Corresponding author: Xiao-Hua Jin (
Academic editor: V. Droissart
Species of the
Pandey TR, Jin X-H (2021) Taxonomic revision of
Of about 208 species of
The study area Pan-Himalaya (also referred to as the PH hereafter) ranges from parts of Afghanistan in the west to the Yunnan Province of China in the east, forming a natural phytogeographic unit; it is further divided into 17 subregions (
This work is based on the review of relevant literature and examination of herbarium specimens, supplemented with observations made on living plants in natural habitats. The specimens of
Biogeographical context of the study, the Pan-Himalaya
More than 500 specimens were sorted and about 200 of them, occurring within the PH, were considered for taxonomic characterisation. Species descriptions are based on vegetative and reproductive features as observed directly and/or through stereomicroscopes (Nikon SMZ1000 and Leica S8 APO) for details. Floral parts were rehydrated in boiling water before their observation and measurements were made under the microscopes. Lip and spur morphology was particularly regarded as taxonomically-informative characters. For species delimitation, morphological species concept (
Data on phenology, habitat and distribution were derived from specimen labels and the distribution maps were prepared from the occurrence locations approximated to the corresponding PH county or district; the list of examined specimens is arranged in the geographical order of the PH (
Floral structure in
Altitudinal range of
1a | Plants 4–20 cm tall; petals entire |
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2a | Rachis and ovary pubescent; sterile bracts absent. |
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3a | Leaves glabrous; floral bracts ca. 10 mm long; ovary and pedicel 8–14 mm long; lateral lobes of lip linear with coiled tips, slightly longer than median lobe; spur 8–15 mm long |
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3b | Leaves papillate; floral bracts ca. 3 mm long; ovary and pedicel ca. 7 mm long; lateral lobes of lip filiform with straight tips, much longer than median lobe; spur ca. 4 mm long |
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2b | Rachis and ovary glabrous; sterile bracts present. |
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4a | Lip 3-lobed; spur distinct and always present. |
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5a | Leaves usually 2; ovary ca. 10 mm long; petals falcate linear-lanceolate |
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5b | Leaf usually one; ovary 4–5 mm long; petals obliquely ovate |
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4b | Lip simple, linear; spur indistinct or absent |
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1b | Plants 10–50 cm tall; petals distinctly 2-lobed. |
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6a | Petals deeply 2-lobed, lower lobe linear to linear lanceolate, more than 4 mm long; lateral lobe of lip linear, with circinate tip. |
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7a | Petal upper lobe falcate-oblong, ca. 4 mm long; lower lobe ca. 10 mm long; floral bracts ca. 7 mm long; ovary cylindric, 12–20 mm long |
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7b | Petal upper lobe spatulate-oblong, 12–15 mm long, lower lobe ca. 4 mm long; floral bracts ca. 15 mm long; ovary terete, 22–35 mm long |
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6b | Petals shallowly 2-lobed, lower lobe like a tooth, less than 2 mm long; lateral lobe of lip linear, retrorse, with bent, but not circinate tip. |
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8a | Sepals 3–7 mm long, 2.5–4 mm broad; petals glabrous; spur reflexed, pendulous. |
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9a | Inflorescence laxly to densely many flowered; ovary with pedicel 7–12 mm long; spur 7–8 mm long |
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9b | Inflorescence subdensely 3–12 flowered; ovary with pedicel 8–10 mm long; spur 12–20 mm long |
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8b | Sepals 7–11 mm long, 3–5 mm broad; petals ciliate; spur spreading horizontally. |
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10a | Leaves with green veins adaxially; lip base with a needle-like appendage near entrance of spur |
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10b | Leaves with white veins adaxially; lip base lacking appendage near entrance of spur |
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Terrestrial herbs, 5–20 cm tall. Tubers ovoid-fusiform. Stems pubescent. Leaves 2, opposite, basal; sheathing at base; leaf blade broadly ovate-orbicular to weakly cordate, 1.5–3.1 cm long, 1–2.5 cm broad, apex apiculate. Inflorescences 4–15 cm long, laxly to subdensely 2- to 6-flowered; rachis minutely glandular, pubescent, 1.5–3.8 cm long; floral bracts narrowly lanceolate, ca. 1 cm long, pubescent, apex acuminate. Flowers green, fragrant; ovary and pedicel curved, 7–13 mm long, pubescent. Dorsal sepal ovate, 5.5–6 mm long, 2.5–3 mm broad, apex acute, forming hood with petals; lateral sepals ovate, reflexed to spreading, 5–7 mm long, ca. 2.6 mm broad, apex acute. Petals obliquely ovate-triangular, base broad, 5–6 mm long, 2–2.5 mm wide, apex acute; lip 3-lobed, clawed, spurred; lateral lobes linear, apex recurved-coiled, 6–9 mm long, ca. 0.6 mm wide; mid-lobe linear, ca. 5 mm long, ca. 0.5 mm broad; spur curved, clavate, 8–15 mm long. Column stout; anther locules parallel; pollinia globose-ovoid; caudicle stout; stigma processes closely parallel, united above mouth of spur, lingulate. (Fig.
