadix-appendage incurved, clavate (Figure
2D)................................A. ilanense
Huang (1972) briefly described pollen morphology of
five Taiwanese species, and illustrated three of them, based
on optical microscopic observation. Ohashi & Murata
(1980) and Ohashi, Murata, & Takahashi (1983) reported
palynological differences among Japanese species and suggested that pollen morphology might be useful in distinguishing between Japanese species. Moreover, Murata
(1984) observed 24 species of Arisaema outside Japan, of
which three are present in Taiwan, and pointed out considerable variation between species in pollen size, pollen
shape, size and density of the pollen spinules, and pollen
surface structure.
Pollen grains of Arisaema are generally spherical,
inaperturate, intectate, echinate or spinulate (Murata,
1984). In the present study, pollen grains of eight taxa in
Taiwan were examined with SEM (Figures 4 & 5). In
general, the results were consistent with the above
description. Pollen grains ranged in size from 12µm (in
A. consanguineum, Table 1) to 21µm (in A. thunbergii
ssp. autumnale) in diameter. The surface of grains is uniformly distributed with conical or echinate spinules. The
length and the density of spinules seem distinct for each
species (Table 1). Generally the spinule density is lower
in larger grains. Therefore, the large grains of A.
thunbergii ssp. autumnale have low spinule density (2.4
spinules in an area 5¡Ñ5µm) while the small grains of A.
consanguineum have high density (9.8 spinules in an area
5¡Ñ5 µm). However, A. taiwanense has large grains (18.
2 µm) but the highest spinule density (15 spinules in an
area 5¡Ñ5 µm). Furthermore, the new species A. ilanense
is palynologically unique (Figures 4, C & D) in having
the rather large grain (18.6 µm), with the shortest spinules
(0.4-0.5 µm) and low spinule density (5.2 spinules in an
area 5¡Ñ5 µm) on the surface among the taxa examined
in Taiwan. A. thunbergii ssp. autumnale was previously
misidentified as A. thunbergii ssp. urashima of Japan by
Wang (1992), but when compared with Ohashi & Murata
(1980), the grains of A. thunbergii ssp. autumnale are
slightly different from those of A. thunbergii ssp.
urashima. The pollen grains of A. thunbergii ssp.
Table 1. Pollen grains of Arisaema in Taiwan.
Taxa
Diameter (µm)
A. sect. Fimbriata
A. grapsospadix
A. sect. Clavata
A. ilanense
A. sect. Tortuosa
A. thunbergii ssp. autumnale
A. heterophyllum
A. sect. Sinarisaema
A. consanguineum
A. formosanum
A. taiwanense
A. sect. Pedatisecta
A. ringens
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Spinule length (µm)
Spinules in area 5×5 µm
Voucher specimen
Figure
15.0
1.2–1.7
6.7
Wang 7079
4-A, B
18.6
0.4–0.5
5.2
Wang 6705
4-C, D
20.9
14.2
1.0–1.2
0.7–1.1
2.4
11.1
Lu & Horng 1361
Wang 6741
4-G, H
4-E, F
12.1
12.7
18.2
0.6–0.8
0.9–1.2
0.8–1.0
9.8
9.6
15.0
Deng 354
Wang 7081
Deng 355
5-A, B
5-C, D
5-E, F
16.7
0.7–1.0
4.9
Wang 6455
5-G, H
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 4. Pollen grains of Arisaema in Taiwan. A & B, A. grapsospadix (Wang et al. 7079); C & D, A. ilanense (Wang et al.
6705); E & F, A. heterophyllum (Wang 6741); G & H, A. thunbergii ssp. autumnale (Lu & Horng 1361). Scale bars equal 5 µm.
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Figure 5. Pollen grains of Arisaema in Taiwan (cont.). A & B, A. consanguineum (Deng 354); C & D, A. formosanum (Wang et al.
7081); E & F, A. taiwanense (Deng 355); G & H, A. ringens (Wang 6455). Scale bars equal 5 µm.
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Wang — Systematics of Taiwanese Arisaema (Araceae)
autumnale have shorter spinules (1–1.3 µm) with a somewhat obtuse apex (Figure 4H) while those of A. thunbergii
ssp. urashima have longer spinules (1.7–1.9 µm) with an
acute apex.
On the whole, our knowledge concerning the
palynology, especially of the infraspecific variation, is still
incomplete. A broad comparative study would be necessary to confirm and clarify the above observations.
Chromosome Numbers
All the Taiwanese taxa, except A. ilanense, A. matsudai
and A. thunbergii ssp. autumnale, have been examined
cytologically by Huang (1962), Hotta (1971), and Murata
(1985, 1990b). These previous studies show that nearly
all the species have the chromosome numbers of 2n=28.
However, some species were reported to have infraspecific heteroploidy (Murata, 1990b). For example, A.
consanguineum was reported to have diploid (2n=28) and
tetraploid (2n=56); A. heterophyllum was reported to have
diploid and hexaploid (2n=84) (Murata, 1990b).
The present study re-examined all the Taiwanese taxa,
except A. matsudai. The preliminary results were consistent with the previous studies. All examined species have
2n=28 chromosomes (diploid), but 2n=56 (tetraploid) was
also found in A. consanguineum, A. formosanum and A.
heterophyllum. The chromosome numbers 2n=28 of A.
ilanense and A. thunbergii ssp. autumnale, and 2n=56 of
A. formosanum and A. heterophyllum are determined for
the first time. The cytological study is still in progress
and will be published elsewhere.
Systematic Treatment
ARISAEMA Mart.
Terrestrial herbs. Stem subterranean, rhizomatous, or
tuberous, bearing a peduncle, 1-3 foliage leaves, and several cataphylls at the top. Foliage leaves trifoliolate,
radiate, or pedate compound. Basal part of petiole of foliage leaf connate into a sheath-like pseudostem encircling
the peduncle. Cataphyll sheath-like, surrounding the
pseudostem and peduncle. Inflorescence forming a spadix enveloped by a foliaceous spathe. Lower part of spathe
forming a cylindrical tube (spathe-tube), usually gradually opening to the mouth; upper part laminar (spatheblade), usually hanging over the spadix-appendage. Upper
part of the inflorescence-axis (spadix-appendage) various
in morphology, naked or partly or entirely covered with
modified sterile flowers (projections). Upper part of spadix-appendage elongate or not; basal part sessile or truncate to subtruncate and stipitate. Flowers unisexual,
without perianth, tightly arranged on lower part of inflorescence-axis, paradioecious or in some species
monoecious. Staminate flowers consisting of two to several anthers growing on a columnar stalk. Pistillate flowers consisting of a unilocular ovary. Ovules orthotropous,
one to many in each locule. Fruit a berry, one to severalseeded.
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Key to the Taxa of Taiwanese Arisaema
1. Leaves trifoliolate or pedately 5-foliolate
2. Spadix-appendage filiform, sessile, with baccate or
linear sterile flowers at the tip (Arisaema sect.
Fimbriata)
3. Subterranean stem rhizomatous; plant evergreen
.............................................1. A. grapsospadix
3. Subterranean stem tuberous; plant deciduous
....................................................2. A. matsudai
2. Spadix-appendage cylindrical conical, stipitate, without modified sterile flowers (A. sect.
Pedatisecta).......................................9. A. ringens
1. Leaves pedate or radiate compound, leaflets more than
7
4. Leaves pedate compound
5. Spadix-appendage clavate, within the spathe (A.
sect. Clavata)...............................3. A. ilanense
5. Spadix-appendage filiform, long-exerted from the
spathe tube (A. sect. Tortuosa)
6. Spathe white to pale yellow with dark stripes; anterior portion of spadix-appendage pendulous;
pseudostem and peduncle shorter than petiole;
terminal leaflet as large as or larger than the adjacent lateral ones (A. subsect.
