Taxonomic revision of Habenaria josephi group (sect. Diphyllae s.l.) in the Pan-Himalaya

Tirtha Raj Pandey; Xiao-Hua Jin

doi:10.3897/phytokeys.175.59849

Abstract

Species of the Habenaria josephi group in the Pan-Himalaya region are revised, based on their morphological characters and results of previous molecular phylogenetics. Eight distinctive species are recognised; key to the species, taxonomic descriptions, illustrations and distribution maps are provided. Habenaria josephi is re-instated, based on morphological and molecular evidence; H. wolongensis is synonymised with H. aitchisonii, a neotype for H. tibetica and the lectotypes for H. balfouriana, H. fargesii, H. glaucifolia and H. clarkei are designated.

Keywords

Distribution, lectotypes, morphological characters, neotype, taxonomy

Introduction

Habenaria Willd. is a large genus in the Orchidaceae (Orchidoideae, Orchideae, Orchidinae), with about 891 species (Govaerts et al. 2020), most of which are terrestrial plants. The genus is distributed along the tropical, subtropical and temperate zones of the Old and New Worlds (Pridgeon et al. 2001; Batista et al. 2011) and has three main centres of diversity, i.e. eastern Asia, central and southern Africa and Brazil (Kurzweil and Weber 1992). The plants are characterised by frequently having simple or bifid petals, a tripartite lip, long rostellar arms, stalked stigmas and a well-developed nectariferous spur (Dressler 1993; Pridgeon et al. 2001).

Of about 208 species of Habenaria occurring in the south to east Asian biodiversity hotspot, roughly one fifth are represented in the Pan-Himalaya (Pearce and Cribb 2002; Chen and Cribb 2009; Rajbhandari and Rai 2017; Govaerts et al. 2020). Amongst the Asian clades of Old World Habenaria, one group with two basal leaves and with a temperate or alpine distribution is of particular taxonomic interest: the Habenaria josephi Rchb.f. complex, as the species belonging to this group formed a close alliance (clade XXIV) in a recent molecular study (Jin et al. 2017). Additionally, they share many morphological traits (e.g. pubescent scape and floral parts, mostly 2-lobed petals) and similar habitats, which often renders the species delimitation difficult; owing to this, some species were either misidentified or assigned different ranks in the past (Lang 1984; Pearce and Cribb 2002; Lucksom 2007; Chen and Cribb 2009; Choudhury et al. 2011; Maity et al. 2019). To address these taxonomic inconsistencies, here we attempt to revise the group in light of the recent molecular works (Jin et al. 2017; Raskoti and Ale 2019) and a broader examination of the herbarium specimens and literature. Our study shows eight species from the group occurring in the region, for which general morphological features, taxonomic description and illustrations are provided. Furthermore, a brief history of Habenaria section Diphyllae Kränzl. and an artificial key to the species are also given.

Materials and methods

Study area

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 (Hong 2015, Fig. 1).

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 Habenaria collected from the PH (Fig. 1), preserved at the herbaria AMES, B, CAL, E, K, KATH, KUN, LD, LE, P, PE, S, TI, TUCH, UPS, W and WU (herbarium acronyms according to Thiers 2020) were thoroughly examined; CAL, KATH, PE and TUCH were personally visited and for others, online catalogues were utilised (e.g. ‘www.cvh.ac.cn’ for Chinese herbaria) and high-resolution images of putative type materials were requested.

Figure 1.

Biogeographical context of the study, the Pan-Himalaya A geographic location of the Pan-Himalaya (shaded with green colour) B subregions of the Pan-Himalaya (Hong 2015) 1 Vakhan 2 N Pakistan 3 Jammu and Kashmir 4 U Ganga and Indus 5 U Yarlung Zangbo 6 W Nepal 7 C Nepal 8 E Nepal 9 Sikkim and Darjiling 10 Bhutan 11 M Yarlung Zangbo 12 L Yarlung Zangbo 13 Yarlung Zangbo-Brahmaputra 14 Tangut 15 N Hengduan 16 S Hengduan 17 U Irrawaddy.

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 (Cronquist 1978; Stuessy 2009), along with recent molecular phylogeny (Jin et al. 2017), was taken into consideration.

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 (Hong 2015). Information on distribution outside the PH and on illustrations available from literature are also provided.

Results

The taxonomic history of Habenaria sect. Diphyllae

Habenaria sect. Diphyllae is one of the 32 sections established by Kränzlin (1892) in his first worldwide revision of Habenaria, including 17 species from Africa and Asia. The sections he assigned were based on the degree of dissection of the petals and labellum and on the structure of the gynostemium. The presence of 3-lobed labellum, simple or bilobed broad petals and thick fleshy stigmatic processes were taken as the diagnostic characters for the section Diphyllae (Kränzlin 1892; Kränzlin 1901). Later, Summerhayes (1968) designated H. diphylla as the type for the section and ascribed 24 species from East Africa to it. Several sectional treatments of Habenaria are available for Neotropics and Africa (Batista et al. 2013; Summerhayes 1968), but comprehensive accounts of Asian species are lacking, though country-level treatments were attempted previously (e.g. Pearce and Cribb 2002). The present study has uncovered the occurrence of 11 species of H. sect. Diphyllae in the PH. Eight of these species are found in high mountain habitats (i.e. collected at an altitude above 1500 m) and represent a monophyletic clade, which is here designated as the H. josephi group. The three remaining species (H. reniformis, H. diphylla and H. acianthoides), in contrast, inhabit the tropical climate and are morphologically distinct (e.g. glabrous scape and floral parts, petals always simple) and will not be considered here.

