Corresponding author: Yu-Min Shui (
Academic editor: X.-H. Jin
Since 2016, KIB (Kunming Institute of Botany) and CPC (Centre for Plant Conservation of Vietnam) have conducted several surveys in the transboundary karst regions in Northern Vietnam and Southern China and seven new species in the genus
Chen W-H, Radbouchoom S, Nguyen HQ, Nguyen HT, Nguyen KS, Shui Y-M (2018) Seven new species of
The Tonkin region is one of the biodiversity hotspots worldwide (
Since 2016, for nearly three months, transboundary surveys have been conducted in Northwestern and Northeastern Vietnam (Bac Kan, Cao Bang, Lao Cai, Phu Tho, Tuyen Quang) and Southern China (Southwestern Guangxi and Southeastern Yunnan). After review of the type specimens and taxonomic publications within the regions (
The geographical distribution of the seven new species of
The new species is similar to
VIETNAM. Phu Tho Province, Xuan Son county, Xuan Son National Park,
Rhizome: 4–6 cm long, 0.5–1 cm in diam. Stipule pale brown to reddish, triangular, glabrous, 0.7–1 cm × 0.2–0.4 cm, margin entire, apex acuminate. Leaves: petiole terete, 7.5–11 cm long, 1–2 mm in diam., densely covered by red lanes; blade greenish, asymmetric, broadly ovate or rotund, 8–14 × 6.5–9 cm; base cordate; apex subacute to obtuse; margin denticulate, short ciliate; adaxially greenish, almost flat, slightly glabrous and extremely sparely strigillose between secondary veins; abaxially red, veins densely red pubescent; venation palmate, 5–6 primary veins, secondary veins brunching dichotomous, tertiary veins reticulate. Inflorescence: dichasial cyme, peduncle erect, 6–9 cm long, red or brownish villous; bracts caducous, triangular to lanceolate, reddish, 5–7 × 2–3 mm, apex acuminate, margin entire. Staminate flower: pedicels 0.5–1.5 cm long; tepals 4, white, adaxially subglabrous, abaxially glabrous; outer tepals 2, broadly ovate, 0.7–1.5 × 0.6–1.2 cm; inner tepals 2, white, oblanceolate to oblong, 0.8–1.6 × 0.2–0.5 cm, base cuneate to rounded, apex acute to obtuse, margin entire; androecium actinomorphic, stamens numerous; filaments fused at base; anthers yellow, obovate, 1–1.2 mm long, apex convex, shorter than filaments, with longitudinal slits. Pistillate flower: pedicel 0.4–0.7 cm long; tepals 5, white; outer tepals 3, broadly ovate, 0.8–1.2 × 0.4–0.6 cm, apex obtuse to rounded, margin entire; inner tepals 2, ovate, 0.5–0.7 × 0.4–0.6 cm, apex obtuse to rounded, margin entire; styles 3, free; stigmas bifid with twisted bands; ovary reddish, 0.2–0.3 cm long, 0.2–0.4 cm in diam., with white papillose and 4 thickened corns; placentation axile, 4-loculed, placentae 2 each locule. Fruit berrylike, white papillose, wingless and with 4 thickened rib-like horns.
Flowering in April–May, fruiting in April–June.
The epithet refers to the white spots on the fruit surface.
The species only grows in deep ground amongst rocks in limestone forests.
The species occurs exclusively in Phu Tho Province, Xuan Son country, Xuan Son National Park of Vietnam.