Flagellarisaema).................4. A. thunbergii ssp.
autumnale
6. Spathe green; spadix-appendage erect;
pseudostem and peduncle longer than petiole; terminal leaflet much shorter than the adjacent late r a l o n e s ( A . s u b s e c t . To r t u o s a )
...........................................5. A. heterophyllum
4. Leaves radiate compound (A. sect. Sinarisaema)
7. Spadix-appendage sessile at the base, smooth on
the apex
8. Spadix-appendage columnar, 4-5 mm thick at
apex..............................6. A. consanguineum
8. Spadix-appendage linear, 1-2 mm thick at
apex...................................7. A. formosanum
7. Spadix-appendage stipitate at the base, rugose on
the apex
9. Peduncle 5-15 cm long, spathe tube 3.8-5 cm
long, limb 5-7 cm long..........8. A. taiwanense
9. Peduncle 1-5 cm long, spathe tube 2-4 cm long,
limb 3-5 cm long........8a. A. taiwanense var.
brevipedunculatum
I. Arisaema sect. Fimbriata Engler, Pfl.-reich IV, 23F:
151. 1920; Li in Wu & Li, Fl. Reipubl. Popularis Sin.
13(2): 123. 1979; J. Murata, J. Fac. Sci. Univ. Tokyo
III 13: 466. 1984.—TYPE: Arisaema fimbriatum Mast.
1. Arisaema grapsospadix Hayata, Icon. Pl. Form. 5: 244,
pl. 17. 1915; Kitamura, Acta Phytotax. Geobot. 10:
186. 1941; Huang, J. Forest NTU 27: 8. 1960; Huang,
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Taiwania 7: 94, fig. 2, 8, 9. 1960; Liu & Huang, Quart.
J. Taiwan Mus. 16: 133. 1963; Liu & Huang in H. L.
Li et al., Fl. Taiwan 5: 803. 1978; Li in Wu & Li, Fl.
Reipubl. Popularis Sin. 13(2): 124. 1979.—TYPE:
Formosa. Arisan, inter Funkiko et Taroyen, ad 6000
ped. alt., Mar 1914, Hayata & Ito s.n. (lectotype: TI,
chosen by H. Ohashi, 1981; figured in Ohashi, Univ.
Mus. Univ. Tokyo, Mater. Rep. 5, pl. 14. 1981); loc.
cit., Jan. 1912, Hayata & Sasaki s.n. (syntype: TI).
Figure 9
Arisaema quinquefoliolum Hayata, Icon. Pl. Form. 9: 146.
pl. 7. 1920 (“quinquefoliola”).—TYPE: Formosa. Ako,
Kuwarusu, 1 Jan. 1917, E. Matuda (“Y. Matsuda”) 46
(lectotype: TI, chosen by H. Ohashi, 1981:16; figured
in Ohashi, Univ. Mus. Univ. Tokyo, Mater. Rep. 5, pl.
36. 1981).
Stem rhizomatous, oblong-ovoid, 3-5 cm long, 1.5-2.2
cm in diameter. Pseudostem 13-25 cm long, usually longer
than petiole. Leaves two or three; petiole terete, 5-15 cm
long; lamina ternate or pedately 5-foliolate; leaflets acute
to acuminate at apex, often mucronate on the tip, round
to cuneate at base, entire or near so on margin; terminal
leaflet elliptic to lanceolate, petiolule 1-2.5 cm long; lateral leaflets lanceolate to oblong, oblique at base, petiolule 3-10 mm long. Peduncle 3-13 cm long. Spathe
greenish on both surfaces with a semilunar white mottle
in throat; tube cylindrical, (3-)4-6 cm long, 1-1.7 cm in
diameter; limb triangular ovate, 1.5-2 cm wide, acute to
apex. Spadix bisexual in mature plants, unisexual (male)
in small plants, 5-9 cm long including appendage; staminate flowers sparsely arranged on the inflorescence-axis,
short stalked to nearly sessile; pistillate flowers ovoid,
ascending; appendage filiform, erect or bent, with a bottle
brush-like tip.
Specimens examined. NANTOU: Chitou, 1924,
Siematsu s. n. (TAIF); Fenghuangshan, Peng & Chen
10524 (HAST); Luku Hsiang: Hsitou Exp’t Forest of NTU,
Peng 15016 (HAST). YUNLIN: Tsaoling, Shihpi, Wang
13865 (HAST). CHIAYI: Alishan highway 18-66K, Wang
10253 (HAST); Dardungshan, Wang et al. 7079 (TNU),
Yang 468 (TNU). KAOHSIUNG: Tengchu, Ohashi et al.
13041 (TAI). PINGTUNG: Chachayalaishan, Yang 20090
(TNU); Chunjih Hsiang: Chachayalaishan protection area,
Liao et al. 220 (HAST); Chaopu to Linchang, Province
Road No.9, 471.5K, Wang 16972 (HAST); Chiaoguolats,
Wang et al. 7426 (HAST, TNU); Daibuzan (Mt.
Tawushan), Soma A229 (TAI); Lilungshan, Yang 8382,
8449 (PPI); Manchou to Chufengku, Wang 7394 (HAST);
Mutan, Chang 4786, 5171 (PPI); Peitawushan, Wang et
al. 9090 (TNU); on the way to Tawushan, Chen et al. 535
(HAST); Pu-an-shan, Wang 19038 (HAST); Shantimen
to Chihpen, Province Road No. 22, 58-61K, Wang 13702
(HAST); Shihtzu Hsiang: Lilungshan, Chen & Leu 837,
850 (HAST); Shoka, Wang 13470 (HAST); Wantao to
Nanzenshan, Yang et al. 4326 (TAI); Nanjenshan, Ohashi
et al. 14579 (TAI), Lu & Chen 1407 (TNU), Wang & Lu
5568 (TNU), Chiu s. n. 1993 (TNU); Kuwarusu, Matuda
448 (TAI), A335 (TAI).
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Distribution. Endemic to Taiwan, in the southern part
from low to medium altitudes (Figure 6).
Notes. The species is characterized by its evergreen
habit, rhizomatous subterranean stem, slanting ovary,
bottle brush-like spadix-appendage, and white semilunar
mottle on the throat of the spathe-blade. The first three
characters are very unique and useful in identifying this
taxa. The last two are shared with A. matsudai, and their
distinctions are discussed below. Its foliage leaves are
rather variable, from trifoliolate to pedately 5-foliolate (A.
quinquefoliolum Hayata), with both sometimes found in
the same individual.
2. Arisaema matsudai Hayata, Icon. Pl. Form. 9: 149.
fig. 55. 1920; Li in Wu & Li, Fl. Reipubl. Popularis
Sin. 13(2): 124. 1979.—TYPE: Formosa. Kuwarusu,
April 1917, E. Matsuda (“Y. Matsuda”) s.n. (holotype:
TI, figured in Ohashi, Univ. Mus. Univ. Tokyo, Mater.
Rep. 5, pl. 24. 1981).
(Figure 10)
Tuber subglobose, 1-2(-3)cm in diameter. Pseudostem
8-25(-30)cm long, usually longer than the petiole.
Cataphyllstwo or three, membranous. Leavesone or two;
Figure 6. Distribution of Arisaema grapsospadix (solid circle),
A. matsudai (star), A. ilanense (circle), and A. heterophyllum
(solid star) in Taiwan.
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Figure 7. Distribution of Arisaema thunbergii ssp. autumnale
(star), A. consanguineum (circle), and A. formosanum (solid
star) in Taiwan.
Figure 8. Distribution of Arisaema taiwanense (solid star), A.
taiwanense var. brevipedunculatum (star), and A. ringens (solid
circle) in Taiwan.
petiole terete, 6-12 cm long; lamina trifoliolate, leaflets
acute to acuminate at apex, often with a short tail on tip,
entire or near so on margin; terminal leaflet ovate-lanceolate to rhomboid-elliptic, cuneate at base, petiolule 0.
6-1.5 cm long; lateral leaflets obliquely ovate to lanceolate,
oblique at base, petiolule 2-4 mm long or sometimes near
sessile. Peduncle 4-10 cm long, often shorter than petiole.