General morphology

Tuber: One or two, globose, oval or elliptic, underground, with few roots dispersed around their junction to the stem. Stem: Erect, terete and slender (sometimes robust in Habenaria aitchisonii), pubescent, often papillate in H. diplonema, H. aitchisonii, H. szechuanica and H. tibetica. Leaves: Typically two, opposite to sub-opposite, appressed to the ground, glabrous (densely papillate hairy in H. diplonema). Leaf blades orbicular or ovate-orbicular, with amplexicaul base and acute or mucronate apex (acuminate in H. diplonema). Inflorescence: Racemose, with few (two to six in H. josephi; up to eight in H. glaucifolia, H. fargesii and H. szechuanica; up to 14 in H. diplonema, H. balfouriana and H. tibetica) to many-flowered peduncle (sometimes reaching up to 40 flowers in H. aitchisonii). Floral bracts are mostly lanceolate with acuminate apex and densely pubescent. Ovary and pedicel: Pubescent, sometimes papillate (e.g. H. aitchisonii, H. balfouriana, H. szechuanica and H. tibetica); curved and twisted. Flower: Generally small-sized (smallest in H. diplonema, larger in H. glaucifolia, H. szechuanica and H. tibetica), greenish to greenish-yellow (Fig. 2), sometimes fragrant (e.g. H. diplonema and H. josephi). Dorsal sepal forms the hood together with the petals, while lateral sepals are deflexed. Six of the species are with distinct 2-lobed petal; H. fargesii has filiform, long anterior lobe; in H. glaucifolia, anterior lobe is lanceolate and smaller than posterior lobe and in H. aitchisonii, H. balfouriana, H. szechuanica and H. tibetica, it is represented by a small tooth at the base of the petals. H. diplonema and H. josephi are with entire petals, albeit the basal part is broadened. Lip shape is a taxonomically-informative feature in this group; though all the species possess 3-lobed lips, the relative size and orientation of especially the lateral lobe is quite variable; H. glaucifolia, H. szechuanica and H. tibetica have much longer lateral lobes which ultimately are coiled at the tip; lateral lobes of H. aitchisonii and H. balfouriana are reflexed up, while those in H. josephi, H. diplonema and H. fargesii are deflexed. A prominent needle-like appendage present at the base of the lip near the opening of the spur is characteristic of H. szechuanica. Spur, too, exhibits remarkable variation; H. diplonema bears a very short spur, in H. aitchisonii, spur is shorter than ovary, while in the rest of the species, the spur is robustly longer than the ovary. The column is well demarcated in all of the species with parallel anthers and stout caudicles; H. josephi and H. glaucifolia have parallel, closely lingulate stigmatic process; the stigma of H. aitchisonii are concave surrounding the opening of the spur.

Figure 2.

Floral structure in Habenaria josephi group A H. josephi B H. glaucifolia C H. aitchisonii (Photographs by X.H. Jin).

Ecology: All species in the Habenaria josephi group are terrestrial herbs growing on moist grasslands, rocky surfaces and alpine gullies. Generally, they occur in the elevation range of 2000–5000 m (Fig. 3).

Figure 3.

Altitudinal range of Habenaria sect. Diphyllae in the Pan-Himalaya.

Taxonomic synopsis

Habenaria sect. Diphyllae Kränzl., Bot. Jahrb. Syst. 16(2): 147. 1892

Key to the species of Habenaria. sect. Diphyllae from the Pan-Himalaya

1a	Plants 4–20 cm tall; petals entire	2a
2a	Rachis and ovary pubescent; sterile bracts absent.	3a
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	1. H. josephi
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	2. H. diplonema
2b	Rachis and ovary glabrous; sterile bracts present.	4a
4a	Lip 3-lobed; spur distinct and always present.	5a
5a	Leaves usually 2; ovary ca. 10 mm long; petals falcate linear-lanceolate	9. H. diphylla
5b	Leaf usually one; ovary 4–5 mm long; petals obliquely ovate	10. H. acianthoides
4b	Lip simple, linear; spur indistinct or absent	11. H. reniformis
1b	Plants 10–50 cm tall; petals distinctly 2-lobed.	6a
6a	Petals deeply 2-lobed, lower lobe linear to linear lanceolate, more than 4 mm long; lateral lobe of lip linear, with circinate tip.	7a
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	3. H. fargesii
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	4. H. glaucifolia
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.	8a
8a	Sepals 3–7 mm long, 2.5–4 mm broad; petals glabrous; spur reflexed, pendulous.	9a
9a	Inflorescence laxly to densely many flowered; ovary with pedicel 7–12 mm long; spur 7–8 mm long	5. H. aitchisonii
9b	Inflorescence subdensely 3–12 flowered; ovary with pedicel 8–10 mm long; spur 12–20 mm long	6. H. balfouriana
8b	Sepals 7–11 mm long, 3–5 mm broad; petals ciliate; spur spreading horizontally.	10a
10a	Leaves with green veins adaxially; lip base with a needle-like appendage near entrance of spur	7. H. szechuanica
10b	Leaves with white veins adaxially; lip base lacking appendage near entrance of spur	8. H. tibetica

Note. Of the 11 species of sect. Diphyllae from the Pan-Himalaya, only the eight species that belong to the high mountain clade (Jin et al. 2017, Fig. 3) are presented here.

