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Taxonomic revision of Habenaria josephi group
(sect. Diphyllae s.l.) in the Pan-Himalaya
Tirtha Raj Pandey1,2,3, Xiao-Hua Jin1
1State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sci-
ences, Beijing 100093, China 2University of Chinese Academy of Sciences, Beijing 100864, China 3National
Herbarium and Plant Laboratories, Godawari, Lalitpur, Nepal
Corresponding author: Xiao-Hua Jin (xiaohuajin@ibcas.ac.cn)
Academic editor: V. Droissart|Received 20 October 2020|Accepted 10 February 2021|Published 06 April 2021
Citation: Pandey TR, Jin X-H (2021) Taxonomic revision of Habenaria josephi group (sect. Diphyllae s.l.) in the Pan-
Himalaya. PhytoKeys 175: 109–135. https://doi.org/10.3897/phytokeys.175.59849
Abstract
Species of the Habenaria josephi group in the Pan-Himalaya region are revised, based on their morphologi-
cal 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, Or-
chidinae), with about 891 species (Govaerts et al. 2020), most of which are terrestrial
plants. e 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). e plants are characterised by frequently having simple or bid
petals, a tripartite lip, long rostellar arms, stalked stigmas and a well-developed nectar-
iferous spur (Dressler 1993; Pridgeon et al. 2001).
Copyright Tirtha Raj Pandey, Xiao-Hua Jin. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
PhytoKeys 175: 109–135 (2021)
doi: 10.3897/phytokeys.175.59849
https://phytokeys.pensoft.net
RESEARCH ARTICLE
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Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
110
Of about 208 species of Habenaria occurring in the south to east Asian bio-
diversity hotspot, roughly one fth 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 oral parts, mostly 2-lobed petals) and similar habitats, which often renders
the species delimitation dicult; owing to this, some species were either misiden-
tied or assigned dierent 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 morpho-
logical features, taxonomic description and illustrations are provided. Furthermore,
a brief history of Habenaria section Diphyllae Kränzl. and an articial key to the
species are also given.
Materials and methods
Study area
e 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).
is work is based on the review of relevant literature and examination of her-
barium specimens, supplemented with observations made on living plants in natural
habitats. e 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 iers 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.
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 stereomi-
croscopes (Nikon SMZ1000 and Leica S8 APO) for details. Floral parts were rehy-
drated in boiling water before their observation and measurements were made under
the microscopes. Lip and spur morphology was particularly regarded as taxonomi-
cally-informative characters. For species delimitation, morphological species concept
Taxonomic revision of Habenaria josephi group 111
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
8ENepal 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.
(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
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
112
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 rst worldwide revision of Habenaria, including 17 species from Africa and Asia.
e sections he assigned were based on the degree of dissection of the petals and la-
bellum and on the structure of the gynostemium. e presence of 3-lobed labellum,
simple or bilobed broad petals and thick eshy 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 avail-
able for Neotropics and Africa (Batista et al. 2013; Summerhayes 1968), but compre-
hensive accounts of Asian species are lacking, though country-level treatments were
attempted previously (e.g. Pearce and Cribb 2002). e 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. e
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
oral 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. aitchiso-
nii, 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). Inorescence: 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-owered peduncle (some-
times reaching up to 40 owers in H. aitchisonii). Floral bracts are mostly lanceolate
with acuminate apex and densely pubescent. Ovary and pedicel: Pubescent, some-
times 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
Taxonomic revision of Habenaria josephi group 113
Figure 2. Floral structure in Habenaria josephi group A H. josephi B H. glaucifolia C H. aitchisonii
(Photographs by X.H. Jin).
together with the petals, while lateral sepals are deexed. Six of the species are with
distinct 2-lobed petal; H. fargesii has liform, long anterior lobe; in H.glaucifolia, an-
terior lobe is lanceolate and smaller than posterior lobe and in H.aitchisonii, H.bal-
fouriana, 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 reexed up, while those in H. josephi, H.diplonema and H.
fargesii are deexed. 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. e 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.
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 eleva-
tion range of 2000–5000 m (Fig. 3).
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; oral 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
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
114
3b Leaves papillate; oral bracts ca. 3 mm long; ovary and pedicel ca. 7 mm
long; lateral lobes of lip liform 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;
oral 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;
oral 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 reexed, pendu-
lous ................................................................................................................... 9a
9a Inorescence laxly to densely many owered; ovary with pedicel 7–12 mm
long; spur 7–8 mm long .....................................................5. H. aitchisonii
9b Inorescence subdensely 3–12 owered; 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 horizon-
tally................................................................................................................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 im-
age!), LE n.v.].