Spathe greenish on both surfaces with a semilunar white
mottle on the throat; tube cylindrical, 2.5-3.5 cm long, 0.
6-1 cm in diameter; limb triangular ovate, 1-1.5 cm wide,
acute at apex. Spadix paradioecious, female in mature
plants, male in small plants, 3-4.5 cm long including
appendage; staminate flowers short stalked, anthers 2;
pistillate flowers ellipsoid, perpendicular to the inflorescence-axis; appendage filiform, erect or bent, with linear
projections at the base (in female spadix) and baccate ones
at the tip.
Specimens examined. CHIAYI: Tsengwen reservoir,
Pinglin Yentzuyai, Wang 11008 (HAST). KAOHSIUNG:
Shanping, Summer collecting team s.n. 1986 (TNU),
Wang et al. 9552, 9687 (TNU), Chen 526 (HAST);
PINGTUNG: Kuwarusu, Matuda 447 (TAI).
Distribution. endemic to the low altitudinal mountainous region of southern part of Taiwan (Figure 6).
Notes. A. matsudai Hayata was originally described
based on an incomplete (lacking subterranean stem) male
plant (Type: Matsuda s.n. 1917, TI photo!). Most taxonomists (e.g. Kitamura, 1941; Huang, 1960; Liu & Huang,
1978) thought it synonymous with A. grapsospadix Hayata
because both taxa share many aspects such as trifoliolate
leaves, a greenish spathe with a semilunar white mottle
on the throat, and the filiform appendage. However, after
comparing numerous fresh, complete, and matured collections in detail, A. matsudai can be distinguished easily from A. grapsospadix in having a deciduous habit, a
subglobose subterranean stem, a unisexual spadix in the
mature plant, many linear sterile flowers born above the
fertile flowers in the female spadix, and pistillate flowers
perpendicular to the inflorescence-axis, etc.
A. matsudai is also similar to A. inclusum N. E. Brown
in Java, but may be distinguished by having baccate sterile flowers at the spadix-appendage tip. The species probably also resembles A. penicillatum N. E. Brown of
southern China, and A. laminatum Blume of Malay
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Figure 9. Arisaema grapsospadix Hayata. 1 & 2, Habit and rhizome; 3, Leaf; 4, Male spadix; 5, Female spadix; 6, Male flowers;
7, Female flower; 8, Longitudinal section of ovary showing ovules. (Wang et al. 9090)
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 10. Arisaema matsudai Hayata. 1 & 2, Habit and tuber; 3, Leaf; 4, Male spadix with spathe; 5, Female spadix; 6, Male
flowers; 7, Female flower; 8, Longitudinal section of ovary showing ovules. (Wang et al. 9552)
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Figure 11. Arisaema ilanense J. C. Wang sp. nov. 1, Habit; 2, Spadix with spathe; 3, Male spadix; 4, Female spadix; 5, Male
flowers; 6, Female flower; 7, Ovules. (Wang et al. 6705)
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Archipelago. An intensive study is necessary to clarify
their classification.
II. Arisaema sect. Clavata Engler, Pfl.-reich IV, 23F:
150 et 171. 1920; J. Murata, J. Fac. Sci. Univ. Tokyo
III 13:470. 1984.—TYPE: Arisaema clavatum Buchet.
3. Arisaema ilanense J. C. Wang, sp. nov. (Figure 11)
Species A. du-bois-reymondiae Engl. affinis, sed spatha
intus atro-purpurea, appendice spadicis claviformibus
sursum amplificatis 9-14 cm longa apice obstipa vel leviter
procurva differt. —TYPE: Taiwan, Ilan Hsien: Ssuyuan
to Nanshan, Wang et al. 6705 (holotype: TNU; isotypes:
HAST, TAI, TNU, TUS).
Tuber depressed-globose, 1.5-3 cm in diameter.
Pseudostem 10-30 cm long, deeply subterranean.
Cataphyllsthree or four, fleshy, whitish to greenish, apex
mucronate. Leaves two; petiole terete, 10-22 cm long;
lamina pedate, leaflets 7-15, sessile, oblanceolate to
elliptic, cuneate at base, acuminate at apex, the terminal
one 10-22 cm long, 2-4.5 cm wide, longer than or equal
to the adjacent one. Peduncle 1.5-7 cm long, usually much
shorter than the petiole. Spathe pale to greenish yellow
outside, dark purple inside; tube cylindrical, slightly auriculate at the mouth, 6-9 cm long, 1.3-2 cm in diameter,
pale to greenish yellow outside, dark purple inside and
gradually paled to reddish brown downward; blade ovate,
6-10 cm long, 3-5 cm wide, acute or sometimes acuminate to the apex. Spadix 1.5-2.4 cm long in male, 1.5-3.4
cm long in female, appendage clavate, 9-15 cm long,
slightly exerted, upper portion dark purple to black, bent
outward or slightly downward, lower portion reddish,
gradually narrowed toward the floriferous part or sometimes stipitate at the base with some echinate projections.
Specimens examined. ILAN: Ssuyuan to Nanshan,
Wang et al. 6705 (holotype TNU, isotypes HAST, TAI,
TNU, TUS); same loc., Wang et al. 8145 (TNU)
Distribution. endemic to northern Taiwan at medium
altitude from 1600 to 1900 m (Figure 6).
Notes. Arisaema sect. Clavata comprises seven species
distributed in Japan and China (Murata, 1984). The section is characterized by having pedate foliolate leaves with
many sessile leaflets and a sessile spadix-appendage with
modified sterile flowers (Murata, 1984). Among those
species belonging the section, A. ilanense closely resembles A. du-bois-reymondiae Engler in China and A.
heterocephalum Koidz. in the Ryukyu Islands, but A.
ilanense also somewhat resembles A. negishii Makino of
Japan and A. hunanense Hand.-Mazt. of mainland China.
The former two have an clavate or cylindrical spadix-appendages which are rather short and often included in the
spathe; while the latter two have an elongate spadix-appendage with a somewhat slender tip. A. ilanense has a
characteristic clavate spadix-appendage which is longer
t h a n t h o s e o f A . d u - b o i s - re y m o n d i a e a n d A .
heterocephalum, but is not as elongated as those of A.
negishii and A. hunanense (Ohashi, pers. comm.).
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A. ilanense was only found in Ssuyuan, northern
Taiwan. This region forms a saddle between the Central
Mountain Range and the Hsuehshan Mountain Range, and
is also the source of the Lanyanghsi River. The habitat is
cool and moist. Thousands of individuals grow on the forest floor under broad-leaved trees. A. taiwanense occurs
at the same locality, but their flowering season seems
different. A. ilanense flowers from February to late March
while A. taiwanense flowers from late March to May. The
two species are possibly isolated by the flowering season.
III. Arisaema sect. Tortuosa Engler, Pfl.-reich IV, 23F:
150 et 185. 1920; Nakai, Bot. Mag. Tokyo 43:524.
1929; Hara, Fl. East. Himalayas 2: 324 et 341. 1971;
Li in Wu & Li, Fl. Reipubl. Popularis Sin. 13(2): 156.
1979; J. Murata, J. Fac. Sci. Univ. Tokyo III 13:471.
1984.—TYPE: Arisaema tortuosum (Wall.) Schott.
IIIa Arisaema subsect. Flagellarisaema (Nakai) J.
Murata, J. Fac. Sci. Univ. Tokyo III 13:471. 1984.—
TYPE: Arisaema thunbergii Blume.
Flagellarisaema Nakai, J. Jap. Bot. 25:6. 1950.
4. Arisaema thunbergii ssp. autumnale J. C. Wang, J.
Murata & Ohashi, ssp. nov.
(Figure 12)
Arisaema thunbergii ssp. urashima auct. non (Hara)
Ohashi & J. Murata: J. C. Wang, Taiwania 37: 54, fig.
1. 1992.