VIETNAM. Phu Tho Province, Xuan Son county, Xuan Son National Park,
Currently six species from
The new species is similar to
VIETNAM. Bac Kan Province, Nari county, An Tinh committee,
Rhizome: densely villous, 9–19 cm long, 0.5–1 cm in diam. Stipule reddish, triangular, 0.8–1.5 × 0.4–0.7 cm, densely villous outside, glabrous inside, margin entire and ciliate. Leaves: petiole terete, greenish to reddish, villous, 14–19 cm long, 1.5–3 mm in diam.; blade greenish, asymmetric, ovate to widely ovate, 11–17 × 8.5–11.5 cm, slightly rugose; base cordate, apex caudate, margin serrulate and long ciliate; adaxially greenish, sparsely brevi-setose, abaxially greenish, sparely villous and densely strigose on veins; venation palmate, 6–8 primary veins, secondary veins brunching dichotomous, tertiary veins slightly reticulate, veins prominent on both sides. Inflorescence: dichasial cyme, peduncle erect, 4–10 cm long, villous; bracts caducous, triangulate to lanceolate, reddish, 5–7 × 2–3 mm, adaxially red villous, apex acuminate, margin serrate and ciliate, abaxially glabrous. Staminate flower: pedice 1–1.5 cm long; tepals 2, white to pink, adaxially red villous, abaxially glabrous; tepals 2, widely ovate, 1.5–2 × 0.7–1 cm, apex obtuse to rounded, margin entire; androecium actinomorphic, stamens numerous, filaments almost free, 1–2 mm long, anthers yellow, obovate, 1–1.2 mm long, apex convex, shorter than filaments, with two lateral longitudinal slits. Pistillate flower: pedicel 1–1.5 cm long; tepals 2, white to pink, adaxially red villous, abaxially glabrous; outer tepals 2, widely ovate, 1–1.5 × 0.7–1.2 cm, apex obtuse to rounded, margin entire; styles 3, free, stigmas bifid with twisted bands; ovary white, sparely red villous; placentation axile with 2-segments per locule. Capsule nodding, 0.35–0.5 cm long, 0.27–0.31 cm in diam. (wings excluded), with 3 subequal wings; abaxial wing 0.6–1 × 0.3–0.7 cm; lateral wings 0.4–0.7 × 0.5–0.7 cm.
Flowering in May–June, fruiting in June–August.
The epithet refers to the habitat of the new species: the bamboo forests along streams.
The new species just grows in bamboo forests along watersides.
CHINA. Guangxi Zhuang Autonomous Region, Longzhou Xian, Chunxiu community,
The new species seems to be a member of the first group with rhizome and without erect stem in
The new species is similar to
VIETNAM. Tuyen Quang province, Lam Binh district, Thuong Lam community, in the secondary tropical evergreen lowland rainforest and broad-leaved forests on the slopes and ridge of crystalline limestone hills with highly eroded earth, grow on ground soil, male and female flowers white, fruit berry, green or pink,
Rhizome: 3–5 cm long, 1.0–1.7 cm in diam. Stipule triangular, 0.8–1×0.2–0.4 cm, pale brown to reddish, margin entire, apex acuminate, adaxially glabrous, abaxially red hirsute. Leaves all basal, alternate; petiole terete, 9–15 cm long, 2–5 mm in diam., densely reddish hirsute; blade dark green or brown, asymmetric, broadly ovate, 7–14× 3.5–9.0 cm, base cordate, oblique, apex subacute to obtuse, margin denticulate, short ciliate, adaxially dark green, almost glabrous, usually with white spots, abaxially dark red, densely red lanate on veins; venation palmate, 6–7 primary veins, secondary veins brunching dichotomous, tertiary veins obviously reticulate, densely velutinous on veins. Inflorescence dischasial cyme, axillary, peduncle 3–5 cm, erect, red to brownish villous; bracts pale greenish, triangular to lanceolate, 1–1.5× 2–3 mm, margin dentate and ciliate, apex acuminate. Staminate flower: pedicel 2.2–3.5 cm, glabrous above the middle, villous below the middle; tepals 4, white-pinkish, glabrous; outer tepals 2, ovate, 1.2–1.5× 0.6–1 cm, apex acute, base cuneate, margin entire; inner tepals 2, oblanceolate to oblong, white-pinkish, 0.8–1.6× 0.2–0.4 cm, base cuneate, apex acute, margin entire; stamens numerous, filaments free, anthers yellow, obovate, 1–1.2 mm long, apex rounded, shorter than filament, with longitudinal slits. Pistillate flower: pedicel 8–1.6 cm long, hirsute; tepals 5, white to pinkish, glabrous; outer tepals 3, ovate, 0.8–1.0× 0.3–0.5 cm, apex obtuse to rounded, margin entire; inner tepals 2, ovate, 0.6–1.0× 0.2–0.4 cm, apex obtuse to rounded, margin entire; styles 3, free, stigmas bilobed, with twisted band; ovary green or pinkish, 1.5–2.0 cm long, 0.3–0.5 cm in diam., cylindric, wingless, puberulent; placentation axile, 3-locular, placentae partly branching 2–4 each locule. Fruit berrylike, wingless.
