ISSN 1346−7565

Acta Phytotax. Geobot. 59 (3): 194−210 (2008)

Taxonomic Revision of the Genus Entada (Leguminosae) in the
Ryukyu Islands, Japan
YOICHI TATEISHI1, NORIHISA WAKITA2 and TADASHI KAJITA2*
1

Department of Science Education, Faculty of Education, University of the Ryukyus, Okinawa 903-0129, Japan;
2
Department of Biology, Graduate School of Science, Chiba University, Chiba 263-8522, Japan.
*tkaji@faculty.chiba-u.jp (author for correspondence)

Two species of Entada are confirmed to occur in the Ryukyu Islands, Japan. One, E. phaseoloides,
occurs in tropical to subtropical Asia, on Pacific islands and in northeastern Australia. The second
species, E. tonkinensis, ranges from the Ryukyu Islands through northern Taiwan and southeastern
China to northern Vietnam. Descriptions and keys are provided for the two species, as well as for E.
rheedii and E. parvifolia which is sometimes confused with E. phaseoloides.
Keywords: Entada phaseoloides, Entada tonkinensis, leaflet number, Ryukyu Islands, seed morphology

Entada Adans. (Leguminosae; Mimosoideae)
comprises about 30 species of lianas and scandent
shrubs or subshrubs distributed in tropical andsubtropical regions (Nielsen 1981, 1992, ILDIS2005). Although more than half of the species occur only in Africa, three species have a very wide
distribution; Entada rheedii Spreng. from Africa
to the Indo-Pacific, E. gigas (L.) Fawc. & Rendle
in the New World, and E. phaseoloides (L.) Merr.
from Southeast Asia to the western Pacific region.
These species have seeds that are dispersed byocean currents, which likely explains their wide
distribution. The northernmost limit of distribution of Entada is in the Ryukyu Islands at thesouthwestern tip of the Japanese archipelago.
The first report of Entada from the Ryukyu
Islands was made by Ito & Matsumura (1899),
who identified a specimen from Ishigaki Island,
Yaeyama Islands as E. scandens Benth. Sakaguchi (1924) followed this treatment. Kanehira(1917), however, assigned these plants to his Entada formosana Kaneh., described from Taiwan,
and treated plants from Yakushima and AmamiOshima as the widespread E. phaseoloides.
Makino & Nemoto (1925, 1931) (Table 1) adopted Kanehira’s (1917) treatment. Later, Takamine(1952) recognized another species, Entada koshunensis Hayata & Kaneh., in the Yaeyama Islands. Sonohara et al. (1952) recognized Entada
formosana from the Yaeyama Islands (Ishigaki,
Iriomote and Yonaguni islands), and E. koshunensis from Ishigaki, Iriomote and Okinawa islands.
Sonohara (1952), moreover, treated plants of Entada on Yakushima and Amami-Oshima Islands as
E. phaseoloides. In addition to Entada koshunensis, Masamune (1955) recognized E. phaseoloides
from Okinawa Island.
Hatusima (1956) treated Entada koshunensis as a synonym of E. parvifolia Merr. from thePhilippines. His treatment (Hatusima 1956) wasNovember 2008

TATEISHI & AL.: Revision of Entada in the Ryukyus

followed by Hatusima & Amano (1958, 1967),
who also recognized Entada phaseoloides on
Okinawa Island. Hatusima (1971) later treated
Entada parvifolia as a synonym of E. phaseoloides. Walker (1976) also considered Entada
phaseoloides to be a widely distributed species

195with large morphological variation, and placed E.
koshunensis, E. parvifolia, and E. formosana in its
synonymy. Walker’s (1976) treatment was adopted in many floristic works and checklists on the
Ryukyu Islands (e.g., Hatusima & Amano 1977,
1994, Shimabuku 1990, 1997). Ohashi’s (1989)

TABLE 1. Summary of taxonomic history of Entada in the Ryukyu Islands.

ReferencesYear

Yaeyama

Ito & Matsumura

1899

E. scandens

Kanehira

1917

E. formosana

Sakaguchi

1924

E. scandens

Makino & Nemoto

1925

E. formosana

E. phaseoloides

Makino & Nemoto

1931

E. formosana

E. phaseoloides

Takamine

1952

E. formosana
E. koshunensis

Sonohara et al.

1952

E. koshunensis
E. formosana

Sonohara

1952

E. koshunensis
E. formosana

Masamune

1955

E. formosana
E. koshunensis

Hatusima & Amano

1958

E. parvifolia
E. phaseoloides

Hatusima & Amano

1967

E. parvifolia
E. phaseoloides

Hatusima

1971

E. phaseoloides

Walker1976

E. phaseoloides

Hatusima & Amano

1977

Ohashi1989

E. phaseoloides

Shimabuku

1990

E. phaseoloides

Hatusima & Amano

1994

Ohashi2001

E. rheedii

E. phaseoloides

Wakita et al.

2005

E. koshunensis

E. phaseoloides

E. phaseoloides

E. tonkinensis

Present study

Okinawa

Amami-Oshima

YakushimaE. phaseoloides

E. phaseoloides
E. phaseoloides

E. phaseoloides

E. phaseoloides

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treatment was fundamentally the same as Walker’s(1976), but different in suggesting the possible
occurrence of E. parvifolia on Ishigaki and Iriomote islands. Later, Ohashi (2001) recognized
the plants from Yaeyama and Okinawa islands as
E. rheedii.
In nearly all the treatments above, the species of Entada on Yakushima and Amami-Oshima(in the northern Ryukyu islands) has been treatedas E. phaseoloides. In contrast, the plants of Entada on Okinawa and southward have either been
considered to be E. phaseoloides or a different
species. The history of the treatment of Entada in
the Ryukyu Islands is summarized in Table 1.
Wakita et al. (2005) confirmed recently that
two species of Entada occurred in the Ryukyu
Islands. The first, recognized from Okinawa andYaeyama islands, has small, convex seeds withangular margins (type S seed) and leaves with thesecond (distal) pinnae consisting of more than
two pairs of leaflets. The second species, from
Yakushima and Amami-Oshima islands, has large,
compressed seeds with rounded margins (type Lseed) and leaves with the second pinnae consistingof two pairs of leaflets. SSCP analysis revealed
different DNA banding patterns in these species.
The molecular difference is a nucleotide substitution and some insertion/deletion differences in
the rps16 intron of the chloroplast DNA. Tentatively, they assigned the plants from Okinawa
and Yaeyama to Entada koshunensis, and plantsfrom Yakushima and Amami-Oshima to E. phaseoloides. Definitive taxonomic decisions were notmade because the sampling was limited only to
the Ryukyu Islands and no phylogenetic analysis
was conducted. Two species have recently been
confirmed by phylogenetic analyses of samples
from a wider distribution area using chloroplast
DNA sequences (Wakita et al. 2008), although
their taxonomy is still in question.
In this paper, we report the results of a taxonomic study on plants from the Ryukyu Islands,

Vol. 59

eastern and southeastern Asia, Oceania, and thePacific islands. Our aim was to clarify the taxonomy of the two species and determine theirgeographic distribution.

Material and Methods
Herbarium specimens preserved in BKF, BO,
HN, HNU, HYO, IBSC, KAG, KYO, L, MAK,
NH, NOU, NY, P, PDA, PNH, RYU, SING, TAI,
TAIF, TI, TNS, TUS, URO, US, and W (seesweetgum.nybg.org/ih/ for the list of herbarium
acronyms) were examined. Morphological characters, particularly of the leaf, pod, and seed, andgeographical distributions were studied.

Results and Discussion
In the survey of herbarium specimens, we
tentatively distinguished Entada phaseoloides on
the basis of the obliquely elliptic and distinctly
asymmetric leaves (Fig. 1, Table 2). We thenstudied a large numbers of specimens of plants
identified as Entada phaseoloides collected
throughout its range of distribution (including the
Ryukyu Islands) using seed shape as the primary
key character (see Appendix for a list of specimens examined). The specimens examined weredivided into two groups corresponding to the two
species recognized in the study by Wakita et al.
(2005). The geographical distributions of the twogroups were then compared (Table 2).
We found that the plants with smaller, convex seeds with angular margins (type S seed) occur over a broad swath of tropical to subtropical
Asia and the Pacific islands, while plants withlarge, compressed seeds with rounded margins(type L seed) occur in a smaller area from Vietnam to Japan (Fig. 2). The distribution of theformer species coincides with that of Entada phaseoloides (Nielsen 1992). Entada phaseoloides
was originally described as Lens phaseoloides

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TATEISHI & AL.: Revision of Entada in the Ryukyus