Ex affinitate A. thunbergii ssp. urashima (Hara) Ohashi
et J. Murata, sed spatha intus et lamina extus alternatim
atropurpureo- et albo-striatus, autumno florens differt. —
TYPE: Taiwan, Taipei Hsien: Sanhsia, Hsiung-kung to
Man-yue-yuan, on the floor of broadleaf forest, Wang,
Sang & Chen 9460 (holotype: TNU; isotype: HAST,
TNU).
Tuber depressed-globose, 3-6 cm in diameter.
Pseudostem 5-15 cm long, much shorter than the petiole.
Cataphylls membranous, purplish to brownish, apex
mucronate. Leaf solitary; petiole terete, 20-55 cm long;
lamina pedate; leaflets 11-15, oblanceolate to elliptic, cuneate at base, acuminate at apex, the terminal one 10-28
cm long, 2-7.5 cm wide. Peduncle 10-20 cm long, usually shorter than petiole. Spathe white to pale yellow with
dark to bronze-purple or reddish purple stripes; tube
cylindrical, auriculate at the mouth, 5-7 cm long, 2-2.5
cm wide, white to pale yellow with longitudinal purple
stripes outside and dark purple stripes inside; blade triangular ovate, 7-10 cm long, 4-5 cm wide, acute to acuminate at apex. Spadix 3-4 cm long in male, appendage
filiform, 40-60 cm long, long-exerted, upper portion bent
downward and pendulous, lower portion gradually thickened to form a smooth long cylindrical enlargement then
gradually narrowed toward floriferous part.
Specimens examined. TAIPEI: Yun-sen Waterfall,
Wang et al. 9035 (TNU); Sanhsia, Hsiung-kung to Manyue-yuan, on the floor of broadleaf forest, Wang, Sang &
C. H. Chen 9460 (type, HAST, TNU); Province Road No.
7, Chihtuan to 67.5K, Wang 11504 (HAST). ILAN:
Fushan Botanical Garden, 1992, Yang s. n. (TNU); Nanao
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Figure 12. Arisaema thunbergii ssp. autumnale J. C. Wang, J. Murata & Ohashi ssp. nov.. 1 & 2, Habit and tuber; 3, Spadix with
spathe; 4, Apex of cataphyll; 5, Male spadix; 6, Male flower from the upper part of spadix, front view and side view; 7, Male
flower from the lower part of spadix, side and front view. (Wang 6747A)
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Hsiang: en route from Shenmihu control station to
Shenmihu, Liu et al. 147 (HAST, TNU). HUALIEN:
Lanshan, Peng 9922 (HAST); Taroko National Park, en
route from Yuehwangting to Yenhai Forest Trail, Peng et
al. 12990 (HAST); Walami, Lu & Horng 1361 (TNU);
Yenhai logging tract, Wang 6747A (TNU), Wang et al.
7640 (TNU).
Distribution. endemic to Taiwan, widespread but scarce
at low altitudes of northern and eastern parts (Figure 7).
Notes. The taxon was recently found in northern and
eastern Taiwan and previously misidentified as Arisaema
thunbergii ssp. urashima of Japan (Wang, 1992). The
gross morphology of the Taiwan plants is closely similar
with those from Japan. However, by a detailed comparison,
the Taiwanese taxon is different from that in Japan in the
pollen morphology, flowering season, and shape and color
of spathe. The palynological differences were described
as above. The flowering season was considered to be important by Murata (1984), who used it as one character
in discriminating the Arisaema section Decipientia.
However, the flowering time may not be so important in
the Taiwanese Arisaema because they flower one after
another all year long, probably due to the warmer climate.
However, the flowering season of A. thunbergii ssp.
autumnale is quite different from that of A. thunbergii ssp.
urashima, the former is exclusively in the autumn (from
October to November) while the latter is in the spring
(from April to May, Ohwi (1965)). Lastly, A. thunbergii
ssp. autumnale is distinct from A. thunbergii ssp.
urashima by the less expanded spathe-limb, and in having distinctly longitudinal striation on the both surfaces
of spathe-blade as well as the inner surface of spathe-tube.
Therefore, it is reasonable to treat them as distinct.
IIIb. Arisaema subsect. Tortuosa (Engler) J. Murata, J.
Fac. Sci. Univ. Tokyo III 13: 471. 1984.
5. Arisaema heterophyllum Blume, Rumphia 1: 110.
1835; Nakai, Icon. Pl. Asiae Orient. tab. 79. 1939;
Kitamura, Acta Phytotax. Geobot. 10: 189. 1941;
Huang, J. Forest NTU 27: 8. 1960; Liu & Huang,
Quart. J. Taiwan Mus. 16: 134. 1963; Liu & Huang in
H. L. Li et al., Fl. Taiwan 5: 805. 1978; Li in Wu &
Li, Fl. Reipubl. Popularis Sin. 13(2): 157, pl. 30: 1-3.
1979.
(Figure 13)
Heteroarisaema heterophyllum (Blume) Nakai, J. Jap. Bot.
25: 5. 1950.
Arisaema takeoi Hayata, Icon. Pl. Form., 5: 246. 1915;
8:132, fig. 59. 1919. —TYPE: Formosa. Inter Heirinbi
et Shokei, 10 May 1916, Hayata s.n. (neotype: TI, chosen by H. Ohashi, 1981: 23; figured in Ohashi, Univ.
Mus. Univ. Tokyo, Mater. Rep. 5, pl. 49. 1981)
Tuber depressed-globose, 1-6 cm in diameter.
Pseudostem 25-60 cm long, much longer than petiole. Leaf
solitary; petiole 5-15 cm long; lamina pedate; leaflets 1119, narrowly elliptic to oblanceolate, cuneate at base,
acuminate at apex, short-petiolate to sessile; terminal leaflet 4-15 cm long, 1-3.5 cm wide, much shorter than adja-
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cent leaflets. Peduncle 15-22 cm long, longer than the
petiole. Spathe green on both sides; tube cylindrical, not
or slightly auriculate at the mouth, 4-10 cm long 1-2 cm
in diameter; limb ovate or ovato-lanceolate, 5-11 cm long,
3-5 cm wide, acuminate at apex. Spadix mostly dioecious,
2.5-3.5 cm long in male, 4-5 cm long in female; appendage filiform, 15-25 cm long, erect, bent outward at tube
mouth and then turned upward and long-exerted, basal
part with few echinate sterile flower or not.
Specimens examined. KEELUNG: Dandan, 1944,
Masamune s. n. (TAI); TAIPEI: Agyokusan, 1934, Suzuki
s. n. (TAI); Chingtienkang to Tsuetsueku, Wang 5284
(TNU), 5332 (HAST, TNU); Chunghsing Farm, Lu 57
(TNU); Ertsuping, Wang et al. 9205 (TNU); Hsiaokoutou,
Huang & Jeng 10823 (TAI); Hsintien: en route from
Hualin to mt.-hiking entrance of Tatungshan, Liu et al.
204 (HAST); Hualin to Tatungshan, Liu et al. 204 (TNU);
Kungliao Hsiang: Chaohui Temple, near Yenliao, Peng
et al. 14965 (HAST); Peichatienshan, Lu 412 (TNU);
Tatungshan, Deng 928 (TNU); Tatunshan to Chingho,
Hsu 5561 (TAI); Shihkuliao, Yang 1253 (TNU); Shinshan,
Chiang 1561 (TAI); MIAOLI: Hsiaoping, Hsu 9322 (TAI);
Hsihu Hsiang Public Cemetery, Peng et al. 13953
(HAST). ILAN: Mt. Oobi, 1938, Masamune s. n. (TAI);
Nanao Hsiang: Chinyang Village, Shenmihu, Chen et al.
675 (HAST); Nan-ao to Hoping, Wang 6741 (TNU);
Tanoorei, Suzuki 7401 (TAI); Taipingshan, Suzuki 174
(TAI); Shenmeihu Huang et al. 5204 (TAI); HUALIEN:
Taroko National Park: Tsankuang Temple to a mt. ridge
above the destroyed Eternal Spring Temple, Peng et al.