1. Habenaria josephi Rchb. f., Trans. Linn. Soc. London, Bot., ser. 2, 3: 114 (1888).

≡ Habenaria aitchisonii var. josephi (Rchb.f.) Hook.f., Fl. Brit. India 6: 152 (1890).

≡ Habenaria diphylla var. josephi (Rchb.f.) N. Pearce & P.J. Cribb, Edinburgh J. Bot. 58: 114. 2001. Type. INDIA, Sikkim, 1849, J.D. Hooker 42 [holotype: K (K000247480 image!); isotypes: K, AMES (00256484 image!), P (P00370608 image!), LE n.v.].

Habenaria clarkei Kränzl., Bot. Jahrb. Syst. 16: 148 (1892). Type. INDIA, Sikkim, J. D. Hooker 42 [lectotype designated here: K (K000247480 image!); isolectotypes: K, AMES (00256484 image!), P, LE n.v.].

Description

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. 4).

Figure 4.

Habenaria josephi A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from FLPH Expedition 13-0845, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering from July to September.

Habitat

Moist grassy hillsides, stream banks, in Betula forest; 3000–4600 m elev.

Distribution

Endemic to the Pan-Himalaya; Sikkim and Darjeeling, Bhutan, M Yarlung Zangbo, Yarlung Zangbo-Brahmaputra and S Hengduan. (Fig. 5).

Figure 5.

Distribution of Habenaria josephi (black circles) in the Pan-Himalaya.

More illustrations

Pearce and Cribb (2002, fig. 41, a–m; as H. diphylla var. josephi).

Additional specimens examined

SIKKIM and DARJEELING: Sikkim, Tungu, 3648–3952 m elev., 1849, J.D. Hooker 42 (K). BHUTAN: Bumthang, Marlungm Tsampa, 4400 m elev., 1949, F. Ludlow, G. Sherriff & J.H. Hicks 19397 (AMES 01946674); Gasa, Upper Mo Chu, 4120 m elev., 1984, I.W.J. Sinclair & D.G. Long 5289 (RENZ); Gasa, Gafoo-la, Upper Pho Chu, 4000 m elev., 1949, F. Ludlow, G. Sherriff & J.H. Hicks 16725 [CAL (CAL0000056823), AMES (01946675)]; Jomolhari (GLORIA site), 4514 m elev., 2010, S. Den et al. 193 (National Biodiversity Centre, Bhutan). M YARLUNG ZANGBO: Yadong, Phari, 1879, Dungboo s. n. [K (K000247481), CAL (CAL0000092705)]; Yadong, North of Phari, 1882, Dr. King’s collectors s. n. (CAL0000092702); Yadong, Tem-la, North of Phari, 1882, G. King’s collectors s. n. (CAL0000092703). YARLUNG ZANGBO-BRAHMAPUTRA: Cona, 3641 m elev., 2013, FLPH Tibet Expedition 13-0957 (PE); Zayü, 4100 m elev., 2013, FLPH Tibet Expedition 13-0845 (PE). S HENGDUAN: Yulong, eastern flank of Lichiang range, 3500 m elev., 1906, G. Forrest 2739 (CAL0000055843).

Note

Hooker (1890) tentatively placed some of the Habenaria species, such as H. reniformis (D. Don) Hook.f., H. diphylla Dalz. and H. aitchisonii Rchb.f., into the section Trimeroglossa. However, H. josephi was assigned a varietal rank under H. aitchisonii. Later, Kränzlin (1892, 1901) proposed the section Diphyllae to accommodate the species with two basal leaves and included H. clarkei, H. glaucifolia, H. diphylla, H. reniformis and H. aitchisonii in the section along with a few other Habenaria species. He followed Hooker’s view regarding the position of H. josephi as a variety of H. aitchisonii, albeit with a note “Die var. Josephi Hook. f. ist nur eine Form, aus den höchsten der oben angegebenen Standorte stammend (the variety josephi is only a form occurring in the highest of the above-mentioned locations)” suggesting that the plant is merely a higher elevation variety of H. aitchisonii. Paradoxically, Kränzlin described H. clarkei in the same publication, based on a duplicate of Hooker 42 (the type specimen of H. josephi) from Sikkim, which was kept at B (distributed from Hooker’s Herbarium at K). After an extensive search, it now appears that Hooker 42 had at least six duplicates: two at K and one each at P, AMES, B and LE, of which the specimen at B was lost during the Second World War, while at LE, the specimen could not be found during a recent search (fide Petr Efimov). The remaining four duplicates are still extant.

In the past, the taxonomic identity of Habenaria josephi became doubtful, often shifting from one name to another, sometimes as Habenaria aitchisoni var. josephi (Rchb.f.) Hook.f. or as H. diphylla var. josephi (Rchb.f.) Pearce & Cribb. Even in recent literature on the orchid species of Sikkim, its type locality is not uniform in this regard. Some botanists treat it as a variety of H. diphylla (Pearce et al. 2001; Pearce and Cribb 2002; Lucksom 2007; Choudhury et al. 2011), while Maity et al. (2019) regard it merely as another synonym of H. diphylla. A closer look at this species reveals it to be not only distinct morphologically, but also well characterised in terms of habitat and distribution. Whereas H. diphylla is predominantly a tropical species of moderate size (10–40 cm tall), broadly distributed from peninsular India to the Philippines, H. josephi is a small-sized (5–20 cm tall) high-elevation temperate to alpine species occurring from Sikkim eastwards to the Hengduan Mountains, i.e. it is endemic to the Pan-Himalaya. Pearce et al. (2001) provided an elaborate discussion on the phenetic variations that delineate this taxon from the other species, yet they assigned it to a varietal rank under H. diphylla. Examination of the type and other dried specimens, as well as living individuals, clearly shows that it is distinct. Short stature, scape without sterile bracts, 2–6-flowered rachis, curved, pubescent ovary, deflexed lateral sepals and lip with stooping (reflexed) lateral lobes that ultimately coil around terminally are amongst the unique diagnostic morphological features. Furthermore, recent molecular studies (Jin et al. 2017; Raskoti and Ale 2019) have consolidated its distinction from similar-looking species. Therefore, a specific rank seems fully justified here.