L. Merrill (1917) and Johnston (1949) chose as
the type the plate of Faba marina by Rumphiusin his Herbarium Amboinense 5: tab. 4 (1747).
The plate had probably been drawn from a plantgrowing in Ambon, or an adjacent area (Scheffer1871), which is within the range of distribution ofplants with type S seeds. Based on Rumphius’s
Faba marina, Scheffer (1871) described E.
rumphii, a name that has been treated as a synonym for E. phaseoloides. The two criticallydrawn plates attached to the original descriptionof Entada rumphii show a convex seed with alateral swelling (tab. 18 of Scheffer 1871), which
can be regarded as a type S seed. From the above
observations, we conclude that plants with type S
seeds should be treated as Entada phaseoloides.
We also conclude that plants with type L seedshould be assigned to Entada tonkinensis Gagnep.
Entada tonkinensis was originally described from
northern Vietnam (Gagnepain 1911), but has beentreated as a synonym of E. phaseoloides (Nielsen
1980, 1981, Lock & Heald 1994), and has neverbeen recorded outside Vietnam. According to theoriginal description (Gagnepain 1911), the seedsof E. tonkinensis are 6 cm long, 5 cm wide, andcompressed. These features are consistent with
type L seeds. We also confirmed that the typespecimen (Balansa 2130) of Entada tonkinensis at
P is identical with specimens from the Ryukyus,
Taiwan, and China. The monophyly of thesesamples has also been confirmed by phylogenetic
analyses (Wakita et al. 2008). The redefined diagnostic characters of Entada tonkinensis and its
distribution are given in the following key andtaxonomic treatment (Table 2 and Fig. 1).
Wakita et al. (2005), assigned plants fromthe Ryukyu Islands with type L seeds to Entada
phaseoloides, and plants with type S seeds to E.
koshunensis. They followed the treatment of Ho(1985) for Entada in Taiwan, which was based onseed characters and leaflet numbers. In contrast
with Wakita et al. (2005), we concluded that the197

plants with type S seeds are Entada phaseoloides,
and those with type L seeds are E. tonkinensis.
Although Wakita et al. (2005) recognized Entada
koshunensis, we considered it to be a synonym ofE. phaseoloides. In works recognizing Entada
koshunensis as a separate species, the diagnostic
character was given as the 3 or 4 pairs of leaflets(Hayata 1921, Makino & Nemoto 1925, 1931,
Nemoto 1936 as E. parvifolia, Ho 1985, Huang &Ohashi 1993). Our herbarium study revealed that
3 or 4 pairs of leaflets appear predominantly inplants from the Philippines through southern Taiwan to the southern Ryukyu Islands, and sporadically over the entire area of distribution of Entada
phaseoloides (see Appendix). Plants with 3 pairsof leaflets were also reported from the Solomon(Verdcourt 1979) and Bismarck islands (Nielsen
1992). Other features of Entada koshunensis,
for example the distinctly asymmetrical leaflets(Fig 1b) and pods with rather thin parchment-like
endocarps, are indistinguishable from E. phaseoloides.
Two other species, Entada parvifolia andE. rheedii, have sometimes been confused with
E. phaseoloides in the Ryukyu Islands (Table 1).
Entada parvifolia Merr., from the Philippines,
has sometimes been applied to plants of E. phaseoloides from the southern Ryukyu Islands andsouthern Taiwan (Hatusima 1956, Hatusima &Amano 1958, 1967, Ho 1985, Huang & Ohashi
1993). The type specimen (Ramos 5067, NY.
http://sciweb.nybg.org/science2/VirtualHerbarium.asp) and the original description of Entada
parvifolia (Merrill 1908), however, show that itis difficult to confuse E. parvifolia with E. phaseoloides. As summarized in Table 2 and in thekey, Entada parvifolia is not a gigantic liana, butrather a trailing or scandent shrub whose leaves
have 8–11 pairs of small, almost symmetricalleaflets (Fig. 1), small pods 15–30 cm long, and
4–5.5 cm wide, a thin parchment-like endocarp,
and small seeds 1.8–2 × 1.6–1.9 cm. Entada parv-

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ifolia is placed in a different subsection, subsect.
Sphaerospermae Brenan, from subsection Entada
in which E. phaseoloides belongs (Brenan 1967,
Nielsen 1992).
Early in the history of taxonomy on Entada
in the Ryukyu Islands, the name E. formosana, asynonym of E. rheedii, was sometimes applied toplants on Yaeyama (Table 1). In the most recent
comprehensive flora of Japan, Ohashi (2001) rec-Vol. 59

ognized Entada rheedii in the southern Ryukyu
Islands and treated E. phaseoloides as its synonym. These two species, however, are clearly different in shape, size, and number of leaflets (Fig.
1), texture of the pod endocarp, and thickness ofseeds (Table 2).
In this study, we show that two Entada species occurred in the Ryukyu Islands of Japan are
E. phaseoloides and E. tonkinensis. In the recent

TABLE 2. Comparison of Entada phaseoloides, E. tonkinensis, E. rheedii, and E. parvifolia.
CharacterE. phaseoloides

Habit

giant woody climber giant woody climber giant woody climber trailing or scandent
shrub

LeafletPod

SeedDistribution

E. tonkinensis

E. rheedii

E. parvifolia

outlineobliquely oblong to
obliquely elliptic or
obliquely obovately
elliptic, distinctly
asymmetricalobliquely elliptic or
obliquely obovately
elliptic, distinctly
asymmetricalobovate to elliptic, or oblong or obliquely
lanceolate, usually oblong, nearly
nearly symmetrical symmetrical

apex

acute to obtuse

acute to roundedobtuse to roundedrounded, truncate orslightly emarginate

no. per 2nd (1–)2–4 pairs
pinna

2–3(–4) pairs

4–5(–6) pairs

8–11 pairs

size of the
distal oneson 2ndpinna

7–11 cm long,
3.5–5.5 cm wide

8.5–12 cm long,
4.4–6 cm wide

6–9 cm long,
2.5–4 cm wide

1.5–3 cm long,
0.6–1.1 cm wide

endocarp

parchment-likeparchment-likewoody

thinly parchment-likelength50–120 cm long

40–130 cm long

40–120 cm long

15–30 cm longwidth

6–9.5 cm wide

9–12 cm wide

8–11 cm wide

4–5.5 cm wide

article

4–7.5 cm long

6–9.5 cm long

5–8.5 cm long

2.5–4 cm long

color

brown

blackish purple

brown

brown–dark brownshapesuborbicular, convex, suborbicular,
margin angularcompressed, marginroundedsize

3.5–5.5 × 3.3–4.5 cm 5.2–7.4 × 4.7–5.5 cm 3.5–5.7 × 3.5–5 cm

1.8–2 × 1.6–1.9 cm

thickness

1–1.5 cm

0.8–1.1 cm

1.6–2.3 cm

Southeast Asia to the Vietnam, China,
South PacificTaiwan, Japan

suborbicular,
irregularly ovoid
compressed, marginrounded2–2.5 cm

Africa, Indo-Pacific Philippines
region, northern
Australia

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TATEISHI & AL.: Revision of Entada in the Ryukyus

Red List of Japan only Entada phaseoloides islisted in the Ryukyu Islands (Kagoshima andOkinawa) as a category IB (EN) plant. Our study
suggests the single species on the Red List in-

199

cludes two different taxa, Entada phaseoloides
and E. tonkinensis, which should be consideredseparately for further conservation.

Key to the species in the Ryukyu Islands, including Entada parvifolia and E. rheedii
A1. Trailing or scandent shrubs; second (distal) pinna with 8–11 pairs of leaflets; distal leaflets of the second
pinna 1.5–3 cm long, 0.6–1.1 cm wide; pods 15–30 cm long, 4–5.5 cm wide; seeds irregularly ovoid, 1.8–
2 × 1.6–1.9 cm ………………………………………………… E. parvifolia (known only from the Philippines)
A2. Large climbing vine; second (distal) pinna with less than 6 pairs of leaflets; distal leaflets of the second
pinna more than 6 cm long, and more than 2.5 cm wide; pods more than 40 cm long, more than 6 cm
wide; seeds compressed laterally, more than 3.5 × 3.5 cm ………………………………………………… B
B1. Leaflets of second pinna 4 or 5(or 6) pairs; distal leaflets 6–9 cm long, 2.5–4 cm wide, usually nearly
symmetrical (equal sided), apex obtuse to rounded; pods with hard woody endocarp, 8–11 cm wide; seeds
more than 2 cm thick ……………………………………… E. rheedii (Africa, Indo-Pacific region, Australia)
B2. Leaflet of second pinna less than 4 pairs; distal leaflets more than 7 cm long, more than 3.5 cm wide, distinctly asymmetrical (unequal sided), apex acute to obtuse; pods with parchment-like endocarp ………… C
C1. Pods 6–9.5 cm wide; seeds 3.5–5.5 cm long, 3.3–4.5 cm wide, convex, 1–1.5 cm thick, brown, margin
angular …………………………………………………………………………………………… 1. E. phaseoloides
C2. Pods 9–12 cm wide; seeds 5.2–7.4 cm long, 4.7–5.5 cm wide, compressed, 1.6–2.3 cm thick, blackish
purple, margin rounded ……………………………………………………………………… 2. E. tonkinensis