12307 (HAST); Tenhsian, Huang 9319 (TAI); Yenhai logging tract, Wang 6747 (TNU); Yuehwangting to Yenhai
logging tract, Wang et al. 7670 (TNU).
Distribution. China, Korea, and Japan. Taiwan, in the
northern parts at low altitude (Figure 6).
IV. Arisaema sect. Sinarisaema Nakai, J. Jap. Bot. 25:
6. 1950; Hara, Fl. East. Himalayas 2:324 et 347. 1971;
Li in Wu & Li, Fl. Reipubl. Popularis Sin. 13(2): 186.
1979; J. Murata, J. Fac. Sci. Univ. Tokyo III 13: 475.
1984. —TYPE: Arisaema formosanum (Hayata)
Hayata.
6. Arisaema consanguineum Schott, Bonplandia 7: 27.
1859; N. E. Brown, J. Linn. Soc. Bot. 36: 176. 1903;
Hayata, Icon. Pl. Form. 5: 241. 1915; Huang, J. Forest
NTU 27: 6. 1960; Huang, Taiwania 7: 102. 1960; Liu
& Huang, Quart. J. Taiwan Mus. 16: 130. 1963; Liu
& Huang in H. L. Li et al., Fl. Taiwan 5: 801. 1978;
Hara, Fl. East. Himalayas 394. 1966; Hara, Fl. East.
Himalayas, ed. 2, 348, pl. 19: c, d, fig. 55: k. 1971.
Figure 14
Arisaema erubescens var. consanguineum (Schott) Engl.
in DC., Monogr. Phan. 2: 558. 1879.
Arisaema kelung-insularis Hayata, Icon. Pl. Form., 5: 246,
fig. 88. 1915; Liu & Huang in H. L. Li et al., Fl. Taiwan 5: 805. 1978; Li in Wu & Li, Fl. Reipubl. Popularis
Sin. 13(2):188. 1979. —TYPE: Formosa. Ins. Kelung,
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Figure 13. Arisaema heterophyllum Blume. 1 & 2, Habit and tuber; 3, Spadix with spathe; 4, Male spadix; 5, Male flowers; 6,
Female flower, upper one showing longitudinal section and inner ovules. (1993, Huang s.n.)
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 14. Arisaema consanguineum Schott. 1 & 2, Habit and tuber; 3, Leaflet; 4, Male spadix; 5, Female spadix; 6, Male flowers;
7, Female flower; 8, Longitudinal section of ovary showing inner ovules. (Chen & Hung 1111)
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May 1910, Sasaki s.n. (holotype: TI, figured in Ohashi,
Univ. Mus. Univ. Tokyo, Mater. Rep. 5, pl. 19. 1981).
Arisaema consanguineum var. kelung-insularis (Hayata)
Huang, Taiwania 7: 102, fig. 7. 1960; Liu & Huang,
Quart. J. Taiwan Mus. 16: 132. 1963.
Arisaema biradiatifoliatum Kitamura, Acta Phytotax.
Geobot. 10: 187. 1941; Li in Wu & Li, Fl. Reipubl.
Popularis Sin. 13(2): 187. 1979. —TYPE: Formosa.
Takao, Daikwanzan, 14 May 1939, Okamoto s.n.
(holotype: KYO? n. v.)
Arisaema erubescens auct. non Schott: Li in Wu & Li,
Fl. Reipubl. Popularis Sin. 13(2): 189. 1979.
Arisaema japonica auct. non Blume: N. E. Brown, J.
Linn. Soc. Bot. 36: 178. 1903; Matsumura & Hayata,
J. Coll. Sc. Univ. Tokyo 22: 456. 1906.
Tuber depressed globose, 2-7 cm in diameter.
Pseudostem 30-80 cm long, white to pale green, sometimes mottled with purple-brown marks. Cataphyllsthree
or four, greenish or reddish, sometimes with purple-brown
marks. Leaf solitary or two; petiole 15-40 cm long, distinctly shorter than the pseudostem; lamina radiately 915-foliolate; leaflets narrowly elliptic to elliptic, 12-25 cm
long, 1-6 cm wide, apex acuminate, base cuneate, green
above, glaucous beneath. Peduncle 6-12-(15)cm long, distinctly shorter than the petiole. Spathe green or red with
white lines along veins on the outer surface; tube
cylindrical, auriculate at the mouth, 5-8.5 cm long, 1-2
cm in diameter, greenish to yellowish green, sometimes
with purplish red stripes on inner surface; blade triangular ovate, 4-8 cm long, 2-4 cm wide, white, green or purplish red, often with pale veins on inner surface, long
acuminate, sometimes with a prolongated filiform tail at
apex. Spadix 6-10 cm long including appendage; appendage clavate, erect, 4-7 cm long, tip more than 2 mm thick,
base with fine bristly projections usually abundant in the
female spadix but scarce in the male; Infructescence recurved downward.
Specimens examined. TAIPEI: Chisingshan, Yang 1368
(TAI), Tang 1249 (TAI); Chunghsing Farm, Lu 53, 59
( T N U ) ; K a b o z a n , F u k u y a m a S T 1 9 2 4 1 ( TA I ) ;
Tsaoshankou, near Chinkuashih, Peng & Chen 10867
(HAST). TAOYUAN: Lalashan, 6th group 32 (TAI);
Parlin to Lalashan, Cheng 8 (TAI). MIAOLI: Kwanwu,
Huang 4751 (TAI); same loc., along the trail to the “Sacred Tree”, Peng et al. 14910 (HAST); 2 km of Kuanwu,
Peng et al. 14936 (HAST); Kuanwu to Chutung, Wang
et al. 9288 (HAST, TNU); Loshan to Kwanwu, Wang &
Yang 4721 (TNU). HSINCHU: Chienshih Hsiang:
Yuanyanghu Natural Preserved Area, Shen et al 739
(HAST), Wang & Hsu 1311 (HAST), Wang et al. 1055
(HAST); Wufeng Hsiang: Kuanwu, Talu forest road 2026K, Wang et al. 9218, 9221 (HAST, TNU); same loc.,
east branch line 7-10K, Wang et al. 9258 (HAST, TNU),
9258A (TNU). TAICHUNG: Anmashan, Huang 8070,
8116, 8120 (TAI); Chihyuanshanchuang, Tang 1251
(TAI); Nanhuhsi-Tochiatunshan-710 track road, Wang et
al. 3848 (TAI); Pilushi 11th watershed, Wang 3096 (TAI).
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NANTOU: Dandar logging tract, 1993, Yang s.n. (TNU);
en route from Tungpu Hot Spring to Kuankao, Peng 8024
(HAST); Kunyang to Yuanfeng, Wang 5093 (TAI);
Meifeng, 1993, Yang s.n. (TNU), Peng 8334 (HAST);
Hsinyi District: Shenmu Forest Trail, Peng & Chen 11403
(HAST); Tsuifeng to Meifeng, Tateishi et al. 17754 (TAI);
Yushan National Park: en route from Lulin Lodge to
Tatachia Saddle, Peng & Chen 11457 (HAST); Wushe to
Chienching, Feung & Kao 4540 (TAI). CHIAYI: Alishan,
Huang 1765 (TAI), 1991, Chang s.n. (TNU), Tateishi et
al. 17806 (TAI); Alishan, Erhwanping, Tateishi et al.
17920 (TAI); Alishan to Erhwanping, Wang 6378 (TNU);
En route to Yushan, Huang et al. 14221, 14231, 14242
(TAI); Tataka, Wang et al. 5483 (TNU). KAOHSIUNG:
Chinching Bridge, Wang 10397 (HAST); Taoyuan Hsiang:
Tienchih, Wang et al. 1152 (HAST), Wang 10393 (HAST);
Tashulinshan, 1919, Matsuda s.n. (TAIF). ILAN:
Taipingshan, Suzuki 187 (TAI), 1918, Sasaki s.n. (TAIF).
HUALIEN: Yenhai logging tract, Wang et al. 7668 (TNU);
Yuli Wildlife Nature Reserve, Deng 433-1, 433-3 (TNU).