Flowering from July to September.
Moist grassy hillsides, stream banks, in
Endemic to the Pan-Himalaya; Sikkim and Darjeeling, Bhutan, M Yarlung Zangbo, Yarlung Zangbo-Brahmaputra and S Hengduan. (Fig.
Distribution of
In the past, the taxonomic identity of
China, Yunnan, 3300–3600 m elev., 1906.
Terrestrial herbs, 4–15 cm tall. Tubers globose-oblong. Stems densely papillate-pubescent. Leaves 2, opposite, basal; sheathing at base; leaf blade ovate to orbicular, 1–2.4 cm long, 1–2.2 cm broad, adaxially with yellowish-white venation, densely papillate, apex acute to acuminate. Inflorescence 3–12 cm long, sparsely 2–14-flowered; rachis 2–5.5 cm long, pubescent; floral bracts lanceolate, 3–6 mm long, apex acuminate. Flowers green, faintly fragrant; ovary and pedicel curved, 6–8 mm long, pubescent. Dorsal sepal broadly ovate, ca. 3.5 mm long, ca. 3 mm broad, glabrous, apex obtuse; lateral sepals oblique, ovate-elliptic, deflexed, ca. 4 mm long, ca. 2.5 mm broad, glabrous, apex obtuse. Petals obliquely falcate-ovate, ca. 3.5 mm long, 2–2.5 mm broad, glabrous, entire; lip 3-lobed, spurred; lateral lobes filiform, ca. 10 mm long; mid-lobe linear-lingulate, ca. 3 mm long; spur pendulous, clavate, 1–4 mm long. Column short; anther apex retuse; caudicles short; stigma processes clavate. (Fig.
Flowering from July to September.
Shady cliffs and rocks; 2700–4300 m elev.
S Hengduan; also in N Fujian of China. (Fig.
Distribution of
The photograph of the type specimen housed at the herbarium of Royal Botanic Garden Edinburgh (
China, Sichuan, 1400 m elev., 1893,
Terrestrial herbs, 13–37 cm tall. Tubers ovoid or oblong. Stems erect or ascending, finely papillate-pubescent. Leaves 2, opposite, basal; base narrowed and amplexicaul; leaf blade spreading horizontally, adaxially yellowish-white marked, ovate or orbicular, 4–6 cm long, 3.5–6 cm broad, apex acute. Inflorescence 10–30 cm long, 4–9-flowered; rachis 5–11 cm long, finely papillate-pubescent; floral bracts lanceolate, ca. 7 mm long, apex acuminate. Flowers yellowish-green; ovary and pedicel twisted, 1.2–2 cm long, pubescent. Dorsal sepal ovate, 3–3.5 mm long, ca. 2.5 mm broad, margins ciliate, apex acute; lateral sepals strongly reflexed, obliquely ovate, 5–7 mm long, ca. 4 mm broad, margins ciliate, apex acute. Petals connivent with dorsal sepal, deeply 2-lobed; upper lobe falcate-oblong, ca. 4 mm long; lower lobe linear, ca. 10 mm long; lip deeply 3-lobed above the base, spurred; lateral lobes divaricate, filiform, ca. 1.5 cm long, apex curled; mid-lobe linear, ca. 1 cm long; spur pendulous, 2–2.5 cm long. Column short and broad, anther connective wide; pollinia granular; caudicles linear; stigmatic processes elongated. (Fig.
Flowering in July to September.
Montane forests, grassy valleys; 1400–3000 m elev.
N and S Hengduan; also in Chongqing and Gansu of China (Fig.
Distribution of
China, Sichuan,
Terrestrial herbs, 12–50 cm tall. Tubers oblong or ovoid. Stems erect, pubescent. Leaves 2, opposite, basal; base obtuse-rounded and amplexicaul; leaf blade spreading horizontally, abaxially tinged with greyish-white, adaxially purplish-green, ovate-orbicular, 2–5 cm long, 1–4.7 cm broad, apex acute-acuminate. Inflorescence 8–45 cm long, 2–8-flowered; rachis 4–18 cm long, pubescent; floral bracts lanceolate or ovate, apex acuminate. Flowers yellowish to yellowish-white; ovary and pedicel twisted, 2.2–3.5 cm long, pubescent. Dorsal sepal forming a hood with petals, erect, ovate or oblong, concave, 10–13 mm long, 6–7 mm broad, apex obtuse; lateral sepals reflexed, obliquely ovate or oblong, 11–14 mm long, 7–7.5 mm broad, apex acute. Petals deeply 2-lobed; upper lobe spatulate-oblong, 12–15 mm long, ca. 6 mm broad, margin ciliate, apex obtuse; lower lobe linear-lanceolate, ca. 4 mm long, ca. 1 mm broad, apex acute; lip reflexed, base with a short claw, deeply 3-lobed, spurred; lateral lobes divaricate, linear, 2.5–4 cm long, circinate toward apex; mid-lobe straight, linear, 1.1–1.4 cm long; spur pendulous, cylindrical-subclavate, 2–3 cm long. Column short; anther parallel, connective wide; pollinia granular; caudicles slender, elongated; stigmatic processes closely parallel, lingulate. (Fig.