Taxonomic treatment
1. Entada phaseoloides (L.) Merr.
Entada phaseoloides (L.) Merr. in Philip. J. Sci. 9: 86(1914); Merr., Interpret. Rumph. Herb. Amb. 253(1917); Merr., Sp. Blancoanae 168, 194 (1918);
Nemoto, Fl. Jap. Suppl. 379 (1936), p.p.; Brenanin Kew Bull. 1955 (2): 164 (1955); Masamune
in Sci. Rep. Kanazawa Univ. 3(1): 120 (1955),
p. min. p.; Hatusima & Amano, Fl. Okinawa: 45(1958); Hatusima & Amano, Fl. Okinawa (2nded.): 49 (1967); Hatusima, Fl. Ryukyus: 345 &
844 (1971), p.p.; Hatusima, Fl. Ryukyus (rev. ed.):345 & 844 (1975), p.p.; Walker, Fl. Okinawa: 541(1976), p.p.; Hatusima & Amano, Fl. Ryukyus:
61 (1977), p.p.; Huang & Ohashi in Fl. Taiwan 3:279 (1977), p. min. p.; I. Nielsen in Adansonia,
ser. 2, 19(3a): 342 (1980); Ohashi in Satake etal., Wild Flow. Jap. Woody Pl. 1: 233 (1989), p.p.
incl. pl. 254: 1–2; Shimabuku, Vasc. Fl. Ryukyu
Isls.: 217 (1990), p. p.; I. Nielsen in Fl. Malesiana,
ser. I, 11(1): 179 (1992); Huang & Ohashi in Fl.
Taiwan (2nd ed.) 3: 169, pl. 77 (1993); Hatusima& Amano, Fl. Ryukyus (2nd ed.): 88 (1994), p.p.;
Shimabuku, Vasc. Fl. Ryukyu Isls. (rev. ed.): 250(1997), p.p. –Lens phaseoloides L. in Stickman
Herb. Amboin. 18 (1754); Amoen. Acad. 4: 128(1759). —Lectotype: Rumphius, Herb. Amboin.
5: 5-8, tab. 4 (1747), (fide Johnston 1949).
Mimosa scandens L., Sp. Pl. ed. 2: 1501 (1763). –E. scandens (L.) Benth. in Hook., J. Bot. 4: 332(1841); Ito & Matsumura in J. Sci. Coll. Imp.
Univ. Tokyo, Sect. 1, 12: 442 (1899); Sakaguchi,
Gen. Index Fl. Okinawa 54 (1924). —Lectotype:
Rumphius, Herb. Amboin. 5: 5-8, tab. 4 (1747),(fide Johnston 1949).
E. koshunensis Kaneh. & Hayata in Hayata, Icon. Pl.
Formosan. 10: 3, fig. 1 (1921); Makino & Nemoto, Fl. Jap.: 722 (1925); Makino & Nemoto, Fl.
Jap. (rev. ed.): 568 (1931); Kaneh., Formos. Tree(ed. 2): 296 (1936); Sonohara et al., Fl. Okinawa:
74 (1952); Sonohara, Useful Trees Shrubs Ryukyus: 32 (1952); Takamine, Fl. Yaeyama Gunto: 48(1952); Masamune in Sci. Rep. Kanazawa Univ.
3(1): 120 (1955); Liu, Ill. Lign. Pl. Taiwan 1: 495(1960). —Type: Taiwan, Pingtung Co.: Chiopeng, alt. ca. 50 m, Dec. 1919, Kanehira 13205(TAIF).
E. formosana auct. non Kaneh. & Hayata: Kaneh., Formos. Tree 195 (1917), p.p.; Makino & Nemoto,200

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Vol. 59

FIG. 1. Outline of leaves of Entada. a, b: E. phaseoloides (a: Papua New Guinea, Carr 16315 (SING). b: Ishigaki Island, theRyukyu Islands, Nakamoto et al. 31 (URO). c: E. tonkinensis from Guangxi, China, Akiyama et al. 1244 (TI). d: E. rheedii
from Taiwan, Tateishi & Hoshi 21620 (URO). e: E. parvifolia from Luzon, Philippines, Ramos & Edano BS44583 (PNH).
Scale bar: 5 cm.

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TATEISHI & AL.: Revision of Entada in the Ryukyus

Fl. Jap.: 722 (1925); Makino & Nemoto, Fl. Jap.
(rev. ed.): 568 (1931); Nemoto, Fl. Jap. suppl.: 379(1936); Sonohara et al., Fl. Okinawa: 74 (1952);
Sonohara, Useful Trees Shrubs Ryukyus: 33(1952). Takamine, Fl. Yaeyama Gunto: 48 (1952);
Masamune in Sci. Rep. Kanazawa Univ. 3(1):3(1): 120 (1955).
E. parvifolia auct. non Merr.: Hatusima in Sci. Bull.
Agric. Home Econ. Div. Univ. Ryukyus 3: 20(1956); Hatusima & Amano, Fl. Okinawa: 46(1958); Hatusima & Amano, Fl. Okinawa (2nded.): 49 (1967); Ho in J. Taiwan Museum 38(1):76, photos 4 & 5, fig. 1C (1985); Huang & Ohashi
in Fl. Taiwan (2nd ed.) 3: 169 (1993).
E. rheedii auct. non Spreng.: Liu et al., Man. Taiwan
Vasc. Pl.: 76 (2000), p.p.; Ohashi in Fl. Jap. 2b:
217 (2001).

Climbers, woody. Stems to 40 m long, often flattened and spiraling, base to 3 m in diam. Leaves
alternate, petiolate, bipinnate; petiole 1.5–3.5 cm
long including basal pulvinus, glabrous or puberulous; rachis, midrib and margins of leaflets glabrous or puberulous; rachis 2–5 cm long, ending
in a bifid tendril; pinnae usually 2 pairs but 1 pairon juvenile plants, 6–20 cm long; leaflets (pinnules) opposite, (1 or)2 or 3 pairs on first (basal)
pinna and (1 or)2–4 pairs (3 or 4 pairs on Ryukyu
plants) on second (distal) pinna, increasing in size
upward, rigidly chartaceous to coriaceous, adaxially shiny, drying considerably darker than abaxial surface), obliquely oblong to obliquely elliptic
or obliquely obovate-elliptic, asymmetrical, baseunequal-sided, apex acute to obtuse and more orless retuse, entire, 4–11 cm long, 2–5.5 cm wide
(4–9.5 cm long, 2–4.5 cm wide in Ryukyu plants),
distal pinnules second pinna 7–11 cm long, 3.5–
5.5 cm wide (7–9.5 cm, and 3.5–4.5 cm wide in
Ryukyu plants). Inflorescences spikes, solitary, inleaf axils or sometimes several spikes from shortshoots, densely flowered, 12–30 cm long including peduncle; peduncle glabrous or puberulous,
1–6 cm long, 1.2–1.5 cm across; rachis rather
densely puberulous. Bracts lanceolate, 0.6–1 mm
long, abaxially puberulous, persistent. Flowers

201

sessile or subsessile, pentamerous, distylous;
flowers with short styles usually on lower half of
spike, flowers with long styles on upper half of
spike. Calyx green, broadly cup shaped, glabrous,
1.2–1.9 mm long; teeth inconspicuous, ciliate.
Petals 5, valvate, green with reddish base, oblongor lanceolate, apex acute, 2.8–4 mm long, 1.1–1.8
mm wide. Stamens 10, filiform, 5.5–8.2 mm long,white, turning yellow after anther dehiscence;
anthers 0.6–1.1 mm long, with a deciduous apical gland. Pistil stipitate; style 0.1–0.2 mm longin short styled flowers, ovary 0.7–1.1 mm long;ovules several, less than 0.1 mm long; style 2.7–4.8 mm long in long styled flowers, ovary 1.4–2.1 mm long, ovules 12–18, 0.1–0.2 mm long.
Pods compressed, straight to slightly curved,
sometimes slightly twisted, 0.4–1.2 m long, 6–10
cm wide, more or less constricted between seeds;
exocarp falling at maturity, endocarp parchmentlike, pale brown to pale grayish brown, splitting
into 1-seeded, turgid segments; segments 6.5–7.5
cm long; sutures woody, thick, continuous, forming a hoop from which the enclosed segments
readily detach at maturity. Seeds 9–16, reddish
brown, suborbicular, compressed, convex, margin
angular, 3.5–5.5 by 3.3–4.5 cm, 1–1.5 cm thick;hilum 0.5–3 mm long; with air filled cavity between cotyledons. Germination hypogeous; cotyledons remaining in seed coat; seedling leafless,first several nodes with only cataphylls.
Japanese name. Koshun-modama (Himemodama).
Distribution. Japan (Ryukyu Islands), Taiwan(Pingtung Co.), Philippines, Malaysia (Sarawak),Indonesia, Papua New Guinea, Australia (E. coastof Queensland), Northern Mariana Islands, Palau,
Micronesia (Yap, Truk, Pohnpei [Ponape] and Kusai Islands), Solomon Islands, Kiribati, Vanuatu,
New Caledonia, Fiji, Tonga, Samoa and French
Polynesia (Fig. 2).
Ryukyu Islands: Okinawa and Yaeyama islands (Ishigaki, Kohama, Iriomote and Yonaguni

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islands). Forests bordering littoral swamps andmangrove and riverine vegetation. Flowering in
June on Okinawa, April to August in Yaeyama Islands.
Entada phaseoloides is widely distributed
in tropical and subtropical areas of Asia and thePacific. Examination of many specimens of thegenus from Asia and the Pacific shows that E.
phaseoloides is not in southeastern continental
Asia, but extends throughout the islands of thebiogeographic province of Malesia, through the
Pacific and along the eastern coast of Queensland(Fig. 2).
2. Entada tonkinensis Gagnep.
Entada tonkinensis Gagnep. in Lecomte, Not. Syst.
2: 60 (1911); Gagnep. in Fl. Indo-Chine 2(2): 65(1913). —Type: Vietnam. Banton village, Tu-vu,
in 1888, Balansa 2130 (P, holo- and iso-).
E. phaseoloides auct. non (L.) Merr.: Kaneh., Formos.
Tree: 193 (1917); Makino & Nemoto, Fl. Jap.: 723(1925); Makino & Nemoto, Fl. Jap. (rev. ed.): 568(1931); Masamune, Fl. Yakushima 241 (1934);
Kaneh., Formos. Tree (rev. ed.): 296 (1936); Sonohara, Useful Trees Shrubs Ryukyus: 32 (1952);
Ohwi, Fl. Jap.: 672 (1953); Masamune in Sci. Rep.
Kanazawa Univ. 3(1): 120 (1955), p.p.; Hatusima& Amano, Fl. Okinawa: 46 (1958); Ohwi, Fl. Jap.
(English ed.): 553 (1965); Hatusima & Amano,
Fl. Okinawa (2nd. ed.): 49 (1967); Hatusima, Fl.
Ryukyus: 345 (1971), p.p.; Kitamura & Murata,
Colour. Ill. Wood. Pl. Jap. 1: 362, pl. 72-442 &fig. 236-2 (1971); Ohwi, Fl. Jap. (rev. ed.): 782(1972); Ohwi, Fl. Jap. (new ed.): 782 (1975); Hatusima, Fl. Ryukyus (rev. ed.): 345 (1975), p.p.;
Walker, Fl. Okinawa 541 (1976), p.p.; Hatusima& Amano, Fl. Ryukyus: 61 (1977), p.p.; Huang &Ohashi in Fl. Taiwan 3: 279, pl. 584 (1977), p.p.;
Hatusima, Fl. Kagoshima: 69 (1978); I. Nielsen
in Adansonia, ser. 2, 19(3): 342 (1980); I. Nielsen
in Fl. Cambodge, Laos & Vietnam. 19: 21, pl. 3-1(1981), p.p.; Ho in J. Taiwan Museum 38(1): 75,
photo 8, fig. 1A (1985); Hatusima, Fl. Kagoshima(rev. ed.): 86 (1986); Wu in Fl. Reipubl. Popularis Sin. 39: 13 (1988), p.p., excl. pl. 5; Ohashi inVol. 59