TAITUNG: Haituan District: between mt.-hiking entrance
and the peak of Takuanshan, Peng et al. 11737 (HAST).
Distribution. Northern India, Nepal, Sikkin, Burma,
northern Thailand, and mainland China. Taiwan, from
low to medium altitudes throughout the island (Figure 7).
7. Arisaema formosanum (Hayata) Hayata, Icon. Pl.
Form. 5: 243, fig. 87. 1915 (“formosana”); Huang, J.
Forest NTU 27: 6, fig. 6, 7. 1960; Huang, Taiwania 7:
102, fig. 1, 3, 4. 1960; Liu & Huang, Quart. J. Taiwan
Mus. 16: 133. 1963; Liu & Huang in H. L. Li et al.,
Fl. Taiwan 5: 803, pl. 1527. 1978; Li in Wu & Li, Fl.
Reipubl. Popularis Sin. 13(2):187. 1979. (Figure 15)
Arisaema alienatum var. formosanum Hayata, J. Coll. Sci.
Univ. Tokyo 30:371. 1911 (“formosana”).—TYPE:
Formosa. Shinchiku, Okunghishan, Apr. 1907,
Kawakami & Watanabe s.n. (holotype: TI, figured in
Ohashi, Univ. Mus. Univ. Tokyo, Mater. Rep. 5, pl. 5.
1981).
Arisaema formosanum forma stenophyllum Hayata, Icon.
Pl. Form. 5: 24. 1915 (“formosana” form.
“stenophylla”); Huang, J. Forest NTU 27:8. 1960; Liu
& Huang, Quart. J. Taiwan Mus. 16: 133. 1963; Liu
& Huang in H. L. Li et al., Fl. Taiwan 5: 803. 1978;
Li in Wu & Li, Fl. Reipubl. Popularis Sin. 13(2):188.
1979. —TYPE: Formosa. Mt. Arisan, May 1913, S.
Sasaki 5 (holotype: TI, figured in Ohashi, Univ. Mus.
Univ. Tokyo, Mater. Rep. 5, pl. 13. 1981)..
Arisaema formosanum var. bicolorifolium Huang,
Ta i w a n i a 2 7 : 3 0 , f i g . 1 , p l . 1 - 5 . 1 9 8 2
(“bicolorifolia”).—TYPE: Taiwan. Mt. Tiger, Huang
8698 (holotype: TAI) syn. nov.
Arisaema oblanceolatum Kitamura, Acta Phytotax.
Geobot. 10: 188. 1941 (“oblanceolata”); Li in Wu &
Li, Fl. Reipubl. Popularis Sin. 13(2): 186. 1979. —
TYPE: Formosa. Takao, Keinanzan, 10 May 1939,
Okamoto s.n. (holotype: KYO. n. v.)
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 15. Arisaema formosanum (Hayata) Hayata. 1 & 2, Habit and tuber; 3, Leaflet; 4, Male spadix; 5, Female spadix; 6, Male
flowers; 7, Female flower. (Wang et al. 9220)
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82
Tuber depressed globose, 1.5-4.5 cm in diameter.
Pseudostem 10-60 cm long, white to pale green, sometimes mottled with purple-brown marks. Cataphyllsthree
or four, greenish or reddish, sometimes with purple-brown
marks. Leaf solitary, rare two; petiole 12-30 cm long,
about equal or shorter than the pseudostem; lamina radiately 7-11-(15)-foliolate; leaflets narrowly elliptic to
elliptic, 10-30 cm long, 0.8-7.5 cm wide, apex acuminate,
base cuneate, green above, glaucous beneath. Peduncle 410-(15)cm long, distinctly shorter than petiole. Spathe
green to reddish green, with white lines along veins, on
the outer surface; tube cylindrical, auriculate at the mouth,
4-6 cm long, 1-1.8 cm in diameter, greenish to yellowish
green, sometimes with purplish red stripes on inner
surface; blade ovate, 4-7 cm long, 2-3.5 cm wide, green
to purplish red, often with pale veins on inner surface,
long acuminate with a prolongated filiform tail at the
apex. Spadix 5-8 cm long including appendage; appendage filiform, (2)-3-4 cm long, less than 1.5 mm wide, base
with fine bristly projections usually abundant in the female spadix but scarce in the male; Infructescence recurved downward.
Specimens examined. TAIPEI: Chunghsing Farm, Lu
58 (TNU); Erhtzuping, Wang et al. 9206 (TNU);
Tsutzuhu, Nakamura 4354 (TAI); HSINCHU: Chienshih
Hsiang: Yuanyanghu Natural Preserved Area, Shen et al.
585 (HAST). MIAOLI: Hushan, Huang 8695 (TAI); Taian
Hsiang: Malapangshan, L i u e t a l . 9 3 (HAST);
TAICHUNG: Anmashan, Huang 8118, 8119 (TAI);
Pilushi 11th watershed, Wang 3095 (TAI); Shuanshan,
1983, Lin & Hsieh s.n. (TAIF); Wuling, en route from
Wuling Farm to Chikashanchuang, Hsu & Moore 633
(HAST). NANTOU: Chitou, Lu 603 (HAST, TNU), Wang
15362 (HAST), Chuang & Kao 3174 (HAST); Chitou
tract, Huang 1401, 1440 (TAI); Fenghuangshan, Peng &
Chen 10517 (HAST); Hoshie tract, Huang 2187 (TAI);
Hsinyi Hsiang: from Patungkuan to Tungpu, Chen 988
(HAST); Patungkuan, Huang 1588 (TNU); Tsuifeng, Hu
1550 (HAST); Tungpu to Kuankao, Huang 1589 (TNU);
Tunyuan to Yunhai, Liao et al. 1289 (TNU); CHIAYI:
Alishan, 1911, Kawakami & Sasaki s.n. (TAIF), Huang
1667, 1668, 1706 (TAI), Chann 5458 (TAI); Alishan to
Erhwanping, Wang et al. 6379 (TNU); Alishan to
Tzushan, Wang et al. 6428 (TNU); Chuchi Hsiang:
Shihcho, Lungyunshanchuang, Wang et al. 725 (HAST);
Erhwanping, 1914, Hayata s.n. (TAIF); Fenchihu, Huang
& Huang 13978 (TAI), Huang 2602 (TAI); Fenchihu to
Dardungshan, Wang et al. 7080, 7081 (TNU); Fenchihu
to Tianchian, Wang et al. 6251 (TNU); Monlu cliff to
Tungpu, Hsu 5389 (TAI); Mt. Morrison, Chuang et al.
4033 (HAST); Niitakayama, Kawakami & Mori 2330
(TAIF); KAOHSIUNG: Tengjy, Wang et al. 7482 (TNU);
Tienchu to Yiakou, Southern E-W Highway, Huang et al.
8859 (TAI). PINGTUNG: Peitawushan, Wang et al. 9166
(TNU); ILAN: Chiliehting Ridge, Huang 7793, 7852
(TAI); Nanao Hsiang: Chinyang Nanao broad-leaved forest preserve, Peng et al. 13796 (HAST); Chinyang: border of Shenmihu Lake in “Nanao broad-leaved forest
bot371-09.p65
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preserve”, Peng et al. 13814 (HAST); Taipingshan, Huang
et al. 10792, 10807 (TAI); Yuanyang Lake, Wang 9292
(TNU); HUALIEN: Chingsuishan, 1983, Chiou & Ho s.
n. (TAIF); Fuli Hsiang: Hsinkangshan, Shen et al. 650
(HAST); Mt. Tarokotaizan, Suzuki 10926 (TAI); Topoko,
1983, Chiou & Lin s.n. (TAIF); Taroko National Park:
Yenhai Forest Trail, Peng et al. 12367 (HAST); Yenhai
logging tract, Wang et al. 6794 (HAST, TNU), Wang et
al. 7667 (HAST, TNU); Yuehwangting to Yenhai logging
tract, Wang et al. 7639 (TNU); Yuli Wildlife Nature
Reserve, Deng 433-2 (TNU); TAITUNG: Sinsuiei, 1932,
Sasaki s.n. (TAI), Suzuki ST8027 (TAI); Southern cross
island highway 155K, Hsiangyang to Liyuan, Wang 14204
(HAST).