Flowering in October–December, fruiting in November–January next year.
The epithet refers to the upward fruit when nearly mature (Fig.
The species just grows on soil within the secondary tropical evergreen lowland rainforest and broad-leaved forests on the slopes and ridge of crystalline limestone with highly eroded earth at an elevation 400–700 m above sea level.
The species occurs exclusively in Tuyen Quang Province in Vietnam.
VIETNAM. Tuyen Quang province, Lam Binh district, Thuong Lam community,
The new species should be a member of
The new species is similar to
CHINA. Guangxi Zhuang Autonomous Region, Jingxi county, Gulongshan,
Rhizome: 0.3–0.5 cm in diam. Stipule triangular, 0.2–0.4 × 0.6–0.8 cm, apex acute, adaxially subglabrous, abaxially hispid. Leaves all basal, alternate; petiole 5–9 cm long, glandular-hispid; blade greenish, asymmetric, ovate or ovate-lanceolate, 12–18 × 5–9 cm, base cordate, auricular, oblique or slightly oblique, apex caudate, margin serrulate; adaxially greenish, tubercular-setose, with white semi-circle in the middle and with dark red patches along the main nerves; abaxially greenish, glandular-pubescent along veinlets, sparsely villous and densely strigose on main veins; venation palmate, ca. 7 primary veins, secondary veins branching dichotomous, tertiary veins slightly reticulate, veins prominent adaxially. Inflorescence dichasial cyme, axillary, peduncle 5–9 cm, glandular-villous; bracts caducous, oblong-lanceolate, ca. 5 × 1 mm, apex acuminate, adaxially glabrous, abaxially long setulose. Staminate flower: pedicel 1.1–1.8 cm long, glandular-villous; tepals 4, pinkish to pink; outer tepals 2, widely ovate, 0.7–1.2 × 0.7–0.9 cm, apex rotund, margin entire, adaxially glabrous, abaxially glandular-villous; inner tepals 2, narrowly oblong or obovate, 0.6–1.0 × 0.2–0.25 cm, apex acute; androecium actinomorphic; stamens numerous, filaments connected to a half, ca. 1 mm long; anthers yellow, obovate, ca. 0.5 mm long, apex emarginate. Pistillate flower: pedicel ca. 1.1 cm long, glandular-villous; tepals 3, pinkish or pink; outer tepals 2, widely ovate, ca. 0.9 × 1.0 cm, apex rotund, margin entire, adaxially glabrous, abaxially glandular-villous; inner tepal 1, oblong, 5–6 × 2–3 mm, base cuneate, apex acute, margin entire; styles 3, fused at base; stigmas spiralled, papillose; ovary green, elliptic, 0.8–0.9 cm long, glandular-villous; placentation parietal, 2-segmented per carpel. Capsule nodding, 3-winged unequally; major wing ca. 3.5 mm long, lateral wings ca. 2.1 mm long.
Flowering in February–May, fruiting in May–June.
The epithet refers to the locality of the type specimens.
The species only grows on the moist surface of shady cliffs at the entrance to shallow caves in a deep valley.
The species occurs exclusively in Jingxi county of Guangxi in China.