2. Habenaria diplonema Schltr., Notes Roy. Bot. Gard. Edinburgh 5: 100. 1912.

Type

China, Yunnan, 3300–3600 m elev., 1906. G. Forrest 2812 [holotype: E (E00381985 image!); isotypes: IBSC (0635875!), CAL (CAL0000000748!), P (P00426408 image!)].

Description

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. 6).

Figure 6.

Habenaria diplonema A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from T.T. Yu 14014, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering from July to September.

Habitat

Shady cliffs and rocks; 2700–4300 m elev.

Distribution

S Hengduan; also in N Fujian of China. (Fig. 7).

Figure 7.

Distribution of Habenaria diplonema (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 192, 7–9).

Additional specimens examined

S. HENGDUAN: Muli, Rangetzantze, 3500 m elev., 1937, T.T. Yü 14014 (KUN, PE); Yulong (Lijiang), 2800 m elev., 1935, C.W. Wang 70748 (PE, KUN); Yulong (Lijiang), 4200 m elev., 1914, C. Schneider 2459 (K).

Note

The photograph of the type specimen housed at the herbarium of Royal Botanic Garden Edinburgh (G. Forrest 2812, E00381985) was published along with the protologue, therefore this specimen is the holotype (Art. 9.1, Note 1 (b), Turland et al. 2018).

3. Habenaria fargesii Finet, Rev. Gen. Bot. 13: 528, t. 18A. 1–8. 1901.

Type

China, Sichuan, 1400 m elev., 1893, Farges 1279 [lectotype designated here: P (P00426411 image!)].

Description

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. 8).

Figure 8.

Habenaria fargesii A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from FLPH Sichuan Expedition 152179, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering in July to September.

Habitat

Montane forests, grassy valleys; 1400–3000 m elev.

Distribution

N and S Hengduan; also in Chongqing and Gansu of China (Fig. 9).

Figure 9.

Distribution of Habenaria fargesii (black circles) in the Pan-Himalaya.

More illustrations

Finet (1901, fig. 18A, 1–8).

Additional specimens examined

N HENGDUAN: Sichuan, T.T. Li 644 (PE). S HENGDUAN: Yanyuan, 3000 m elev., 2015, FLPH Sichuan Expedition 152179 (PE).

4. Habenaria glaucifolia Bureau & Franch., J. Bot. (Morot) 5: 152. 1891.

≡ Senghasiella glaucifolia (Bureau & Franch.) Szlach. J. Orchideenfreund 8: 365. 2001.

Type

China, Sichuan, Prince Henry D’Orleans s. n. [lectotype designated here: P (P00426784 image!)].

Description

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. 10).

Figure 10.

Habenaria glaucifolia A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from K.Y. Lang et al. 945, PE B–G drawn from the same by T.R. Pandey).

Phenology

Flowering from July to September.

Habitat

Montane forests, grasslands; 2000–4400 m elev.

Distribution

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. 11).

Figure 11.

Distribution of Habenaria glaucifolia (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 194, 1–2).

Additional specimens examined

YARLUNG ZANGBO-BRAHMAPUTRA: Mishmi Hills, 1911, Bailey s. n. (E); Zayü, Guyu, Luoma, 2996 m elev., 2009, X.H. Jin et al. SET-ET344 (PE); Zayü, Shang, 3400 m elev., 2013, Jin et al. ST-2554 (PE). TANGUT: Sêrxü, 3900 m elev., 1974, Vegetation Team of Sichuan 5771 (PE). N HENGDUAN: Danba, 3000 m elev., 1940, Qu 7523 (PE); Heishui, 3200 m elev., 1957, Li 73181 (PE); Heishui, 1959, Chuan 1432 (PE); Hongyuan, 2900 m elev., 1957, Zhang & Zhou 22665 (PE); Jinchuan, 2450 m elev., 1957, Li 75398 (PE); Litang-Yalong, 1921, F. Kingdon-Ward 4466 (E); Maoxian, 1952, He & Zhou 13230 (PE); Markam, 2700 m elev., 1957, s. coll. 22969 (PE); Markam, 2900 m elev., W.L. Chen 766 (PE); Xiaojin, Gasiling, 3100 m elev., 1957, J. Zhou 204 (IBSC); Xiaojin, 2450 m elev., 1958, Zhang & Wang 5907 (PE); Yajiang, 3700 m, elev.1961, Jiang 9863 (PE); Yajiang, 2875 m elev., 2006, Boufford et al. 35947 (PE, KUN). S HENGDUAN: Daocheng, 3100 m elev., 1973, Sichuan Vegetation Survey Team 2391 (PE): Daocheng, 4300 m elev., 1973, Sichuan Vegetation Survey Team 3645 (PE): Dêqên, 3100 m elev., 1937, T.T. Yü 9867 (PE); Eryuan, 2900 m elev., 1963, NW Yunnan Team 6339 (PE); Ganluo, 2000 m elev., 1959, Chuan 4191 (PE); Kangding, 1951, Hu & He 10401 (PE); Kangding, 3500 m elev., 1963, Guan & Wang 808 (PE); Kangding, 3650 m elev., 1982, K.Y. Lang et al. 945 (PE); Luding, 2420 m elev., 1959, Jiang & Jin 1949 (PE); Meigu, 2100 m elev., 1959, Chuan 1113 (PE); Meigu, 2300 m elev., 1976, s. coll. 13083 (PE); Mianning, 3300 m elev., 1940, Qu 7353 (PE); Muli, 1921, F. Kingdon-Ward 4571 (E); Muli, Ye-tze, 3100 m elev., 1937, T.T. Yü 7022 (PE); Shangri-la, 3200 m elev., 2010, Kham Expedition 10-3079 (PE); Tianquan, 1959, Chuan 853 (PE); Weixi, 3200 m elev., 1935, C.W. Wang 68011 (PE); Yanyuan, 2500–2600 m elev., 1960, Jiang 5991 (PE); Yulong (Lijiang), 2900 m elev., 1981, Hengduanshan Team of Beijing Institute 02666 (PE); Yulong (Lijiang), 1910, G. Forrest 6050 (PE).