Flowering from July to September.
Montane forests, grasslands; 2000–4400 m elev.
Yarlung Zangbo-Brahmaputra, Tangut, N and S Hengduan; widely spread in eastern part of Pan-Himalayan Region. Also found in Gansu, Guizhou and Shaanxi of China. (Fig.
Distribution of
Afghanistan (now Pakistan), Darban Valley, Kuram District, 2280 m elev., 1880,
Terrestrial herbs, 10–50 cm tall. Tubers oblong or ellipsoid. Stems erect, papillate-pubescent. Leaves 2, opposite, basal; base narrowed and amplexicaul; blade ovate or ovate-orbicular, 2–7 cm long, 1.5–6 cm broad, apex acute. Inflorescence 8–45 cm long, laxly to densely many (up to 40) flowered; rachis 1.5–8 cm long, papillate-pubescent; sterile bracts ovate to lanceolate, 0.7–1.5 cm long, acuminate; floral bracts ovate-lanceolate, margins ciliate, ca. 0.7 cm long, apex acuminate. Flowers bright green, with often faintly yellowish lip; ovary and pedicel arcuate, 0.7–1.2 cm long, papillate-hairy. Dorsal sepal forming a hood with petals, erect, ovate, concave, 3–5 mm long, 2.5–3.5 mm broad, apex obtuse or acute; lateral sepals reflexed, obliquely ovate-oblong, 3.5–5.5 mm long, 2.5–3 mm broad, apex obtuse or acute. Petals 2-lobed, glabrous; upper lobe obliquely falcate-lanceolate, 3–5 mm long, 1.5–2 mm broad; lower lobe a tooth at the base of upper lobe, ca. 0.5 mm long; lip deeply 3-lobed near the base, spurred; lateral lobes linear, retrorse, embracing erect sepals, 6–12 mm long, apex slightly bent; mid-lobe reflexed, linear, 5–9 mm long, ca. 1 mm broad; spur pendulous, cylindrical-clavate, 7–8(-10) mm long. Column stout, anthers diverging, connective wide; pollinium granular; caudicles short, stout; stigmatic processes converging, subclavate. (Fig.
Flowering from July to September.
Open
N Pakistan, U Ganga and Indus, U Yarlung Zangbo, W, C and E Nepal, Sikkim and Darjeeling, Bhutan, M Yarlung Zangbo, L Yarlung Zangbo, Yarlung Zangbo-Brahmaputra, N and S Hengduan. (Fig.
Distribution of
This species is distributed along the whole range of the Himalaya up to the Hengduan Mountains at elevations between 2000 and 4500 m (temperate to alpine) and thus is the most widespread
China, Yunnan, 1910,
Terrestrial herbs, 10–24 cm tall. Tubers oblong. Stems densely pubescent. Leaves 2, opposite, basal; leaf blade ovate or ovate-orbicular, 2–4.5 cm long, 2–4 cm broad, fleshy, apex acuminate or acute. Inflorescence 8–20 cm long, subdensely 3–12-flowered; rachis 5–10 cm long; floral bracts lanceolate, apex acuminate. Flowers yellowish-green; ovary and pedicel arcuate, fusiform, 0.8–1 cm long, finely papillate-hairy. Dorsal sepal forming a hood with petals, erect, ovate, concave, 5–6 mm long, 3.5–4 mm broad, margin ciliate-denticulate, apex obtuse; lateral sepals oblique, ovate-oblong, reflexed, 6–7 mm long, 3.5–4 mm broad, apex subacute. Petals 2-lobed, glabrous; upper lobe obliquely ovate-lanceolate, 5–6 mm long, 2–2.2 mm broad; lower lobe a tooth at base of upper lobe, ca. 0.5 mm long; lip deeply 3-lobed above base, spurred; lateral lobes linear, retrorse, almost embracing ovary, linear, 1–1.2 cm long, apex bent; mid-lobe linear, reflexed, ca. 1 cm long; spur pendulous, slightly curved, clavate, 1.2–2 cm long. Column stout, anthers parallel, connective wide; stigmatic processes sub-oblong. (Fig.
Flowering in July and August.