Satake et al., Wild Flow. Jap. Woody Pl. 1: 233,
(1989), p. p., excl. pl. 254-1–2; Shimabuku, Vasc.
Fl. Ryukyu Isls.: 217 (1990), p.p.; Hatusima, Fl. N.
Ryukyus: 103 (1991); I. Nielsen in Fl. Males. Ser.
I, 11(1): 179 (1992), p.p.; Huang & Ohashi in Fl.
Taiwan (2nd ed.) 3: 169, pl. 77 (1993); Hatusima& Amano, Fl. Ryukyus (2nd ed.): 88 (1994), p.p.;
Shimabuku, Vasc. Fl. Ryukyu Isls. (rev. ed.): 250(1997), p.p.; Ohashi in Fl. Jap. 2b: 217 (2001).
E. scandens auct. non (L.) Benth.: Matsum. in Bot.
Mag. Tokyo 16: 102 (1902), p.p.; Matsum. & Hayata in J. Coll. Sci. Imp. Univ. Tokyo, Sect. 1, 22:
116 (1906); Matsum., Index Pl. 2(2): 261 (1912);
Tutcher in Rep. Bot. For. Dept. Hongk. 1914: 30(1915).
E. parvifolia auct. non Merr.: Hatusima & Amano, Fl.
Okinawa: 46 (1958), p.p.; Hatusima & Amano, Fl.
Okinawa (2nd ed.): 49 (1967), p.p.

Climbers, woody. Stems often flattened and spiraling, base to 60 cm across. Leaves alternate,petiolate, bipinnate, glabrous; petiole 1.5–4 cmlong including basal pulvinus; rachis 3–6.5 cm
long, ending in a bifid tendril; pinnae usually
2 pairs, but 1 pair on juvenile plants, 10-22 cm
long; leaflets (pinnules) opposite, 2 pairs on first(basal) pinna, 2 or 3 pairs on second (distal) pinna, increasing in size upward, rigidly chartaceous,adaxially shiny, sometimes drying considerably
darker than abaxial surface, obliquely elliptic, or
obliquely obovate-elliptic, asymmetrical, baseunequal sided, apex acute to obtuse, entire, 5–
12 cm long, 2.5–6 cm wide, distal pinnules on
second pinna 8.5–12 cm long, 4.4–6 cm wide.
Inflorescences spikes, solitary in leaf axils orseveral spikes from short shoots, densely flowered, 9–25 cm long including glabrous peduncle;
rachis puberulous. Peduncle 1–7 cm long, 1.3–
1.5 cm across. Bracts lanceolate, 0.6–1 mm long,
abaxially puberulous, persistent. Flowers sessile
or subsessile, ca. 6 mm long, pentamerous, distylous; short styled flowers usually on lower half of
spike; long styled flowers on upper half of spike.
Calyx green, broadly cup shaped, glabrous, 1.2–
2 mm long; teeth inconspicuous. Petals 5, valvate,

105

120

135

150

165

180

0

165

1000

150

2000
3000 km

30

15

0

15

30

TATEISHI & AL.: Revision of Entada in the Ryukyus

FIG. 2. Distribution map of Entada phaseoloides ( ● from herbarium specimens; ○ from literature) and E. tonkinensis ( ▲ ).

90

E. phaseoloides
E. tonkinensis

November 2008
203

204

Acta Phytotax. Geobot.

pale green with reddish base, elliptic to oblong,apex acute, 3–3.2 mm long, 1.3–1.5 mm wide.
Stamens 10, filiform, 5.5–7 mm long, white, turning yellow after anther dehiscence; anthers 0.5–1
mm long, apically with a deciduous gland. Pistil
stipitate; style of short styled flowers 0.1–0.6 mm
long, ovary 0.5–1 mm long; ovules several, lessthan 0.1 mm long; style of long styled flowers 4–4.7 mm long, ovary 1.8–2 mm; ovules 12–18,
0.1–0.2 mm long. Pods compressed, straight toslightly curved, 0.5–1.5 m long, 9–12 cm wide,
exocarp falling at maturity; endocarp parchmentlike, pale brown, splitting into 1-seeded, turgid
segments 6.5–7.5 cm long; sutures woody, thick,continuous and forming a hoop from which enclosed segments readily detach at maturity. Seeds
9-16, blackish purple, compressed, suborbicular,
margin rounded, 5.2–7.4 by 4.7–5.5 cm, 1.6–2.3
cm thick. Germination hypogeous, cotyledonsremaining within seed coat; seedling leafless, first
several nodes with only cataphylls.
Japanese name. Modama.
Distribution. Japan (Ryukyu Islands), Taiwan (northern and central counties), China (HongKong, Fujian, Guangdong, Guangxi, Yunnan),
North Vietnam (Fig.2).
Ryukyu Islands: Yakushima and AmamiOshima. Riverine vegetation. Flowering in June.
In the Ryukyu Islands, Entada tonkinensis
has always been treated as E. phaseoloides (Table1), but can be clearly distinguished by seed shapeand size (Table 2). The type specimen (Balansa
2130) of E. tonkinensis at P has leaves with twopairs of obliquely elliptic and distinctly asymmetrical leaflets, but has no seeds. According tothe original description (Gagnepain 1911), theseeds of E. tonkinensis are compressed, but not at
all convex.
Entada tonkinensis ranges from the northern
Ryukyu Islands through northern and central Taiwan and southeastern China to northern Vietnam.

Vol. 59

Entada tonkinensis grows in inland evergreen forests, especially along streams at low and medium
elevations.
We thank Drs L.K.C. Chan, K. Chayamarit, N. Fukuoka, H. Funakoshi, K. Iwatsuki, K. Kurima, S. Matsu­
mura, C. Nyomdham, N. N. Thin, M. Tuda, K. Watana­be, T. Yamashiro, Irawati and J. Yip and T. Yukawa
for their help in obtaining samples. We also thank the
curators of the herbaria BKF, BO, HN, HNU, HYO,IBSC, KAG, KYO, L, MAK, NH, NOU, NY, P, PDA,
PNH, RYU, SING, TAI, TAIF, TNS, TUS, US and W
for permission to examine specimens. We also thankDrs J. Murata, T. Ohi-Toma, T. Denda, Y. Watano, T.
Asakawa, and K. Takayama for their valuable comments and useful suggestions during this study. Great
thanks are also due to Dr. D. Boufford for critical reading the manuscript. This work was supported by JSPS
KAKENHI Nos. 12575011 and 14405015 to Y.T. and16370043 and 19370032 to T.K.

References
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Bull. 10: 161–192.
Brenan, J. P. M. 1967. Notes on Mimosoideae: XI. The
genus Entada, its subdivisions and a key to theAfrican species. Kew Bull. 20(3): 361–378.
Gagnepain, F. 1911. Mimosees nouvelles. Notul. Syst.
(Paris) 2: 56–61.
Hatusima, S. 1956. New and noteworthy plants fromthe Ryukyu Islands and Formosa. Sci. Bull. Agric.
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Taiwan Museum. 38(1): 75–80.
Huang, T.-C. & H. Ohashi. 1977. Leguminosae. In:
Huang, T.-C. et al. (eds.), Fl. Taiwan 3: 148–421.
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Huang, T.-C. et al. (eds.), Fl. Taiwan (2nd ed.) 3:
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Service). 2005. ILDIS World Database of Legumes. version 10, November 2005. http://www.
ildis.org
Ito, T. & J. Matsumura. 1899. Tentamen Florae Lutchuensis, Sectio 1. J. Sci. Coll. Imp. Univ. Tokyo 12:
263–541.
Johnston, I. M. 1949. The botany of San Jose Island(Gulf of Panama). Sargentia no. 8. Arnold Arboretum of Harvard Univ., Jamaica Plain.
Kanehira, R. 1917. Formosan Tree. Bureau of Productive Industries, Government of Formosa, Taipei (inJapanese).
Lock, J. M. & J. Heald. 1994. Legumes of Indo-China.
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Makino, T. & K. Nemoto. 1925. Flora of Japan. Nipponshokubutsusoran Publishing Association,
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Makino, T. & K. Nemoto. 1931. Flora of Japan, 2nd ed.
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Nielsen, I. C. 1992. Mimosaceae. Fl. Males., Ser. 1,

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Flowers of Japan, Woody Plants I, pp. 229–256.
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Rumphius, G. E. 1747. Herbarium Amboinense vol. 5.
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Scheffer, R. H. C. C. 1871. Obserbationes Phytographicae. Natuurw. Tijdschr. Ned.-Indie 32: 387–427.
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Ryukyu Islands. Hirugi Sha, Naha. (in Japanese)
Shimabuku, K. 1997. Check List Vascular Flora of the
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Sonohara, S., S. Tawata, T. Amano & E. H. Walker.
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Ryukyu For. Exp. Sta. no. 1. (in Japanese)
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Kajita. 2008. Two species of Entada in Japan as
evidenced by cpDNA phylogeny. Acta Phytotax.
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Washington D.C.