Taxonomic revision of Habenaria josephi group 115
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 pubes-
cent. 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. Inorescences
4–15 cm long, laxly to subdensely 2- to 6-owered; rachis minutely glandular, pu-
bescent, 1.5–3.8 cm long; oral 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, reexed 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).
Phenology. Flowering from July to September.
Habitat. Moist grassy hillsides, stream banks, in Betula forest; 3000–4600 m elev.
Figure 3. Altitudinal range of Habenaria sect. Diphyllae in the Pan-Himalaya.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
116
Figure 4. Habenaria josephi A habit B oral 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).
Distribution. Endemic to the Pan-Himalaya; Sikkim and Darjeeling, Bhutan, M
Yarlung Zangbo, Yarlung Zangbo-Brahmaputra and S Hengduan. (Fig. 5).
More illustrations. Pearce and Cribb (2002, g. 41, a–m; as H. diphylla var. josephi).
Additional specimens examined. SIKKIM and DARJEELING: Sikkim, Tun -
gu, 3648–3952 m elev., 1849, J.D. Hooker 42 (K). BHUTAN: Bumthang, Mar-
lungm Tsampa, 4400 m elev., 1949, F. Ludlow, G. Sherri & 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.
Sherri & J.H. Hicks 16725 [CAL (CAL0000056823), AMES (01946675)]; Jomol-
hari (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 ank of Li-
chiang range, 3500 m elev., 1906, G. Forrest 2739 (CAL0000055843).
Taxonomic revision of Habenaria josephi group 117
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. aitch-
isonii. 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 spe-
cies. 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 speci-
men of H. josephi) from Sikkim, which was kept at B (distributed from Hooker’s Her-
barium atK). 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 (de Petr Emov). e 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 re-
cent 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 moder-
ate size (10–40 cm tall), broadly distributed from peninsular India to the Philippines,
Figure 5. Distribution of Habenaria josephi (black circles) in the Pan-Himalaya.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
118
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-owered rachis, curved, pubescent ovary, deexed lateral sepals and
lip with stooping (reexed) 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. erefore, a specic rank seems fully justied here.
2. Habenaria diplonema Schltr., Notes Roy. Bot. Gard. Edinburgh 5: 100. 1912.
Type. C, 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. Inorescence 3–12 cm long,
sparsely 2–14-owered; rachis 2–5.5 cm long, pubescent; oral bracts lanceolate,
3–6 mm long, apex acuminate. Flowers green, faintly fragrant; ovary and pedicel
curved, 6–8mm long, pubescent. Dorsal sepal broadly ovate, ca. 3.5 mm long, ca.
3 mm broad, glabrous, apex obtuse; lateral sepals oblique, ovate-elliptic, deexed, 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
liform, ca. 10mm long; mid-lobe linear-lingulate, ca. 3 mm long; spur pendulous,
clavate, 1–4mm long. Column short; anther apex retuse; caudicles short; stigma pro-
cesses clavate. (Fig. 6).
Phenology. Flowering from July to September.
Habitat. Shady clis and rocks; 2700–4300 m elev.
Distribution. S Hengduan; also in N Fujian of China. (Fig. 7).
More illustrations. Wu et al. (2010, g. 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. e photograph of the type specimen housed at the herbarium of Royal Bo-
tanic Garden Edinburgh (G. Forrest 2812, E00381985) was published along with the pro-
tologue, therefore this specimen is the holotype (Art. 9.1, Note 1 (b), Turland et al. 2018).
Taxonomic revision of Habenaria josephi group 119
Figure 6. Habenaria diplonema A habit B oral 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).
Figure 7. Distribution of Habenaria diplonema (black circles) in the Pan-Himalaya.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
120
3. Habenaria fargesii Finet, Rev. Gen. Bot. 13: 528, t. 18A. 1–8. 1901.
Type. C, 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, nely 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. Inorescence 10–30 cm long,
4–9-owered; rachis 5–11 cm long, nely papillate-pubescent; oral 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 reexed, 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, liform, ca. 1.5cm 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).
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).
More illustrations. Finet (1901, g. 18A, 1–8).
Additional specimens examined. N HENGDUAN: Sichuan, T.T. Li 644 (PE).