5. Habenaria aitchisonii Rchb.f., Trans. Linn. Soc. London, Bot., ser. 2, 3: 113. 1888.

Habenaria wolongensis K.Y. Lang, Acta Phytotax. Sin. 22(4): 314, 1984, syn. nov. Type. CHINA, Sichuan, Wolong, 2200 m elev., 1982, K. Y. Lang, L. Q. Li & Y. Fei 1528 [lectotype designated by Pandey and Jin (in press): PE (01147127!), isolectotype: PE (01147128!)].

Habenaria diceras Schltr., Notes Roy. Bot. Gard. Edinburgh 5: 101, t. 78. 1912. Type. CHINA, Yunnan, Lijiang Range eastern flank, 2700–3000 m elev., 1906, G. Forrest 3074 [holotype: E (E00381986 image!), isotype: P (P00426380 image!].

Habenaria bihamata Kränzl., Repert. Spec. Nov. Regni Veg. 17: 106. 1921. Type. CHINA, Yunnan, Pe yen tsin, S. Ten s. n. (type: B n.v., probably lost).

Habenaria pubicaulis Schltr., Acta Horti Gothob. 1: 139. 1924. Type. CHINA, Sichuan, ca. 3900 m elev., 1922, Harry Smith 3858 [holotype: UPS (V-091292 image!); isotypes: PE (01516965!), E (E00381983 image!), LD (1073030 image!), S (S07-285 image!)].

Type

Afghanistan (now Pakistan), Darban Valley, Kuram District, 2280 m elev., 1880, Aitchison 413 [holotype: K (K000247484 image!), isotype: AMES (00256482 image!)].

Description

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. 12).

Figure 12.

Habenaria aitchisonii A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from K.Y. Lang et al. 944, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering from July to September.

Habitat

Open Juniperus/Larix montane forests, thickets, grasslands; 1800–4400 m elev.

Distribution

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. 13).

Figure 13.

Distribution of Habenaria aitchisonii (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 192, 4–6).