Distribution. Endemic to Taiwan, from low to medium
altitudes throughout the island (Figure 7).
Notes. A. consanguineum and A. formosanum are very
similar in morphology as well as habit. The two species
often occur in the same locality (see Figure 7). Their typical forms are easily distinguished by the thickness of the
spadix-appendage tip, the only character distinguishing
them. In general, the former has a thicker spadix-appendage (typically more than 3 mm in diameter) while the later
has a slender one (typically less than 1.5 mm in diameter).
However, both taxa show great variation in plant size, leaf
shape, spathe color, and the thickness of spadix-appendage tip, all of which result in some infraspecific taxa. Also,
some intermediate plants with 2-3 mm thick spadix-appendage tips were occasionally found. Murata (1985)
pointed out that these species may be isolated seasonally
and/or geographically. However, their flowering season
changes continuously from species to species. During the
transition period, hybridization, which may confuse the
delimitation of species, possibly occurs. In this situation,
any nomenclatural change and infraspecific classification
await an extensive comparative study of both species.
8. Arisaema taiwanense J. Murata, J. Jap. Bot. 60: 353,
fig. 1, 2, 3. 1985. —TYPE: Taiwan. Erhwanping, Apr.
19, 1985, Murata 17248 (holotype: TI, n.v.; isotype:
TAI).
(Figure 16)
Arisaema consanguineum auct. non Schott: Hayata, Icon.
Pl. Form. 5: 241. 1915.
Tuber depressed globose, 2-6 cm in diameter.
Pseudostem 2-30 cm long, white to pale green, usually
mottled with purple-brown marks. Cataphylls greenish or
reddish, often with purple-brown marks. Leaf solitary;
petiole 10-40 cm long, usually longer than the
pseudostem; lamina radiately 7-15-foliolate; leaflets
oblanceolate, apex filiform-caudate, with a thread-like
pendulous tail at tip, green above, glaucous beneath. Peduncle 5-15 cm long, distinctly shorter than the petiole.
Spathe purplish red to dark purple, with fine green dots
between veins on outer surface; tube obconical, gradually
opening to the auriculate mouth, 2-6 cm long, 4-8 cm
wide, white to pale yellowish green with purple stripes
on inner surface; blade ovate to broadly ovate, 3-8 cm
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 16. Arisaema taiwanense J. Murata. 1 & 2, Habit and tuber; 3, Leaflet; 4, Male spadix; 5, Female spadix; 6, Male flowers;
7, Female flower. (Wang et al. 9377)
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84
long, 4-8 cm wide, dark purple, scarcely with pale veins
on inner surface, acute to acuminate with a prolongated
filiform tail up to 40 cm long at apex. Spadix 3-10 cm
long including appendage; appendage stipitate or sometimes sessile, thick cylindrical, 2-5 cm long, 0.5-1.5 cm
wide, apex slightly enlarged, strongly to weakly rugose,
base narrower, with fine bristly projections, usually abundant in the female spadix but scarce in the male;
Infructescence recurved downward.
Specimens examined. TAOYUAN: Lalashan, 1990,
Chiang s.n. (TAI); HSINCHU: 8.7 km N of Kuanwu, Peng
et al. 14937 (HAST). TAICHUNG: Anmashan, Huang
8114, 8115 (TAI); Chihyuanshanchuang, Tang 1250
(TAI); Hoping Hsiang: Lishan, Ho 328 (HAST); same
loc., Hsu 3849 (TAI); Pilushi 11th watershed, Wang 3095
(TNU); NANTOU: Chiti, near Chitou, 1968, Hsu s.n.
(TAI); Juiyenchi, Lu et al. 1450 (TNU); CHIAYI: Alishan,
1914, Hayata s.n. (TAI), 1924, Ishizaki s.n. (TAI), Wang
2303 (TAI), 1957, Chuan & Kao s.n. (TAI); Alishan
highway, Shihcho, Tinghuti-Tadungshan-Furungshan,
Wang 18495 (HAST); Alishan to Erhwanping, Tateishi
et al. 17901 (TAI); Chihchung, Wang et al. 5406 (TNU);
Erhwanping, Murata 17248 (isotype of A. taiwanense J.
Murata, TAI!); Erhwanping to Shinmu, Murata & Huang
17667 (TAI); Fenchihu to Dardungshan, Wang et al. 7082
(HAST, TNU); KAOHSIUNG: Tengchih, Huang et al.
13849, 13850 (TAI); ILAN: Ssuyuan Yakou, Yang &
Hsieh 4070 (TAI); Ssuyuan to Nanshan, Lu 70 (HAST,
TNU), 71, 224 (TNU); Taipingshan (Taiheizan), Lin et
al. 23 (TNU), Suzuki 187 (TAI); Taipingshan, around
Taipingshanchuang, Lin et al. 23 (HAST).
Distribution. Endemic to Taiwan, throughout the island at medium altitudes, common (Figure 8).
8a. Arisaema taiwanense var. brevipedunculatum J.
Murata, J. Jap. Bot. 60:356, fig. 3A. 1985. —TYPE:
Taiwan. Peitawushan, Kuaikushanchuang - the base,
May 5 1985, Murata 17252 (holotype: TI. n.v.; isotype:
TAI).
Pseudostem 2-10 cm long, greenish, without colored
marks. Cataphylls greenish pale brown, without colored
marks. Peduncle 1-5 cm long. Spathe tube 2-4 cm long,
3.5-6 cm wide; spathe-blade concave and arched, 3-5 cm
long not including a thread-like tail. Spadix-appendage
2-4 cm long, 0.5-1.2 cm wide.
Specimens examined. PINGTUNG: Peitawushan,
Kuaikushanchuang - the base, Murata 17252 (isotype of
A. taiwanense var. brevipedunculata J. Murata, TAI!),
Tateshi et al. 19562, 19564 (TAI); HUALIEN: Yuli Wildlife Nature Reserve, Deng 444-1, 444-3 (TNU).
Distribution. Endemic to Taiwan, at medium altitudes
of southern and eastern parts (Figure 8).
V. Arisaema sect. Pedatisecta Schott ex Engler in DC.,
Monogr. Phan. 541. 1879 pro minor part.; J. Murata,
Kew Bull. 46(1): 127. 1991. Lectotype: Arisaema
japonicum Blume, chosen by J. Murata 1991: 127.
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9. Arisaema ringens (Thunb.) Schott, Melet. 17. 1832;
N. E. Brown, J. Linn. Soc. Bot. 36: 179. 1903; Hayata,
Icon. Pl. Form., 5: 246. 1915; 9: 147, fig. 54. 1920;
Huang, J. Forest NTU 27: 8, fig. 8. 1960; Huang,
Taiwania 7:101. 1960; Liu & Huang, Quart. J. Taiwan
Mus. 16: 134. 1963; Liu & Huang in H. L. Li et al.,
Fl. Taiwan 5: 805. 1978; Li in Wu & Li, Fl. Reipubl.
Popularis Sin. 13(2): 155, pl. 29:4-6. 1979; Chang, J.
Phytogeogr. Taxon. 32: 111. 1984.
(Figure 17)
Arum ringens Thunb., Act. Soc. Linn. Lond. 2: 337. 1794.
Ringentiarum ringens (Schott) Nakai, J. Jap. Bot. 25: 5.
1950.
Arisaema arisanense Hayata, Icon. Pl. Form. 6:100. 1916;
Huang, J. Forest NTU 27:6. 1960; Li in Wu & Li, Fl.
Reipubl. Popularis Sin. 13(2): 156. 1979. —TYPE:
Formosa. Arisan, 2500m alt., June 1914, Faurie s.n.