In
The new species is similar to
VIETNAM. Bac Kan Province, Nari county, Kim Hy National reserve,
Rhizome: slender, 2.5–7 mm long, 1.5–2 mm in diam. Stipule reddish, triangular, subglabrous, 1–1.8 × 8–1.2 mm wide. Leaves: petiole reddish, erect, villous, 1.5–3.5 cm long, 0.5–1 mm in diam.; blade asymmetric, widely ovate or rotund, 2.2–2.6 × 1.6–1.8 cm, base cordate, apex obtuse to acute, margin slightly serrulate, ciliate; adaxially dark green with white line or stripes along veins, strigose; abaxially slightly greenish striated and short strigose along primary and secondary veins; venation palmate, 4–5 primary veins, 2 or 3 secondary veins brunching dichotomous, tertiary veins reticulate. Inflorescence: dichasial cyme; peduncle 2–3.5 cm long, glabrous; bracts caducous, greenish, triangular, glabrous, apex acute, margin entire. Staminate flower: pedicel ca. 1 cm long; tepals 4, widely ovate, white inside, pinkish outside, both sides glabrous; outer tepals 2, broadly ovate, glabrous, base cuneate, apex obtuse or rounded, margin entire; inner tepals 2, ovate, base cuneate, apex acute to obtuse, margin entire; androecium actinomorphic, stamens numerous, filaments longer than anthers, fused at base; anthers yellow, obovoid, apex emarginate, with longitudinal slits. Pistillate flower: pedicel ca. 1 cm long; tepals 3, white, both sides glabrous; outer tepals 2, broadly ovate, ca. 5 × 6 mm, abaxially slightly pinkish with red spots, base rounded, apex obtuse or rounded, margin entire; inner tepal 1, elliptic to oblong, ca. 6 × 4 mm, base cuneate to rounded, apex obtuse, margin entire; styles 3, free, stigmas bifid with twisted bands; ovary slightly pinkish with red minute spots, ca. 5 mm long, ca. 3 mm in diam. (wings excluded), glabrous, 1-loculed; placentation parietal, 2-branched each placenta; Capsule nodding, trigonous-ellipsoid, subequally 3-winged; abaxial wing lunate, ca. 5 ×3 mm, slightly larger than lateral wings; lateral wings narrowly lunate, ca. 5 × 2 mm. Seeds ca. 0.15 mm long, oblong.
Flowering in April–May, fruiting in May–June.
The epithet refers to the small size of leaves.
The species just grows in limestone crevices of cliffs.
The species occurs exclusively in Nari county of Bac Kan Province in Vietnam.
In
The new species is similar to
VIETNAM. Bac Kan Province, Cho Ra county, Ba Be National Park,
Rhizome, densely white villous, 2–3 cm long, 0.7–1 cm in diam. Stipule green, 1.5–2 × 1–1.5 cm, widely ovate, apex acute to obtuse, margin ciliate. Leaves: petiole erect, densely white villous, 6–13 cm long, 1–3 mm in diam.; blade green, asymmetric, widely ovate, 8.5–18 × 7–12 cm, rugose, usually with one obtuse lobe; base cordate, apex acuminate or acute, margin slightly denticulate, ciliate; adaxially green, densely tubercular-based setose, abaxially pale green, reddish when young, densely villous on veins; venation palmate, 5–6 primary veins, 2 or 3 secondary veins brunching dichotomous, tertiary veins reticulate. Inflorescence: dichasial cyme, peduncle 12–15 cm long, villous; bracts not seen. Staminate flower: pedicels 0.5–1 cm long; tepals 4; outer tepals 2, broadly ovate, 7–11 × 6–8 mm, apex rounded to acute, margin entire sometimes serrate and ciliate; inner tepals 2, oblanceolate, 5–7 × 1.8–2 mm, apex obtuse to acute, margin entire; androecium actinomorphic, stamens numerous; filaments longer than anthers, slightly fused at base; anthers yellow, 1–1.2 mm long, shorter than filaments, with two longitudinal slits. Pistillate flower: pedicel 1.2–2 cm long; tepals 2, white or pinkish, widely obovate to orbicular, 0.5–1 × 0.5–1 cm, apex acute, margin entire; styles 2–3, free; stigmas bilobed with twisted bands; ovary green or pinkish, trigonous ellipsoid, 1–1.5 cm long, 0.5–1 cm in diam. (wings excluded), white or red villous, 1-loculed; placentation parietal, placentae 3, 2-branched. Capsule pale green to pink, trigonous-ellipsoid, 1–1.5 cm long, 0.5–1 cm in diam. (wings excluded), villous, brownish when dried, 3-winged unequally; abaxial wing lunate, 0.9–1.2 × 0.3–0.7 mm; lateral wings lunate, 0.9–1.2× 0.2–0.5 cm. Seeds oblong, ca. 0.2 mm long.
Flowering in March–May, fruiting in April–June.
The epithet refers to the dense villous trichomes of the new species.
The species only grows on cliffs in limestone forests.