Additional specimens examined

N PAKISTAN: Hazara, Rawalpindi district, 2500 m elev., 1975, J. Renz 10800 (RENZ). U GANGA & INDUS: Mussoorie, 2128 m elev., 1898, P.W. Mackinnon s. n. (CAL0000092710); Mussoorie, 1824 m elev., 1899, P.W. Mackinnon 22991 (CAL0000092691); Mussoorie, 2300 m elev., 1983, J. Renz & Y.P.S. Pangtey 13641 (RENZ). U YARLUNG ZANGBO: Dinggyê, 4151 m elev., 2013, PE-Tibet team 3421 (PE); Gyirong (Jilong), 3700 m elev., 2013, PE-Tibet team 3825 (PE); Gyirong (Jilong), 2950 m elev., 1975, Qinghai-Tibet Team 7034 (PE). W NEPAL: Karnali, Mugu, Pina-Ghurchi, 2800 m elev., 1985, P.R. Shakya, M.N. Subedi, R.K. Uprety 8783 (KATH); Jumla to Dori Lekh, 3200 m elev., 1979, K.R. Rajbhandari & B.Roy 3378 (KATH). C NEPAL: Dhawalagiri, Mustang, Muktinath-Thorungse, 3900 m elev., 1983, K.R. Rajbhandari 8173 (KATH); Mustang, Dhampus, 2450 m elev., 1988, M. Suzuki et al. 8881592 (KATH); Mustang, Larjung, 2550 m elev., 1996, T. Hoshino et al. 9668062 (KATH); Mustang, 1999, S. Ishizawa et al. 990912020 (TI). E NEPAL: Sagarmatha, Solukhumbu, 3570 m elev., 2005, Watson et al. DNEP3 BX107 (KATH); Solukhumbu, Thame, 2800 m elev., 2013, B.B. Raskoti 00987 (KATH). SIKKIM and DARJEELING: Sikkim, Lachen Valley, 3648 m elev., 1895, R. Pantling 398 (CAL); Lachen, 2584 m elev., 1909, Smith & Cave 2669 (CAL0000092699). BHUTAN: Gasa, Upper Mo Chu, Laya, 3950 m elev., 1983, C. Sargent 114 (RENZ 16003). M YARLUNG ZANGBO: Yadong, Phari, Kang me, 1882, Dr. King’s Collector s. n. (CAL0000092702). L YARLUNG ZANGBO: Mainling, 3100 m elev., 1972, Tibet Chinese Medicinal Plants Survey team 3858 (PE); Nyingchi, near township, 3200 m elev., 1964, Anonymous 793 (PE). YARLUNG ZANGBO-BRAHMAPUTRA: Lhünzê, 3889 m elev., 2013, FLPH Tibet Expedition 13-0890 (PE); Zayü, 4100 m elev., 1973, Qinghai-Tibet Team 1218 (PE); Zayü, 3200 m elev., 1935, C.W. Wang 65772 (PE). N HENGDUAN: Batang, 3520 m elev., 1983, K.Y. Lang et al. 2410 (PE); Danba, 3000 m elev., G.L. Qu 7522 (PE); Luhuo Xian, NW of Luhuo along highway 317, 3385 m elev., D.E. Bufford et al. 33485 (K); Wenchuan, Wolong Nature Reserve, 2220 m elev., 1982, Lang et al. 1528 (type of H. wolongensis, PE). S HENGDUAN: Cawarong, 3450 m elev., 1982, Qinghai-Tibet Team 10547 (PE); Dali, 3000 m elev., 2001, H.K. Kadoorie Team s. n. (PE); Daocheng, 3100–3200 m elev., 1981, Qinghai-Tibetan Team 4235 (PE); Dêqên, 2200 m elev., 1935, C.W. Wang 69920 (PE); Heishui, 3200 m elev., Q. Li 73191 (PE); Heqing, 2730 m elev., 1996, Y.B. Luo 40 (PE); Jinyang, 3500 m elev., K.Y. Lang 14900 (PE); Jiulong, Tanggu Xiang, 3700 m elev., 2005, D.E. Bufford 33420 (K); Kangding, 3470 m elev., 1982, Lang et al. 1534 (PE); Kangding, 3650 m elev., 1982, K.Y. Lang et al. 944 (PE); Luding, 2500 m elev., 1938, Z.B. Wang 9772 (PE); Muli, 3200–3400 m elev., 1983, Qinghai-Tibet Team 14063 (PE); Shangri-la, 4200–4300 m elev., 1981, Hengduan Team 3322, 2832 (PE); Shangri-la, Napa Hai, amongst shrubs, 3350 m elev., 2002, H. Sun 08 (K); Shangri-la (Zhongdian), 3000 m elev., 1937, T.T. Yu 12539 (PE); Weixi, 3600 m elev., 1935, C.W. Wang 68356 (PE); Xiangcheng, 3500 m, elev. Qinghai-Tibetan Team 004810 (PE); Yulong (Lijiang), 3040 m elev., 1922, G. Forrest 22208 (K); Yulong (Lijiang), 2700–3000 m elev., 1913, G. Forrest 10985 (PE); Yulong (Lijiang), 2750 m elev., 1981, Qinghai-Tibet Team 213 (PE).

Note

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 Habenaria species in the Pan-Himalaya. The type material was collected from the Darban Valley along the Pakistan-Afghanistan border in the western end of the distribution (Aitchison 1888). Similar plants from the Hengduan Region were described as new species by Schlechter (1912, 1924) and Kränzlin (1921), respectively, but were later reduced to synonyms (Chen and Cribb 2009). There is considerable variation in the size of the plant, in the colouring of the leaves, with yellowish-white markings that occasionally give a false impression of this being a new species; the peduncle is also quite variable in length as is the density of the inflorescence, from subdensely few-flowered to densely many-flowered. Habenaria wolongensis from Sichuan also falls within the range of variation of H. aitchisonii and is here reduced to synonymy.

6. Habenaria balfouriana Schltr., Repert. Spec. Nov. Regni Veg. 20: 381 (1924).

Type

China, Yunnan, 1910, G. Forrest 6149 [lectotype designated here: E (E00381989 image!); isolectotypes: PE (00340644!), IBSC (0636129!), K (K000796932 image!), P (P00370551 image!)].

Description

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. 14).

Figure 14.

Habenaria balfouriana A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from the isolectotype specimen G. Forrest 6149, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering in July and August.

Habitat

Montane forests, shrubby grasslands, alpine meadows; 3000–4300 m elev.

Distribution

Endemic to the Pan-Himalaya, found only in Hengduan Mountains; S Hengduan. (Fig. 15).

Figure 15.

Distribution of Habenaria balfouriana (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 194, 3–4).

Additional specimens examined

S HENGDUAN: Daocheng, 4236 m elev., 2007, X.H. Jin 9194 (PE); Yanyuan, 3600 m elev., 1983, Qinghai-Tibet Team 12529 (PE); Yulong (Lijiang), 3000 m elev., 1937, T.T. Yu 15316 (PE); Yulong (Lijiang), mountain meadows, 3344 m elev., 1906, G. Forrest 2739 (K).

Note

This species grows above 3000 to 4300 m in grassy alpine meadows. Habenaria balfouriana has a restricted distribution in the southern Hengduan Mountains, sharing the habitat with the similar-looking H. aitchisonii. Though H. aitchisonii and H. balfouriana were found closely allied in a recent molecular study (Jin et al. 2017), they are distinct morphologically; both have yellowish-green flowers with bilobed petals, but the former has an elongated spur that exceeds the length of the ovary and pedicel (versus spur shorter than ovary in H. aitchisonii).