Received November 26, 2007; accepted April 19, 2008

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APPENDIX. Specimens examined. Number following herbarium acronym shows number of largest pair of leaflets on second pinnae; letters indicate seed type (S: type S; L: type L).

Entada phaseoloides (L.) Merr.
JAPAN. Ryukyu Islands. No precise locality: Aug. 1909, Miyagi s.n. (MAK)3; fr, Hatusima s.n. (RYU). —Okinawa I.: Kunigami, 27 Jan. 1924, Tashiro s.n. (KYO, TAIF)4; Kunigami-son, Ibu, Ibu river, 5 Apr. 1974, fl.bud,
Shinzato s.n. (RYU)3; ibid., May 1984, fr, Murata s.n. (URO); ibid., 11 Nov. 1984, fr.s, Murata 17088 (TI, TUS,
URO) 3/S; ibid., 25 Apr. 1995, Tateishi et al. 40529 (URO)4; ibid., 29 Apr. 1995, fr, Nakamoto 8 (URO); ibid., 27
May 1995, fl, Nakamoto & Nakamoto 25 (URO)4; ibid., 28 May 1995, fl, Kanna 1 (URO); ibid., 27 Jun. 1995, fl,Nakamoto & Ogawa 2 (URO)3; ibid., 27 Jun. 1995, fl, Nakamoto 7 (URO)4; ibid., Sep. 1995, fr, Yamashiro s.n.
(URO); ibid., alt. 60–80 m, 30 Nov. 1995, Nakamoto 26 (URO)4; ibid., 26 Dec. 1995, Kurima et al. 126 (URO);
ibid., 5 Feb. 1997, Kurima & Tateishi 242 (URO)3-4/L; ibid., 3 Jun. 1997, Kurima & Matsumura 117 (URO)3-4;
ibid., 6 Jun. 1998, Kurima & Matsumura 118 (URO)3; ibid., 26 May 2001, Wakita 210526-1 (TI, URO)3-4/S;
Onna-dake, 27 Feb. 1935, Tawada 348 (KYO)4. —Ishigaki I.: No precise locality, fr.s, Kuroiwa s.n. (TI)S; ibid.,
11 Aug. 1894, fl, Ito 1280 (TI), 25–31 Jul. 1923, fl, Koidzumi s.n. (KYO)3, in 1923, Sakaguchi s.n. (KYO)3; Suriyama, 19 Mar. 1895, fl, Ito 1394 (TI); Yuriyama, 6 Aug. 1919, Kawagoe s.n. (KAG); Ota, 11 Nov. 2002, Wakita& Tateishi 231111 (URO)3; Upper stream of Miyara river, above Maesato reservoir, 2 Mar. 1995, fr, Murata
s.n. (URO)S, 18 Mar. 1995, fr.s, Tateishi et al. 40542 (URO)3/S; Maesato reservoir, alt. 40–50 m, 4 Jan. 1996,
Nakamoto et al. 29 (URO)3; ibid., 8 Jul. 1997, fl, Kurima & Matsumura 116 (URO)3; ibid., 30 Jul. 1997, fl, Kurima & Matsumura 105 (TI, URO)3; Mt. Fukai-omoto, 3–6 Jul. 1974, Miyagi et al. 37811 (RYU)3; Omoto, 8 Dec.
1996, Kurima & Matsumura 234 (URO)3; Sokohara, 14 Jun. 2002, Wakita 210614 (TI, URO)3; ibid., 16 Jul. 2003,
Wakita & Tateishi 230716 (URO)3; upper stream of Nagura river, 7 Dec. 1996, Kurima & Matsumura 233 (URO)3;
ibid., 2 Aug. 1997, fl, Kurima & Matsumura 108, (TI, URO)3; ibid., 3 Aug.1998, Kurima & Matsumura 101 (URO)3;
Nagura, 28 Jul. 1973, fl, Furuse 3671 (RYU, TUS)3; Nagura, 22 Jul. 2001, Wakita & Matsumura 210722b (URO)3;
upper stream of Miyara river, 2 Aug. 1923, fl, Tawada 347 (KYO)3; east of Takada, upper stream of Miyara
river, 23 Feb. 1988, seedl & fr, Murata & Murata 15580 (TI)3; Motonagura, 22 Jul. 2001, Wakita & Matsumura
210722C (URO)3; Kawahara, alt. ca. 20 m, 16 Sep. 1994, fr, Tateishi et al. 40278 (URO)4, & 40281 (URO)S;
ibid., 14 Jan. 1996, Nakamoto et al. 30 (URO)3-4; ibid., 8 Jul. 1997, fl, Kurima & Matsumura 112 (URO)4; Bannadake, 30 Jul. 1959, Oka 13172 & 13184 (KAG), 27 Mar. 1974, fr, Furuse 5441 (RYU); ibid., 18 Mar. 1995, Tateishi et al. 40543 (URO)3; ibid., 4 Jan. 1996, Nakamoto et al. 31 (URO)3; ibid., 7 Dec. 1996, Kurima & Matsumura
235 (URO)3; ibid., 8 Jul. 1997, fl, Kurima & Matsumura 111 (URO)3; ibid., 30 Jul. 1997, fl, Kurima & Matsumura
104 (TI, URO)3; Arakawa, seaside, 8 Aug. 1973, fr, Kawakami s.n. (RYU)S; Kabira–Yoshihara, 21 Aug. 1961,
fl, Migo 25830 (RYU)3; Uganzaki 30 Jul. 1997, fl, Kurima & Matsumura 113 (TI, URO)3; ibid., 3 Aug. 1997, fl,Kurima & Matsumura 107, (TI, URO)3-4; ibid., 22 Jul. 2001, Wakita & Matsumura 210722 (URO). —Kohama I.:
E. part, sandy beach, 4 Mar. 1997, Yamashiro 3034 (URO)3; ibid., 26 Mar. 1998, Shinjo & Tateishi 10950 (URO)3;
ibid., 13 Jun. 2002, Wakita 220613 (URO)3. —Iriomote I.: No precise locality, 1–20 Jul. 1923, fl, Koidzumi s.n.
(KYO)3, 28 Aug. 1966, fl, Miyagi 3645 (RYU)3, 17 Aug. 1967, Miyagi 3696 (RYU)3, Apr. 1973, yfr, Wada s.n.
(TI); Funaura, around the Tropical Biological Research Center, Ryukyu Univ., 123°48´00–10˝E, 24°23´30˝N, alt.
30–40 m, 16 Apr. 1998, fl, Yonekura et al. 98073 (TUS)3; Haemi-dake, May 1937, fl, Sonohara s.n. (KYO, TI)3;
Kuira-gawa River, 123°50´47.5˝E, 24°23´54.5˝N, alt. 20 m, 21 Jul. 2003, fl, Kokubugata 1571 (TNS)2; Nakama
river, 11 Aug. 1969, fr, Tokeshi 33 (RYU)3, 16 Jun. 1971, s, Yamazaki s.n. (TI)S, alt.0–30 m, in mangrove swamp,
29 Apr. 1982, fr, Fukuoka 11349 (KYO)3; mouth of the Yutsun river, alt. 10 m, 5 Jan. 1996, Nakamoto et al. 32(URO)3; Yutsun river, 4 Dec. 1996, Kurima & Matsumura 124 (URO)S; ibid., 21 Jul. 2001, Wakita 210721, (URO);
Yoshikera, 14 Aug. 1998, Kurima & Matsumura 128 (URO)3-4; Kura, 14 Aug. 1998, Kurima & Matsumura 141(URO)3; Inda-zaki, 12 Jun. 2002, Wakita 210612 (URO)3. —Yonaguni I.: 11–13 Jul. 1923, Koidzumi s.n. (KYO)4;
4–13 Jul. 1967, Shinjo 5313 (URO); Urabu, 24 Jan. 1993, Miyara s.n. (URO)4; Higawa, alt. 70 m, 6 Jan. 1996, fr.s,
Nakamoto et al. 33 (URO)3/S; ibid., 10 Jun. 2002, Wakita 210610 (TI, URO)3; ibid., 9 Nov. 2002, Wakita & Tateishi 231109 (URO)S.
TAIWAN. Pingtung Co.: Koshifo, Aug. 1981, Hsu s.n. (TUS)S; Chiopeng, alt. ca. 50 m, Dec. 1918, fr.s,
Kanehira 13205 (TAIF, holotype of E. koshunensis)S; ibid., alt. ca. 50 m, 1 Sep. 1984, Tateishi et al. 18446

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TATEISHI & AL.: Revision of Entada in the Ryukyus