SHENGDUAN: 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. C, 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. Inorescence
8–45 cm long, 2–8-owered; rachis 4–18 cm long, pubescent; oral bracts lanceo-
late 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
reexed, 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 reexed, base with a short claw, deeply 3-lobed, spurred; lateral
Taxonomic revision of Habenaria josephi group 121
Figure 8. Habenaria fargesii A habit B oral 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).
Figure 9. Distribution of Habenaria fargesii (black circles) in the Pan-Himalaya.
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; stig-
matic processes closely parallel, lingulate. (Fig. 10).
Phenology. Flowering from July to September.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
122
Figure 10. Habenaria glaucifolia A habit B oral 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).
Figure 11. Distribution of Habenaria glaucifolia (black circles) in the Pan-Himalaya.
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).
Taxonomic revision of Habenaria josephi group 123
More illustrations. Wu et al. (2010, g. 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, Bouord 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. King-
don-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, 2500–2600m
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 ank, 2700–3000 m elev., 1906, G. For-
rest 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, Si-
chuan, 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!)].
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
124
Type. A (now P), 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. Inores-
cence 8–45 cm long, laxly to densely many (up to 40) owered; rachis 1.5–8 cm long,
papillate-pubescent; sterile bracts ovate to lanceolate, 0.7–1.5 cm long, acuminate;
oral bracts ovate-lanceolate, margins ciliate, ca. 0.7 cm long, apex acuminate. Flowers
bright green, with often faintly yellowish lip; ovary and pedicel arcuate, 0.7–1.2cm
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 reexed, oblique-
ly 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, 1.5–2mm
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 reexed, 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 con-
verging, subclavate. (Fig. 12).
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, Yar-
lung Zangbo-Brahmaputra, N and S Hengduan. (Fig. 13).
More illustrations. Wu et al. (2010, g. 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-orungse,
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, ame, 2800 m elev., 2013, B.B. Raskoti 00987 (KATH). SIK-
KIM and DARJEELING: Sikkim, Lachen Valley, 3648 m elev., 1895, R. Pantling 398
(CAL); Lachen, 2584 m elev., 1909, Smith&Cave 2669 (CAL0000092699). BHU-
Taxonomic revision of Habenaria josephi group 125
Figure 12. Habenaria aitchisonii A habit B oral 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).
Figure 13. Distribution of Habenaria aitchisonii (black circles) in the Pan-Himalaya.
TAN: 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.,
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
126
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). NHENG-
DUAN: Batang, 3520 m elev., 1983, K.Y. Lang et al. 2410 (PE); Danba, 3000m elev.,
G.L. Qu 7522 (PE); Luhuo Xian, NW of Luhuo along highway 317, 3385 m elev.,
D.E. Buord et al. 33485 (K); Wenchuan, Wolong Nature Reserve, 2220 m elev., 1982,
Lang et al. 1528 (type of H. wolongensis, PE). S HENGDUAN: Cawarong, 3450m
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. Buord 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, 3500m,
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. is species is distributed along the whole range of the Himalaya up to
the Hengduan Mountains at elevations between 2000 and 4500 m (temperate to al-
pine) and thus is the most widespread Habenaria species in the Pan-Himalaya. e
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). ere 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 inorescence, from subdensely few-owered to densely many-owered. 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. C, 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 pu-
bescent. Leaves 2, opposite, basal; leaf blade ovate or ovate-orbicular, 2–4.5 cm long,
Taxonomic revision of Habenaria josephi group 127
Figure 14. Habenaria balfouriana A habit B oral bract C pedicellate ovary with spur D petal Edor-
sal sepal F lateral sepal G lip (A photographed from the isolectotype specimen G. Forrest 6149, PE
B–Gdrawn from the same specimen by T.R. Pandey).
2–4 cm broad, eshy, apex acuminate or acute. Inorescence 8–20 cm long, subdensely
3–12-owered; rachis 5–10 cm long; oral bracts lanceolate, apex acuminate. Flowers
yellowish-green; ovary and pedicel arcuate, fusiform, 0.8–1 cm long, nely 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,
reexed, 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 lin-
ear, reexed, ca. 1 cm long; spur pendulous, slightly curved, clavate, 1.2–2 cm long. Col-
umn stout, anthers parallel, connective wide; stigmatic processes sub-oblong. (Fig. 14).
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 Moun-
tains; S Hengduan. (Fig. 15).