7. Habenaria szechuanica Schltr., Acta Horti Gothob. 1: 140. 1924.

Type

China, Sichuan, 3200 m elev., 1922, Harry Smith 2916 [holotype: UPS (UPS-V-109140 image!); isotypes: E (E00381982 image!), LD (1072390 image!), PE (01516964!), S (S-G-7344 image!)].

Description

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. 16).

Figure 16.

Habenaria szechuanica A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from Hengduan Mountain Team 02687, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering in July and August.

Habitat

Montane forests with Pinus yunnanensis Franch. and Picea sp.; 2900–4000 m elev.

Distribution

N and S Hengduan; also in Shaanxi of China. (Fig. 17).

Figure 17.

Distribution of Habenaria szechuanica (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 195, 1–4).

Additional specimens examined

N HENGDUAN: Songpan, 3450 m elev., 2002, Y.B. Luo 850 (PE). S HENGDUAN: Xiangcheng, 3900 m elev., 1981, Team of Qinghai-Tibetan Plateau 4782c (PE). Yulong (Lijiang), 2900 m elev., 1981, Hengduan Mountain Team 02687 (PE, 4 duplicates).

8. Habenaria tibetica Schltr. ex Limpricht, Repert. Spec. Nov. Regni Veg. Beih. 12: 338. 1922.

Type

China, Kangding, 3600 m elev., 13 July 1953, X.L. Jiang 36260 (neotype designated here: PE 00340847!) (Fig. 18).

Figure 18.

Neotype of Habenaria tibetica Schltr. ex Limpricht X.L. Jiang 36260 (PE 00340847) (Photographed by T.R. Pandey).

Description

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. 19).

Figure 19.

Habenaria tibetica A habit B floral bract C pedicellate ovary with spur D petal E dorsal sepal F lateral sepal G lip (A photographed from the neotype X.L. Jiang 36260, PE B–G drawn from the same specimen by T.R. Pandey).

Phenology

Flowering from June to August.

Habitat

Thickets, alpine grasslands; 3200–4900 m elev.

Distribution

N and S Hengduan, also in NE Qinghai of China. (Fig. 20).

Figure 20.

Distribution of Habenaria tibetica (black circles) in the Pan-Himalaya.

More illustrations

Wu et al. (2010, fig. 195, 5–7).

Additional specimens examined

N HENGDUAN: Gonjo, 3200 m elev., Vegetation Team of Qinghai-Tibet Plateau 9671 (PE); Gonjo, 3800 m elev., 2010, Kham Expedition 10-1872 (PE); Songpan, 3835 m elev., 2002, Y.B. Luo 827 (PE); Xiaojin, Hanniu Region, 3400 m elev., 1959, Z.G. Liu 0359 (PE). S HENGDUAN: Kangding, Muya Region, A-Tai Xiang, 3560 m elev., 1982, Lang et al. 981 (PE, KUN).

Note

According to the protologue, Habenaria tibetica was described by Schlechter, based on two specimens from China: East Tibet, Ta tsien lu, 3600 m elev., Limpricht 2303 and Batang-Litang, 4800–4900 m elev., Limpricht 2277 (Limpricht 1922); consequently, both of these are syntypes as per Art. 9.6 of the ICN (Turland et al. 2018). Many of the type specimens, described by Schlechter together with H. tibetica (e.g. Platanthera minax Schltr. & P. winkleriana Schltr.), were believed to be kept at B and later, their duplicate specimens were recovered at other European herbaria WU and WRSL; however, despite an extensive search, none of the type materials of H. tibetica could be located in any of the world’s major herbaria and could have been destroyed during the Second World War at B. Furthermore, we were unable to find any other original material related to the species. Thus, assuming that all the original material of H. tibetica is lost, it warrants designating a neotype, which is here accomplished. For that purpose, X.L. Jiang 36260 (PE) is designated the neotype according to Art. 9.8 of the ICN (Turland et al. 2018); this specimen was also collected from the original type locality (Kangding, Sichuan). The designated neotype specimen conforms to the description in the protologue and is consistent with the current application of the taxon name (e.g. Chen and Cribb 2009).

Acknowledgements

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.

References

Aitchison JET (1888) The Botany of the Afghan Delimitation Commission. In: The Transactions of the Linnean Society of London, Series 2. Botany 3: 1–139. https://doi.org/10.1111/j.1095-8339.1888.tb00623.x

Batista JAN, Bianchetti LB, González-Tamayo R, Figueroa XMC, Cribb PJ (2011) A synopsis of new world Habenaria (Orchidaceae) I. Harvard Papers in Botany 16(1): 1–47. https://doi.org/10.3100/025.016.0101

Batista JAN, Borges KS, Faria MW, Proite K, Ramalho AJ, Salazar GA, van den Berg C (2013) Molecular phylogenetics of the species-rich genus Habenaria (Orchidaceae) in the New World based on nuclear and plastid DNA sequences. Molecular Phylogenetics and Evolution 67(1): 95–109. https://doi.org/10.1016/j.ympev.2013.01.008

Chen SC, Cribb PJ (2009) Habenaria. In: Wu ZY, Raven PH, Hong DY (Eds) Flora of China (Vol. 25). Science Press, Beijing and Missouri Botanical Garden Press, St. Louis, 144–160.

Choudhury S, Mukherjee SK, Chowdhery HJ (2011) Distribution and diversity of the genus Habenaria Willdenow (Orchidaceae) in India (conference paper). In: Ghosh C, Das AP (Eds) Recent Studies in Biodiversity and Traditional Knowledge in India. Gour Mahavidyalaya, Malda, 81–90.