207(TAI, TUS)3; Chiopeng, Chiopeng river, 22°06´29.9˝N, 120°52´55.5˝E, alt. 30–50 m, 22 Sep. 2003, Wakita
240922-1(URO)3/S, ibid., 22°06´32.0˝N, 120°52´51.9˝E, alt. 30–50 m, 22 Sep. 2003, Wakita 240922-2 (URO)3/S,
ibid., 22°06´44.1˝N, 120°52´46.2˝E, alt. 30–50 m, 22 Sep. 2003, Wakita 240922-3 (URO)4/S; Nanjen-shan, 22°05´
68.7˝N, 120°53´47.7˝E, alt. 20–50 m, 23 Sep. 2003, Wakita 240923 (URO)3-4; Nanjen-shan–Chiopeng, alt. ca. 200
m, 2 Nov. 1982, Ohashi et al. 14406 (TUS), 14410 (TUS, URO)3, fr, 14449 (TUS) & fr,14557 (TAI, TUS)3.
PHILIPPINES. Luzon I.: Rizal Prov., Mar. 1905, Ahern’s collector FB2901, fl (BO, P, SING)3; Tayabas
Prov., Casiguran, May-Jun. 1925, yfr, Ramos & Edano BS45227 (IBSC, SING)3; Quezon Prov., Ilavac, alt. 1025 ft,
14 Jul. 1961, Lagrimas & Rojo SH-323 (PNH)3; Sorsogon Prov., Irosin, Mt. Bulusan, Nov. 1915, fr, Elmer 15305(KAG, P, W)3, Jul. 1916, fl, Elmer 14723 (KAG, W)3; Isabella Prov., San Mariano, Feb.–Mar. 1926, Ramos &Edano BS47055 (IBSC, P)3; Camarines Prov., Albay, Jun. 1908, fl, Curran FB12256 (P)3. —Polillo I.: Karlagan,alt. 30 m, 19 Nov. 1948, yfr, Fox 21 (PNH)3. —Mindoro I.: Apr.–May 1908, Merritt FB41423 (L)3. —Samar I.:
Mt. Sohoton Natural Bridge National Park, Basey, 31 Jan. 1992, fr.s, Barbon et al. PPI-5782 (PNH)3/S. Leyte I.:
Hubasan, VISCA, Baybay, alt. 731 m, 1 Mar. 1993, yfr, Gaerlan et al. PPI-10648 (PNH)3. —Palawan I.: Quezon,
Ampaplot, 1 Oct. 1964, fr.s, Espiritu PNH 91467 (PNH)2. —Mindanao I.: Davao Distr., Todaya, Mt. Apo, Aug.
1909, Elmer (BO); Zamboanga del Norte, Dilsus, 31 Jan. 1958, fl, Frake 710 (PNH)3; ibid., alt. 500 m, 12 Feb.
1958, fl, Frake 773 (PNH); Bukidnon Prov., Mt. Katanglad, alt. 1700 m, 11 Apr. 1949, fl, Sulit 3458 (PNH)3. —Sulu Prov., Tawitawi I.: Jul.–Aug. 1924, fl, Ramos & Edano BS44221 (BO, P, SING)2-3.
PALAU. Kayangel I.: 28 Aug. 1939, drifted seed, Tuyama s.n. (TI)S. —Palau I.: Airai, inner edge of mangrove area, alt. 10 m, 4 Nov. 1969, fl, Fischer 4 (L)2. —Korror I.: Armijt, 10 Aug. 1939, fl, Tuyama s.n. (TI)2;
Arumizu, 10 Aug. 1939, fr, Tuyama s.n. (TI)S; Campus of Tropical Biology Institute, 13 Sep. 1939, Tuyama s.n.
(TI)2.
NORTHERN MARIANA ISLANDS. Saipan: Angaur I., seedl, Tuyama s.n. (TI).
MICRONESIA. Pohnpei (Ponape): Jul.–Aug. 1929, Kanehira 750 (TI)2.
MALAYSIA. Sarawak: Kuching Div., Bako National Park, Dec. 1956, Murthy S.7678 (SING)2, 17 Jul. 1966,
fl, Hou 533 (L)2; ibid., Telok Asam, 31 May 1963, fl, Ashton S.17814 (L, SING)2-3; ibid. 8 Aug. 1979, fl, Chai
S.41005 (L)2; ibid. 29 Mar. 1975, fl, Paie S.36418 (L)2.
INDONESIA. Sumatra. 14 Nov. 1920, Lorzing 7871 (BO). —Mentawai Is.: Siberoet I., 11 Sep. 1924,
yfr, Iboet 65 (BO, L, SING), 9 Sep. 1924, yfr, Boden-Kloss 10597 (BO, SING). —Krakatau I.: 20 Jul. 1924, van
Leeuwen 7985 (BO)3. Java. 24 Nov. 1914, yfr, Backer 17467 (BO). —Jawa Barat: Kediri, Prigi, 22 Feb. 1914,
fr, Backer 11773 (BO, L)2/S; Banten Prov., Udjung Kulon Nature Reserve, 27 Jan. 1984, yfr, Wirawan s.n. (L)2.
Sulawesi. Sulawesi Tengah: east slope of Benteng bay, Togean, 0°21´S, 122°00´E, alt. 50 m, 21 Dec. 1999, fl, Sidiyasa 1839 (BO, L)2; Sopu valley, c. 80 km SSE of Palu, c. 1°16´S, 120°18´E, alt. 1000 m, 26 May 1979, fl, vanBalgooy et al. 5556 (BO, KYO, L)3; Kampong Kadolo, in 1913, fl, Rachmat 729 (BO, L)2; Wuasa, Lowe Utara
I. 1°25´18.3˝S, 120°18´48.8˝E, alt. 1100 m, 28 Mar. 2001, fl, Kessler et al. PK3096 (L)2. —Sulawesi Tenggara:
Kendari, G.Atolanu, Sampara, Kab.Unaha, alt. 80 m, 6 Feb. 1986, fr, Amir 64 (BO)2/S. Nusa Tenggara Timur.
Lesser Sunda Is. Flores I.: 28 Jun. 1974, yfr, Verheijen 4317 (L)3; Manggarai, alt. 700 m, 16 Apr. 1936, fl, de
Voogd 2838 (BO, L)2; S. part, Mt. Ndebi, alt. 500 m, 14 Apr. 1965, Kostermans & Wirawan 230 (L)2. —Sumba
I.: Maumaru–Kenanggar, alt. 750 m, 25 Jan. 1975, yfr, Wiriadinata 479 (BO)2. Maluku (Moluccas). Halmahera
I.: fl, de Vriese s.n. (L)2; Galela, alt. ± 20 m, 20 Sep. 1921, fr, Beguin 1739 (BO, L)2/S. —Ternate I.: Toramadiahi,alt. ± 400 m, 20 Dec. 1920, fl, Beguin 1241 (L)2, & 1242 (BO)2. —Tanimbar Is.: Yamdena I.: 12 Mar. 1938, yfr,
Buwalda 4127 (BO, L)2, near Ranarmoje River, near Norkese, 5 Apr. 1956, fl, van Borssum 3284 (BO, L)2; Selaru I.,
13 Mar. 1956, yfr, Pleyte 77 (BO). —Boeroe I., Kajeli, Teysmann 1871 (BO); Wa’Katin, alt. 600 m, 21 Apr. 1921,
Toxopeus 158 (BO). —Ceram I.: 19 Nov. 1993, fr, Suharno IRA61 (BO)2/S; Sungei Mala, Uweth, Taniwel, alt. 10
m , 15 Dec. 1984, fl, Ramlanto 491 (BO, L)2-3; Desa Gah, Kec. Seram Timur, alt. 3 m, 17 Feb. 1986, Mirmanto& Ruskandi ERI.101 (BO)2. —Ambon I.: Teysmann s.n. (BO), de Vriese & Teysmann s.n. (L)2, Tochehoc, 11 Jul.
1900, Boerlage 202 (BO)2; Benteng, 24 Oct. 1931, s, Rant 528 (BO)S. —Wetar I. Masapun: S. coast, alt. 2 m,
13 Apr. 1939, fl, Bloembergen 3713 (PNH, SING)2-3. —Aru Islands: 19 Apr. 1993, Nooteboom 5688 (L)2; Trangan I., W. of Sia, 6°49´S, 114°16´E , 5 Oct. 1994, fl, van Balgooy 6649 (L)3. Papua (Irian Jaya). Andai, south of
Manokwari, 31 Mar. 1943, fl, Tuyama 997 (TI)2; Merauke Distr., Agats, Kec. Sawaerma, Ds. Sawa, alt. 10 m, 23