More illustrations. Wu et al. (2010, g. 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
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
128
(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. is species grows above 3000 to 4300 m in grassy alpine meadows. Habe-
naria balfouriana has a restricted distribution in the southern Hengduan Mountains,
sharing the habitat with the similar-looking H. aitchisonii. ough 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 owers with bilobed pet-
als, 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. C, 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, nely 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. Inorescence 15–26 cm long, 3–8-owered;
rachis 5–12 cm long, papillate-hairy; oral bracts linear or lanceolate, apex acumi-
nate. 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, 7–8mm
long, 3–4 mm broad, apex obtuse; lateral sepals reexed, 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 ob-
tuse; lower lobe a tooth at the base of upper lobe, ca. 1.5 mm long; lip reexed, spurred,
Figure 15. Distribution of Habenaria balfouriana (black circles) in the Pan-Himalaya.
Taxonomic revision of Habenaria josephi group 129
Figure 16. Habenaria szechuanica A habit B oral 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).
adaxially with a needle-like 5–7 mm long appendage near the entrance of spur, deeply
3-lobed; lateral lobes linear-liform, 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 17. Distribution of Habenaria szechuanica (black circles) in the Pan-Himalaya.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
130
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).
More illustrations. Wu et al. (2010, g. 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. C, Kangding, 3600 m elev., 13 July 1953, X.L. Jiang 36260 (neotype des-
ignated here: PE 00340847!) (Fig. 18).
Description. Terrestrial herbs, 12–40 cm tall. Tubers narrowly oblong to ellip-
soid. 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. Inorescence 10–35 cm long, laxly 2–10-owered; rachis 2–15 cm long,
papillate-hairy; oral 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 reexed, 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 reexed over base;
lateral lobes linear-liform, 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).
Phenology. Flowering from June to August.
Habitat. ickets, alpine grasslands; 3200–4900 m elev.
Distribution. N and S Hengduan, also in NE Qinghai of China. (Fig. 20).
More illustrations. Wu et al. (2010, g. 195, 5–7).
Additional specimens examined. N HENGDUAN: Gonjo, 3200 m elev., Veg -
etation 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: Kang-
ding, 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
Taxonomic revision of Habenaria josephi group 131
Figure 18. Neotype of Habenaria tibetica Schltr. ex Limpricht X.L. Jiang 36260 (PE 00340847) (Pho-
tographed by T.R. Pandey).
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
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
132
Figure 19. Habenaria tibetica A habit B oral 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).
Figure 20. Distribution of Habenaria tibetica (black circles) in the Pan-Himalaya.
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
Taxonomic revision of Habenaria josephi group 133
world’s major herbaria and could have been destroyed during the Second World War at
B. Furthermore, we were unable to nd any other original material related to the species.
us, 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 desig-
nated 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). e 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
is research was supported by a grant from the National Natural Science Founda-
tion of China (No. 31670194). e Curators and stas 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. Specically,
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
typication, Dr. P. Ormerod for comments on the manuscript and Dr. H. Æ. Pedersen
(C) for providing some literature. anks 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) e Botany of the Afghan Delimitation Commission. In: e Trans-
actions 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 Evolu-
tion 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 Mahavidya-
laya, Malda, 81–90.
Tirtha Raj Pandey & Xiao-Hua Jin / PhytoKeys 175: 109–135 (2021)
134
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 classication of the orchid family. Dioscorides Press, Port-
land, 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, Cam-
pacci 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) e 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, Panzengeschichte und Panzengeographie 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. Habenari-
inae. 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. Reper-
torium Specierum Novarum Regni Vegetabilis. Beihefte 12: 1–515.
Lucksom SZ (2007) e 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 typication of some Habenaria species in
the Pan-Himalaya. Phytotaxa.
Pearce N, Cribb PJ (2002) e 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). Edin-
burgh Journal of Botany 58(1): 99–122. https://doi.org/10.1017/S0960428601000488
Taxonomic revision of Habenaria josephi group 135
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). De-
partment of Plant Resources, Kathmandu, Nepal.
Raskoti BB, Ale R (2019) Molecular phylogeny and morphology reveal a new epiphytic spe-
cies 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.1651-22 27.1924.tb14521.x
Stuessy TF (2009) Plant taxonomy: e 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.
iers B (2020) Index Herbariorum: A global directory of public herbaria and associated sta.
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 [Eds] (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 [Eds] (2010) Flora of China Illustrations (Vol. 25). Science Press,
Beijing and Missouri Botanical Garden Press, St. Louis.
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