The species occurs exclusively in Ba Be National Park (Cho Ra county) of Bac Kan Province in Vietnam.
In
The new species is similar to
CHINA. Guangxi Zhuang Autonomous Region, Jingxi county, Banliang community,
Rhizome: stout, 7–10 cm long, 1–1.5 cm in diam. Stipule reddish, triangular, adaxially glabrous, abaxially villous, 1–1.5 × 0.5–0.7 cm. Leaves: petiole terete, dark red or brown, densely reddish expanding villous or strigose, 5–20 cm long, 0.5–0.7 cm in diam.; blade asymmetric, widely suborbicular to reniform, 7.5–14 × 6–11 cm; base cordate, apex subacute to obtuse; margin long ciliate; adaxially greenish, extremely sparsely short setulose; abaxially dark reddish, velutinous, densely strigose on reticulate veins; venation palmate, 5–6 primary veins, secondary veins brunching dichotomous, tertiary veins netted and obviously reticulate. Inflorescence: dichasial cyme, peduncle 13–16 cm long, villous; bracts caducous, broadly ovate to orbicular, margin serrate-ciliate, apex obtuse to rounded. Staminate flower: pedicel 0.6–1.0 cm long; tepals 3–4, white to pink, adaxially glabrous, abaxially hirsute-villous; outer tepals 2, broadly ovate, 1.2–1.5 × 1–1.3 cm, abaxially white to pale pink, glabrous to sparsely red setulose, base cuneate to rounded, apex obtuse to rounded, margin entire; inner tepals 2, white to pale pink, oblanceolate, 1–1.5 × 0.2–0.5 cm, base cuneate to rounded, apex acute to obtuse, margin entire; androecium actinomorphic, stamens numerous; filaments longer than anthers, slightly fused at base; anthers yellow, obovate, 0.5–1 mm long, apex truncate, shorter than filaments, with longitudinal slits. Pistillate flower: pedicel 1.5–2 cm long; tepals 3, pinkish or white; outer tepals 2, widely obovate to orbicular, 1.5–1.7 × 1.2–1.5 cm, base cuneate to rounded, apex rounded, margin entire; inner tepal 1, elliptic to oblanceolate, 1–1.3 × 0.2–0.4 cm, base cuneate to rounded, apex acute, margin entire; styles 3, fused at base; stigmas spirally twisted; ovary pinkish, trigonous ellipsoid, 0.6–1.2 cm long, 0.4–0.7 cm in diam. (wings excluded), red villous, 1-loculed; placentation parietal upper and axile at base. Capsule pinkish when fresh, brownish when dried, trigonous-ellipsoid, 1–1.5 mm long, 0.5–0.7 mm in diam. (wings excluded), 3-winged unequally; abaxial wing lunate, 0.6–1 ×0.3–0.6 cm; lateral wings narrowly lunate, 0.6–1 × 0.2–0.4 cm; Seeds oblong, ca. 0.2 mm long.
Flowering in August–November, fruiting in December–March next year.
The epithet refers to the reticulate pattern of nerves on the abaxial surface of leaves.
The species only grows on rocks at the entrance to caves or on the shady slope in limestone forests.
The species distributes to the border region between China and Vietnam, e.g. Jingxi county of Guangxi in China and Cao Bang and Tuyen Quang Province in Vietnam.
CHINA. Guangxi Zhuang Autonomous Region, Jingxi county, Renzhuang community, 13 May 2017,
In
We are thankful to Mr. Liu C., Kunming Institute of Botany, Chinese Academy of Sciences and Mr. Chuong Q.N. of Centre for Plant Conservation of Vietnam (CPC), and people committee of Bac Kan, Cao Bang, Lao Cai, Phu Tho, Tuyen Quang provinces, for their help in the field. We also thank Mr. Wei Y.G. in Guangxi Institute of Botany, Mr. Dong W.K. in Beijing Florascape Company for providing collection information in the field and S. W. Guo in Kunming Institute of Botany for taking some photographs. This research is supported by the National Natural Science Foundation of China (grant nos. 31370228, 31770251, 31070174 and 30270109), Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (2015CASEABRI001), the Critical Ecosystem Partnership Fund through Fauna and Flora International (FFI) and People Resources and Conservation Foundation (PRCF).