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Oct. 1992, fl, Widjaja et al. EW6219 (L)2; Vogelkop, Manokuwari subdistr. Warnapi, 15 km north of Ransiki, alt. ±20 m, 27 Sep. 1948, Kostermans 2804 (BO, L, SING)2; W. New Guinea, Servei, 26 Sep. 1939, yfr, van Dijk 998a
(BO, L)2; Mamberamo region, Idenburg River, alt. 125 m, 1 Sep. 1914, fl, Feuilletau de Bruyn 79 (BO)2; Wandarmon Pen., Wondiwoi Moutains, alt. 800 m, 28 Feb. 1962, fl, Schran 13311 (KYO, L)3; Div. South N. Guinea, along
river Maro, between Merauke and Tajam, 12 Jul. 1957, fr.s, Kalkman BW3732 (BO, L)2/S; South New Guinea, Sg.
Aoendoena near Oeta, alt. 3 m, 9 Jul. 1941, fl, Aet 492 (BO, L)2; Kp. Tajam, 29 Jul. 1941, fl, Anta 204 (L)2.
PAPUA NEW GUINEA. Northern Div., ca. 1/2 km inland of Cape Killerton, 9 Jul. 1953, yfr, Hoogland 3260
(L)2. East Sepik Prov., Hunstcin range (Mt. Sansan), 4°28´S, 142°43´E, alt. 100–200 m, 5 Aug. 1990, fl, Takeuchi
6458 (L)4. —Sepik Distr.: Aitape Subdistr., near Wantipi village, along Bliri River, alt. ca. 650 ft, 29 Jul. 1961, fl,Darbyshire & Hoogland 8315L (PNH, SING)4. —Eastern Papua, 500 ft, Jan. 1941, s, Shaw Mayer s.n. (SING)S.
—Lower Fly River, E. bank opp. Sturt I., Oct. 1936, fl, Brass 8082 (BO, L)2. —Western Distr.: Kiunga Subdistr.,
Base Camp, Ok Tedi River, 5°14´S, 141°12´E, alt. 2500 ft, 2 Nov. 1969, fl & yfr, Foreman & Galore NGF 45772(L)2; Oriomo Creek, south of Yakup Cr., 8°50´S, 143°00´E, alt. 50 ft, 26 Sep. 1963, Womersley NGF 17763 (L)2;
Oriomo River, 8°50´S, 143°15´E, alt. 70 ft, Jan. 1959, White & Gray NGF 10444 (L)2. —Morobe Distr. Bupu
River, 6°45´S, 147°00´E, alt. c. 100 ft, 29 Sep. 1960, fl, Henty NGF 13620 (L, SING)2, ibid., 5 Sep. 1963, fl, Henty
NGF 16699 (L)2; N. coast of Huon Gulf at Singaua Plantation, 147°09´E, 06°40´S, 30 Oct. 1962, fl, Hartley 10866(L)2; Lae Subdistr., Singawa, 20 miles from Lae, 6 45´S, 147 00´E, alt. 10 m, 4 Apr. 1974, fr, Katik NGF 37993(L)2; Lae Subdistr., Apo beach, 6°44´S, 147°00´E, alt. sea level, 29 Sep. 1978, fl, Kerenga et al. LAE 74302 (L)2;
Labu, 6°45´S, 146°55´E, alt. 10 ft, 13 Dec. 1965, fl, Streimann & Kairo NGF 26084 (L)2. —New Britain: Kandrian Subdistr., Piriloma Village, 6°06´S, 150°45´E, alt. c. 1300 ft , 11 Mar. 1965, fl & fr, Sayers NGF 21913 (L)3;
E. New Britain Distr., Pomio Subdistr., 5°04´S, 151°48´E, alt. 900 m, 9 Jun. 1973, fl, Stevens & Lelean LAE 58669(L, PNH)2. —Sudest Island: Jac Landing, 16 Aug. 1956, fl & yfr, Brass 27715 (L)2. —Kokoda, ca. 1200 ft, 30
Mar. 1936, fl, Carr 16315 (SING)2, ca. 2000 ft, 28 Apr. 1936, fl, Carr 16474 (PNH, SING)2. —Bougainville I.:
Karngu, Buin, sea level, 17 Oct. 1930, fr.s, Kajewski 2265 (BO)2; Lower south slope of Lake Loloru Water, c. 14
miles north of Buin, alt. c. 2300 ft, 13 Aug. 1964, fl, Craven & Schodde 259 (PNH)2; Siwai, Dec. 1932, fl, Waterhouse 821-B (L); Maisua, 18 Sep. 1931, Waterhouse 548-B (L).
SOLOMON ISLANDS. Shortland Is.: South East Oema I., 2 Apr. 1969, fl, Mauriasi et al. BSIP 13807 (SING)2.
—New Georgia Is.: Bambari area, east Kolombangara I., ridge top 240´ above sea level, 3 Jan. 1968, fl, Mauriasi
et al. BSIP 8583 (L, SING)3; Vangunu I., Gevala River, 12 Dec. 1962, fl, Whitmore BSIP 1246 (L); Vangunu I.,
Balavaeni, 10´ above sea level, 19 Jul. 1965, fl, Maenu’u BSIP 6123 (L, SING)3; S.E. New Georgia, Mango River,10´ above sea level, 22 Jun. 1965, fl, Maenu’u BSIP 6056 (SING)2; Totepari I., east of Maoesango Pt., ridge top
650´ above sea level, 23 Jul. 1969, fl, Mauriasi et al. BSIP 16121 (L)2. —Santa Ysabel I.: Allardyce Harbour, alt.
50 ft, behind the mangrove, 16 Oct. 1963, fl, Whitmore BSIP 2210 (L)2. N.W. Santa Cruz, Graciosa Bay (Kauri
Area), ridge top 400´ above sea level, 11 Nov. 1969, fl, Mauriasi et al. BSIP 17209 (L, SING)3. —Malaita I.: S.W.
Malaita, Are Are Distr., west coast, Kiu, 13 Dec. 1963, fr, Whitmore BSIP 3876 (L, SING)3. —N.E. Guadalcanal I.:
Rere River, c. 3 miles inland, 24 Nov. 1963, fr, Whitmore BSIP 2780 (L, SING)3.
AUSTRALIA. Queensland. Near Noah Head. Daintree National Park, 16°07´25.9˝S, 145°27´25.0˝E, alt. ca.
100 m, 13 Dec. 2004, fr.s, Kajita & Takayama 04121301 (TI)2; Kulki area, Cape Tribulation, Daintree National
Park, on sandy beach. 16°04´S, 145°30´E, alt. 0 m, 13 Dec. 2004, fr.s, Kajita & Takayama 04121302 (TI)2 &04121303 (TI)2/S; Cook Distr., Mission Beach, near Clump Point jetty, 17°05´S, 146°00´E, alt. 10 m, 25 Aug.
1978, fl, Thorsborne et al. 467 (L)2; Cow Bay, 16°08´35.4˝S, 145°27´14.2˝E, 15 Nov. 2000, Tateishi & Omine
53108 (TI, URO)4.
KIRIBATI. Phoenix Islands. Kanton I.: Jetsam, Feb. 1958, drifted seeds, Degener & Degener 24628 (TI)S.
VANUATU. Malekula I.: Malekula mainland, opposite Awei I., Maskelynes, alt. 5 m, 16 Nov. 1992, fl, Curry
910 (L)2. —Efate I.:Mt. Bernier, fr, Konishi 1299 (TNS)1-2/S.
NEW CALEDONIA. Auf den Bergen bei Oubatche, alt. 1000 m, 13 Dec. 1902, fl, Schlechter 15398 (L, W)2;
Hienghene, 26 Jun. 1971, fl, McKee 23920 (NOU); Cours moyen et inferieur de la Tchamba, 17 Aug. 1966, fr,
Nothis 242 (NOU); Basse Valee de la Tchamba, 13 Feb. 1968, fl, McKee 18393 (NOU); Vallee de la Nessepue, a 5
km de la mer, 20 Oct. 1970, fl, Schmid 3447 (NOU); Poindimie: Ina, 10 Sep. 1976, fl, McKee 31941 (NOU)2.