(holotype: TI, figured in Ohashi, Univ. Mus. Univ.
Tokyo, Mater. Rep. 5, pl. 8. 1981).
Arisaema taihokense Hosokawa, J. Jap. Bot. 12: 214, fig.
2, 3, 5: a-c. 1936; Li in Wu & Li, Fl. Reipubl. Popularis
Sin. 13(2): 155. 1979.—TYPE: Taiwan. Taihoku,
Koko, 1934, Ohnuma s.n. (holotype: TAI).
Tuber depressed globose, 2-5 cm in diameter.
Pseudostem 4-30 cm long, white to pale green or sometimes brownish pale green. Cataphylls greenish to
purplish. Leavestwo; petiole 10-35 cm long, usually longer
or equal to pseudostem; lamina trifoliolate; terminal leaflet
ovate-lanceolate to rhomboid-elliptic; lateral leaflets obliquely ovate to lanceolate, apex filiform-caudate, usually
with a thread-like tail at tip, green above, glaucous
beneath. Peduncle 3-6 cm long, distinctly shorter than
petiole. Spathe yellowish green to green; tube cylindrical,
gradually opening to an auriculate mouth, 3-5.5 cm long,
1-2 cm in diameter, yellowish green to green on outer
surface, purplish red with white to pale yellowish green
striations on inner surface; blade concave and revolute,
saccate, caudate at the apex. Spadix 3-8 cm long including appendage; appendage white, 2-6 cm long including
a short stalk of 3-10 mm long, attenuate, base truncate,
apex obtuse, without modified sterile flower.
Infructescence erect.
Specimens examined. KEELUNG: Campus of National
Taiwan Coll. Marine Science and Technology, Peng 6361
(HAST); Chingjenhu, Liao et al. 1116 (HAST);
Hopingtao, Lu 604 (TNU); Insl. Keelung, Masamune et
al. 42 (TAI); Keelung, 1913, Sasaki s.n. (TAIF), Kou &
Kao 4509 (TAI); TAIPEI: Chihnankung, Hoshanyueh,
Wang 15307 (HAST); Chihsingshan to Erhtzuping, Lin
et al. 78 (TNU); Chinshan, 1993, Wang s.n. (TNU), Lu
511, 512 (TNU); Chunghsing Farm Huang 13454, 13455
(TAI), Lu 55, 56, 61, 62, 68 (TNU); en route from Hsintien
to Pinglin, Leu & Yang 24 (HAST); Homei, Meiyienshan,
Huang & Huang 13974 (TAI); Hsiaokoutou, Huang &
Jeng 10822 (TAI), 1991, Chen et al. s.n. (TNU), Ou 2
(TNU); Hsiaokoutou to Shihting, Wang 5280 (TNU);
Huangtitian, Kao 388, 389, 390 (TNU), Wang et al. 700
(HAST); Koko, Ohnuma s.n. 1934 (Holotype of A.
taihokense Hosokawa, TAI!); Kuanyinshan, 1915,
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Wang — Systematics of Taiwanese Arisaema (Araceae)
Figure 17. Arisaema ringens (Thunb.) Schott. 1, Habit and tuber; 2, Male spadix; 3, Female spadix; 4, Male flowers; 5, Female
flower. (Wang 6455)
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Shimada s.n. (TAIF), 1931, Mori s.n. (TAI), Huang 2007,
2 0 1 5, 2 3 5 5 (TAI), Fukuyama s.n. 1940 ( TA I ) ;
Manyuehyuan, Wang 5675 (HAST, TNU); Maso (Wanli),
Kawakami & Shimada 4348 (TAIF); Mt. Sitisei, Suzuki
3503 (TAI), Suzuki 6747 (TAI); Nankang: en route from
Chunghua Inst. of Technology to Nakangshan, Peng &
Chan 10425 (HAST); Nanpanliao, Huang 10453 (TAI);
Pei-Highway, Huang 15930 (TAI); Peiyi highway
(Province Road No.9) 53.5K, Wang 15308, 15309, 15310
(HAST); Pichashan, Yang 1196 (TAI); Pihu, Tsou & Lin
652 (HAST); Pinglin Hsiang: Jenlipan, Peng et al. 10585
(HAST); Sabosan, 1914, Shimada s.n. (TAIF);
Sandiaoling, Masamune 2643 (TAI); Shamaoshan, Chung
705 (PPI); Sekitei, Kawakami & Fuji 1914 (TAIF);
Shantzuchiao, 1934, Sasaki s.n. (TAI); Shihting Hsiang:
Leikungpo, Hsiaokotou, Peng & Hu 12286 (HAST); Sozan
(Yangmingshan), 1929, Suzuki s.n. (TAI), Hsu 5303
(TAI), Chuang & Lin 4809 (HAST); Tatunshan, Lu 15216
(TAIF), Murata & Yang 17861 (TAI); Tsaoshan, Hsu
10403 (TAI); Wulai, Ruan 42 (HAST, TNU), 1972, Chang
s.n. (TNU), 1981, Chen s.n. (PPI); Yangmingshan National Park, Peng & Hsiao 10459 (HAST); Yangmingshan
National Park: on the way from Erhtzuping to
Hsiangtienchih, Chen et al. 431 (HAST, TNU);
Yangmingshan National Park: Tsutzuhu, Leu 387
(HAST); Yehliu, Wang et al. 7310 (TNU), Lu 599 (TNU);
Yinhotung, 1972, Chien s.n. (TNU), Wang 6455 (HAST,
TNU); TAOYUAN: Lalashan Chen 13 (TAI); HSINCHU:
Neiwan, 1916, Sasaki s.n. (TAIF). MIAOLI: Shihtan,
Peng et al. 13937 (HAST). ILAN: Insl. Kizan, 1932,
Masamune & Suzuki s.n. (TAI); Nanao Hsiang: Wushihpi,
Wang et al. 675 (HAST); Taipingshan, 1918, Sasaki s.n.
(TAIF); Taipingshan to Tuchang, Suzuki 1140 (TAI);
HUALIEN: Antong, Kao 6147 (TAI); Dangi-Forest,
Taroko, Suzuki 9253, 9305 (TAI);TAITUNG: Is. Lanyu,
1935, Hosokawa s.n. (TAI), Chang 6158, 11481, 11482
(PPI), Yeh 768 (PPI); Is. Lanyu, Hurngtourshan, Wang et
al. 6100 (TNU), Chung 572 (PPI).
Distribution. Mainland China, south Korea, Japan, and
the Ryukyu Islands. Taiwan, in northern part and Lanyu
Is. at low altitudes (Figure 8).
Acknowledgments. I wish to express my gratitude to Dr. H.
Ohashi, Professor of Tohoku University, and Dr. J. Murata, Tokyo Metropolitan University, for their critical reading of the
manuscript, valuable comments and assistance in sending me
useful literature, and also to Dr. D. H. Nicolson, Smithsonian
Institution, Dr. B. Bartholomew, California Academy of Science,
and Dr. C. I. Peng, Academia Sinica, for their critical reading
of the manuscript and helpful suggestions for improvement. I
am indebted to Mr. S. F. Huang, Herbarium of Department of
Botany, National Taiwan University, for providing the “Catalogue of the type specimens preserved in TI”, and also to Ms.
M. J. Deng and Y. C. Lu for their assistance in SEM
photography. Mr. K. C. Yang of the Taiwan Forestry Research
Institute, Dr. S. T. Chiu of the National Museum of Natural
Science, Ms. M. F. Kao of the Department of Botany, National
Taiwan University and Mr. W. H. Hu of the Institute of Botany,
Academia Sinicaall kindly sent me living materials. Thanks are
also due to the directors and curators of the herbaria for allow-
bot371-09.p65
86
ing me to examine the specimens. Last but not least, I am grateful to Dr. T. C. Huang, Professor of Department of Botany, National Taiwan University, for his encouragement. This work
was supported in part by grant NSC79-0211-B003-02 and
NSC83-0211-B003-007 from the National Science Council,
ROC.
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