Cronquist A (1978) Once again, what is a species? In: Romburger JA (Ed.) Beltsville Symposia in Agricultural Research. [2] Biosystematics in Agriculture. Invited papers presented at a symposium held May 8–11, 1977, at the Beltsville Agricultural Research Center (BARC), Beltsville, Maryland 20705. Allenheld, Osmun & Co., Montclair, 3–20.

Dressler RL (1993) Phylogeny and classification of the orchid family. Dioscorides Press, Portland, Oregon.

Finet AE (1901) Les Orchidées de L’Asie Orientale. Revue Générale de Botanique 13: 497–534. [t. 18A. 1–8.]

Govaerts R, Bernet P, Kratochvil K, Gerlach G, Carr G, Alrich P, Pridgeon AM, Pfahl J, Campacci MA, Holland Baptista D, Tigges H, Shaw J, Cribb P, George A, Kreuz K, Wood J (2020) World checklist of Orchidaceae. Facilitated by the Royal Botanic Gardens, Kew. http://wcvp.science.kew.org/

Hong D-Y (2015) Flora of Pan-Himalaya (Vol. 47). Science Press, Beijing.

Hooker JD (1890) The Flora of British India, Orchideae to Cyperaceae (Vol. 6), Part 17. L. Reeve & Co., London, 224 pp. https://doi.org/10.5962/bhl.title.678

Jin WT, Schuiteman A, Chase MW, Li JW, Chung SW, Hsu TC, Jin XH (2017) Phylogenetics of subtribe Orchidinae s.l. (Orchidaceae; Orchidoideae) based on seven markers (plastid matK, psaB, rbcL, trnL-F, trnH-psba, and nuclear nrITS, Xdh): implications for generic delimitation. BMC Plant Biology 17(1–14): e222. http://dx.doi.org/10.1186/s12870-017-1160-x

Kränzlin F (1892) Beiträge zu einer Monographie der Gattung Habenaria Willd. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 16: 52–223.

Kränzlin F (1901) Orchidacearum genera et species (Vol. 1). Mayer & Müller, Berlin.

Kränzlin F (1921) Orchidaceae Ténianae Yunnanense. Repertorium Specierum Novarum Regni Vegetabilis 17(4–7): 99–112. https://doi.org/10.1002/fedr.19210170411

Kurzweil H, Weber A (1992) Floral morphology of southern African Orchideae II. Habenariinae. Nordic Journal of Botany 12(1): 39–61. https://doi.org/10.1111/j.1756-1051.1992.tb00200.x

Lang KY (1984) New species of Orchidaceae from western Sichuan, China. Zhiwu Fenlei Xuebao 22(4): 312–315.

Limpricht W (1922) Botanische Reisen in den Hochgebirgen Chinas und Ost-Tibets. Repertorium Specierum Novarum Regni Vegetabilis. Beihefte 12: 1–515.

Lucksom SZ (2007) The Orchids of Sikkim and North East Himalaya. S.Z. Lucksom, Siliguri.

Maity D, Ghosh J, Pradhan N, Mukherjee SK, Maiti GG (2019) Enumeration of Orchids of Sikkim. Pleione 13(2): 355–384. https://doi.org/10.26679/Pleione.13.2.2019.355-384

Pandey TR, Jin XH (in press) Taxonomic notes and typification of some Habenaria species in the Pan-Himalaya. Phytotaxa.

Pearce N, Cribb PJ (2002) The Orchids of Bhutan. Flora of Bhutan (Vol. 3(3)). Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Pearce N, Cribb PJ, Renz J (2001) Notes relating to the Flora of Bhutan: XLIV. Taxonomic notes, new taxa and additions to the Orchidaceae of Bhutan and Sikkim (India). Edinburgh Journal of Botany 58(1): 99–122. https://doi.org/10.1017/S0960428601000488

Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN (2001) Genera Orchidacearum (Vol. 2). Orchidoideae, part 1. Oxford University Press, New York.

Rajbhandari KR, Rai SK (2017) A Handbook of the Flowering Plants of Nepal (Vol. 1). Department of Plant Resources, Kathmandu, Nepal.

Raskoti BB, Ale R (2019) Molecular phylogeny and morphology reveal a new epiphytic species of Habenaria (Orchidaceae; Orchideae; Orchidinae) from Nepal. PLoS ONE 14(10): e0223355. https://doi.org/10.1371/journal.pone.0223355

Schlechter R (1912) Enumeration and Description of Species of Orchidaceae. Notes from the Royal Botanic Garden Edinburgh 5: 99–113.

Schlechter R (1924) Acta Horti Gothoburgensis 1(3): 139–140. https://doi.org/10.1111/j.1651-2227.1924.tb14521.x

Stuessy TF (2009) Plant taxonomy: The Systematic Evaluation of Comparative Data. Second Edition. Columbia University Press, New York.

Summerhayes VS (1968) Orchidaceae (Part 1). In: Milne-Redhead E, Polhill RM (Eds) Flora of Tropical East Africa. Crown Agents for Overseas Governments and Administrations, London, 236 pp.

Thiers B (2020) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/science/ih/ [accessed: 10 September 2020]

Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ, Smith GF (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten, Koeltz Botanical Books. https://doi.org/10.12705/Code.2018

Wu ZY, Raven PH, Hong DY (2010) Flora of China Illustrations (Vol. 25). Science Press, Beijing and Missouri Botanical Garden Press, St. Louis.