Montane forests, shrubby grasslands, alpine meadows; 3000–4300 m elev.
Endemic to the Pan-Himalaya, found only in Hengduan Mountains; S Hengduan. (Fig.
Distribution of
This species grows above 3000 to 4300 m in grassy alpine meadows.
China, Sichuan, 3200 m elev., 1922,
Terrestrial herbs, 18–30 cm tall. Tubers subglobose or ellipsoid. Stems erect, finely papillate-hairy. Leaves 2, opposite, basal; base obtuse-rounded, abruptly narrowed and amplexicaul; leaf blade broadly ovate or orbicular, 3–5 cm long, 3–5 cm broad, apex shortly acuminate or acute. Inflorescence 15–26 cm long, 3–8-flowered; rachis 5–12 cm long, papillate-hairy; floral bracts linear or lanceolate, apex acuminate. Flowers yellowish-green; ovary and pedicel twisted, fusiform, 1.2–1.5 cm long, papillate-hairy. Dorsal sepal forming a hood with petals, erect, ovate, concave, 7–8 mm long, 3–4 mm broad, apex obtuse; lateral sepals reflexed, obliquely ovate, 7–9 mm long, 3.5–4 mm broad, apex subobtuse. Petals shallowly 2-lobed; upper lobe obliquely oblong-lanceolate, 7–9 mm long, 2–2.5 mm broad, margin papillate-ciliate, apex obtuse; lower lobe a tooth at the base of upper lobe, ca. 1.5 mm long; lip reflexed, spurred, adaxially with a needle-like 5–7 mm long appendage near the entrance of spur, deeply 3-lobed; lateral lobes linear-filiform, 2.4–2.8(–4) cm long, apex often curled; mid-lobe linear, 1.2–2 cm long; spur cylindrical-clavate, 1.6–2.4 cm long, horizontally curved. Column stout, anther resupinate; stigma processes narrow, lingulate. (Fig.
Flowering in July and August.
Montane forests with
N and S Hengduan; also in Shaanxi of China. (Fig.
Distribution of
China, Kangding, 3600 m elev., 13 July 1953,
Neotype of
Terrestrial herbs, 12–40 cm tall. Tubers narrowly oblong to ellipsoid. Stems mostly underground, papillate-hairy. Leaves 2, nearly opposite, basal; prominent white veins on adaxial surface, base abruptly narrowed and amplexicaul; leaf blade wide ovate or orbicular, 2.5–6.5 cm long, 2.5–7 cm wide, apex obtuse or acute. Inflorescence 10–35 cm long, laxly 2–10-flowered; rachis 2–15 cm long, papillate-hairy; floral bracts lanceolate or linear-lanceolate, apex acuminate. Flowers yellowish-green; ovary and pedicel strongly arcuate, 1.5–2 cm long, pubescent. Dorsal sepal forming hood with petals, ovate, concave, 7–9 mm long, 4–5 mm wide, apex subobtuse; lateral sepals reflexed, obliquely ovate, 8–11 mm long, 4–5 mm wide, apex acuminate. Petals shallowly 2-lobed; upper lobe oblique, oblong-lanceolate to ovate-lanceolate, 8–10 mm long, 3–4 mm wide, margin papillate-ciliate, apex subacute; lower lobe ca. 1.5 mm long; lip deeply 3-lobed, spurred, lobes reflexed over base; lateral lobes linear-filiform, 2–4 cm long, curled at apex; mid-lobe linear, 1–2 cm long; spur cylindrical-clavate, 1.5–2.5 cm long, often horizontal and curved upwards. Column stout, anthers parallel; pollinia granular; caudicles stout, elongated; stigma processes lingulate. (Fig.
Flowering from June to August.
Thickets, alpine grasslands; 3200–4900 m elev.
N and S Hengduan, also in NE Qinghai of China. (Fig.
Distribution of
According to the protologue,
This research was supported by a grant from the National Natural Science Foundation of China (No. 31670194). The Curators and staffs of the herbaria AMES, B, BM, CAL, E, IBSC, K, KATH, LD, LE, P, PE, RENZ, S, TI, TUCH and UPS are gratefully acknowledged for granting access to their specimen collections. Specifically, Dr. P.J. Cribb (K), Dr. M. Hjertson (UPS) and Dr. R. Vogt (B) were of vital help for ascertaining the type specimens at the respective herbaria, for which they deserve our sincere gratitude. We are grateful to Dr. J. McNeill (E) for his generous suggestions for typification, Dr. P. Ormerod for comments on the manuscript and Dr. H. Æ. Pedersen (C) for providing some literature. Thanks are also due to Til Prasad Pangali Sharma for helping in the maps and Dr. R.Y. Yu for useful suggestions on the manuscript. Special thanks to all the reviewers for their constructive comments on the manuscript.