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FIJI. Viti Levul I.: near Naivi-dula village, NE. part of island, 19 Oct. 1960, fl, Hotta 3359 (KYO, TI)2; village
of Nasau, NE. part of island, alt. ca. 250 m, 13 Oct. 1960, fl, Hotta 3062 (KYO)2; Mba, Nandala Creek, alt. ca.780 m,
9-25 Sep. 1947, fl, Smith 6253 (L)2; Tailevu, hills east of Waininbuka River, vicinity of Ndakuivuna, alt. 100–200 m,
15–27 Apr. 1953, fl, Smith 7034 (L)2; Namosi, hills bordering Wainavindrau Creek, vicinity of Wainimakutu, alt.
150–250 m, 17 Sep.–8 Oct. 1953, fl, Smith 8851 (L)2; Sleua, hills between Navua River and Wainiyaku Creek, alt.
100–200 m, 20 Oct. 1953, fl, Smith 8981 (L)2; Sleua, hills west of Waivunu Creek, between Ngaloa and Korovoa,
alt. 50–150 m, 23 Nov.–7 Dec. 1953, fl, Smith 9497 (L)3; Naitasiri Prov., vicinity of Nasinu, 9 mile from Suva, alt.
150 m, 20 Oct. 1927, Gillespie 3401 (BO)2. —Vanua Levu I.: Thakaundrove, hills between Vatukawa and Watningio Rivers, Ndrekeniwai Valley, 24 Nov. 1933, Smith 583 (BO)2. —Kandavu I.: Namalata isthmus region, alt. 0–
30 m, 11–18 Oct. 1933, fl, Smith 19 (BO)2. —Uounivanoua I.: Tailevu Distr., alt. 50 m, 21 Jul. 1955, yfr, McKee
2819 (L)2.
TONGA. Vava’u I., north part of island, east of Mataki Niu’a, alt. ca. 140 m, 12 Aug. 1960, fl-bud, Hotta
4957(KYO)2 & 4965 (KYO)2; Mt Mo’ybgakafal; transect from SW corner of Lake ‘Ano, Lat/Long: 18°39´37˝S,
174°02´72˝W, alt. 800 ft, 22 Oct. 1997, Wood et al. 6942 (PTBG). —Eua Isl., west of Mt. Koloaki-lupe-tonga, eastof Pngai Village, 10 Sep. 1960, fr, Hotta 5480 (KYO)2.
SAMOA. Upolu I. Southern coast, Safata,Saanapu, on trail in mangrove forest, alt 0 m, 26 Oct. 2002, fr.s,
Kajita et al. 02102605 (TI)2/S; Satatoa mangrove forest, alt 0 m, 28 Oct. 2002, fr.s, Kajita et al. 02102801 (TI)2/S.
Entada tonkinensis Gagnep.
JAPAN. Ryukyu Islands. Yakushima I.: Yaku-cho, Anbo, 11 Aug. 1911, Nakano s.n. (MAK)2; ibid., Sep.
1921, Koidzumi s.n. (KYO)2; ibid., 3 Sep. 1926, Masamune s.n. (TI)2; ibid., 11 Aug. 1954, Takeuchi s.n. (TI)2;
ibid., 10 Jul. 1968, Hatusima 31557 (KAG)2; ibid., Jun. 1971, fl, Akahoshi s.n. (KAG)2; ibid., 4 Sep. 1984, fr.s,
Murata et al. 17162 (TI, TUS, URO)2/L; ibid., 23 May 1983, Moto s.n. (TI)2; ibid., 7 Jun. 1996, fl, Kurima &Tateishi 131 (TI, URO)2; ibid., 20 Jun. 1997, fl, Kurima et al. 121 (TI, URO)2; ibid., 22 Jun. 2001, Wakita & Tateishi 210622 (URO)2/L. —Amami-Oshima I.: Sumiyo-son, Higashinakama, 7 Dec. 1900, Uchiyama s.n. (TI)3-4;
ibid., 13 Apr 1910, Kamiya s.n. (MAK)3; ibid., 3 Jun. 1911, Ueda s.n. (MAK)3; ibid., 16 Jul. 1919, Kawagoe s.n.
(KAG); ibid., 6 Mar. 1924, Tashiro s.n., (KYO, TAIF)2; ibid., 15 Nov. 1927, Saito 1567, (TI)3; ibid., Jun. 1982, fl,
Tabata s.n. (KAG); ibid., 2 Apr. 1988, Iwano 19845 (TUS)3; ibid., 14 Feb. 1990, fr, Murata & Nemoto s.n. (TUS)3;
ibid., 29 Oct. 1996, Kurima et al. 120 (TI, URO)3/L; ibid., 20 Jun. 1997, Kurima & Matsumura 127 (URO)L; ibid.,
1 Aug. 1998, fr, Tateishi & Yamashiro s.n. (URO)3/L; ibid., 30 Jun. 1999, Yamazaki s.n. (TI); ibid., 21 Nov. 1999,
fr, Tateishi & Kurosawa 52005 (URO)3/L; ibid., 24 Jun. 2001, fl & fr, Wakita & Tateishi 210624-1 (URO)2-3/L &210624-2 (URO)2-3/L; ibid., 1 Jun. 2002, Wakita 210601 (URO)3.
TAIWAN. Taipei Co.: Hsintien, alt. 100–200 m, 8 May 1985, fl, Huang 2803 (TAI, TUS)2; Hsintien–Urai,
27 Mar. 1927, Suzuki s.n. (KAG)2; Taipei Forest Exp. Station, 24 Aug. 1914, Kawagoe s.n. (KAG)2. —Illan Co.:
Tatung Village, Dulishan–Nioudou, alt. 300–400 m, 15 Sep. 1983, fr, Ohashi & Nemoto 16849 (TAI, TUS, URO)2/
L; Tahtung, 24°40´42.9˝N, 121°35´97.7˝E, 26 Sep. 2003, Wakita 240926A (URO)2/L, ibid., 24°40´32.4˝N, 121°
36´29.3˝E, alt. 300 m, 26 Sep. 2003, Wakita 240926B (URO)2/L; Shuanglienpei~Juntou, alt. 100–500 m, 15 Oct.
1984, Tateishi & Nemoto 20677 (KYO, TI, TUS)2-3; Juntou, 24°45´03.6˝N, 121°39´92.3˝E, alt. 250–300 m, 25
Sep. 2003, Wakita 240925A & 240925B (URO)2; Xiaonan’ao, Unknown collector, s.n. (MAK)2. —Nantou Co.:
Yuchih-hsiang, Lienhuachih, 120°53´E, 23°55´N, in 1926, Araki s.n. (TUS)3; Puri, 2 Nov. 1929, Saito 7810 (L,TI)2.
CHINA. Fujian. Longxi Diqu, Xinxu Gongshe, Maping, alt. 400 m, 4 Jun. 1959, Huang 190443 (IBSC)2.
Hong Kong. Tai Po Kau Nature Reserve, Man Ping, Hoi Ha, 22°25´33˝N, 114°10´55˝E, alt. 170 m, 4 May 2005,
fl, Ohi-Toma 20050504 (TI)2. Guangdong. North River, Man Sze Hop, 11 Apr. 1914, fl, Tutcher 10781 (IBSC);
Luofo Shan, in valley, 6 May 1978, fl, K’tung 6063 (IBSC, L)2. —Yunfu Shi: Zhucun, 15 May 1934, fl, Wang
36902 (IBSC)2; Dayonghe, 20 May 1934, fl, Wang 36901 (IBSC)2; Huangdongkeng, 13 Feb. 1929, fr.s, Huang
1841 (IBSC)L. —Zhaoqing Shi: Dinghu, Dongkeng, 28 Jun. 1976, Shi 12324 (IBSC)2; Dinghu, Jiukeng–Taikeng,
alt. 400 m, 23 Jun. 1964, fr.s, Ting & Shi 10251 (IBSC)2/L; Dinghu, Qingyunsi, Houshan, 13 Jul. 1979, fl, Shi
13831 (IBSC)2/L. —Huizhou Shi, Boluo Xian, Mt. Luofu, Baishuimen, 27 May 1930, fr, Chun 41127 (IBSC)2;

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Boluo Xian, Mt. Luofu, Sulaoguan, 26 Feb. 1930, fr.s, Gao 50107 (IBSC) 2/L; Boluo Xian, Mt. Luofu, Niujiaokeng, 6 May 1978, fl, Staff of IBSC 6063 (IBSC)2. —Jiangmen Shi, Taishan Xian, Dalongdong, Niuyuanshan, 31
Aug. 1930, fr, Tso 22363 (IBSC)2. Guangxi. in 1958, fr.s, Chang 619, (IBSC)2. —Liuzhou Diqu, Luoxiu Gongshe, Luoxiang Shan, alt. 300 m, 4 Apr. 1982, fl, Dayaoshan Exped. Team 13883 (IBSC)2. —Baise Diqu, Lingyun
Xian, Lingzhan, Baiji Cun, Langguangou, alt. 450 m, 15 Sep. 1989, fr, South China Exped. Team 1609 (IBSC)2;
Xiancheng–Yuhong, alt. 1300 m, 7 Dec. 1957, fl, Chang 11106 (IBSC)2; Napo Xian, Poman, alt. 850 m, 17 Oct.
1997, Akiyama et al. 1244 (TI, URO)2; Napo Xian, Nonghua Reserve, alt. 900 m, 10 May 1989, fl, South China
Exped. Team 505 (IBSC)2. —Quinzhou Diqu, Shangsi Xian, Nangui, Muxi, 9 Nov. 1958, Chang 13107 (IBSC);
S.E. of Shangsi, near Iu Shan village, Mts. Shiwan Dashan, 7 May 1933, fl, Tsang 22232 (IBSC, P)2; Naliang–
Damian, 15 Apr. 1956, fl, Academia Sinica Guangdong Bot. Exped. 2368 (IBSC)2; Mts. Shiwan Dashan, Fuqin
Xiang, alt. 400 m, 23 Mar. 1944, fl, Chun 4765 (IBSC)2; Shiwan Dashanqu, Dong’an Xiang, Zhumu He, alt. 200
m, 8 Jan. 1944, fr, Chun 4325 (IBSC)2/L; Guitai, Wugushan , alt. 380 m, 12 Apr. 1959, fl, Gan 40035 (IBSC)2. —Nanning Diqu, She-feng Dar Shan, alt. 2000 ft, 28 Oct. 1928, fr, Chang 8279 (IBSC)2, Longjin, Daqing Shanq,
Nadian, alt. 450 m, 25 Dec. 1957, fr, Tam 57616 (IBSC)2; near Daming Shan, alt. 300–350 m, 10 May 1957, fl,
Chun 12428 (IBSC)2; Longzhou Xian, Daqing Shan, 1 Jun. 1935, Gao 55206 (IBSC)2. Lo-hsiang, 1 Dec. 1928,
Sin 3800 (IBSC)2, 18 May 1929, fl, Sin 8277 (IBSC)2. Yunnan. No precise locality, in 1940, Chang 1475 (IBSC)2.
—Honghe: Dawei Shan, below Jinzhu, Dashutang, alt. 2300 ft, 4 Apr. 1940, fl, Wang et al. 100381 (IBSC)2; Dawei
Shan, Laozhai, alt. 2600 ft, 25 Feb. 1940, fr, Wang et al. 100161 (IBSC)2/L; Hekou, 27 Apr. 1940, fl, Chang 1544
(IBSC)2.
VIETNAM. No precise locality, 23 May 1962, fr, Phanchung 1563 (HNU)L. —Vinh Phu Prov.: Near Tu
Vu, Valley of Banton, in 1888, Balansa 2130 (P, holotype and isotype of E. tonkinensis)2; Mt. Tam Dao, 1 Aug.
1917, fr, Hayata s.n. (TI); ibid., on the way to Dihn Rung Rinh Peak, alt. 950–1000 m, 10 Dec. 1996, fr.s, Tateishi
44188 (URO)2/L; ibid., 16 Aug. 1997, Tateishi 1283 (URO)2/L. Ha Son Binh (Soy Tay) Prov.: Mt. Bavi, alt. 700m, 24 May 1940, fl, Petelot 2468 (HNU)2; Bavi, Dinh Ngoc Hoa Peak, 11 Aug.1997, Tateishi 1228 (URO)2/L. —Ha Nam Ninh Prov.: Cuc Phuang, 13 Jan. 1983, Russian & Vietnamese Research Group 1258 (HN), Cuc Phuang
National Park, 20°21´00˝N, 105°35´37˝E, alt. 365 m, Murata et al. 20041218-39 (TI)2/L.