PhytoKeys 128: 97–119 (2019)
A peer-reviewed open-access journal
Synopsis of the tribe Stipeae (Poaceae) in Nepal
doi: 10.3897/phytokeys.128.34637
97
RESEARCH ARTICLE
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Synopsis of the tribe Stipeae (Poaceae) in Nepal
Marcin Nobis1,2, Polina D. Gudkova2,3, Colin A. Pendry4
1 Institute of Botany, Faculty of Biology, Jagiellonian University, Gronostajowa 3 st., 30-387 Kraków, Poland 2 Research laboratory “Herbarium”, National Research Tomsk State University, 36 Lenin ave., 634050,
Tomsk, Russia 3 Faculty of Biology, Altai State University, 61 Lenin ave., 6560496, Barnaul, Russia 4 Royal
Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR, Scotland, UK
Corresponding author: Polina D. Gudkova (pdgudkova2017@yandex.ru)
Academic editor: Maria Vorontsova | Received 18 March 2019 | Accepted 17 July 2019 | Published 19 August 2019
Citation: Nobis M, Gudkova PD, Pendry CA (2019) Synopsis of the tribe Stipeae (Poaceae) in Nepal. PhytoKeys 128:
97–119. https://doi.org/10.3897/phytokeys.128.34637
Abstract
In Nepal the Stipeae consists of six genera: Achnatherum, Orthoraphium, Piptatherum, Ptilagrostis, Stipa,
Trikeraia, and 15 species. Two new combinations, Ptilagrostis duthiei (Hook. f.) M.Nobis & P.D.Gudkova
and Achnatherum staintonii (Bor) M.Nobis & P.D.Gudkova, are proposed, and new country records for
Stipa klimesii, Ptilagrostis dichotoma, Ptilagrostis concinna and Achnatherum jacquemontii are reported. The
records of Stipa roborowskyi, S. przewalskyi, S. capillata, S. consanguinea, S. mongholica, and S. sibirica,
previously thought to occur in Nepal were based on misidentifications, and these have been excluded from
the list of Nepalese Stipeae. We present keys for the identification of genera and species, and a checklist including information on nomenclatural types, regional and national distribution, and habitat. A lectotype
is designated for Stipa brandisii Mez.
Keywords
Checklist, Identification key, Nepal, Poaceae, Stipeae
Introduction
The tribe Stipeae L. (feather grasses) is composed of extratropical and high-mountain
grasses consisting of about 680 species distributed on all continents except Antarctica
(Tzvelev 1977; Barkworth 2007; Romaschenko et al. 2008, 2010, 2011, 2012; Soreng
et al. 2003, 2015, 2017, Nobis et al. 2019). It is an early divergent, highly specialized,
monophyletic lineage within the subfamily Pooideae Benth. The Stipeae are characterized by their single-flowered spikelets without rachilla extensions, lemmas with apical
awns where the awn is the result of fusion between the central and two lateral vascular
Copyright Marcin Nobis et al. 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.
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Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
traces, florets with three (rarely two) lodicules, and usually the palea is concealed by
the lemma (if the palea is exposed when the floret is closed, then the palea is coriaceous
(Roshevitz 1934; Tzvelev 1976; Freitag1985).
Although agrostologists have maintained a broad concept of the genus Stipa L.
since its description (Hitchcock 1935, Clayton and Renvoize 1986; Freitag 1985; Columbus et al. 2019 and others), recent molecular phylogenetic studies suggest that
ca. 34 genera should be recognized within the tribe (Hamasha et al. 2012; Nobis et
al. 2019; Peterson et al. 2019). However, the species composition of some genera still
requires further study.
All previous treatments of the Stipeae in Nepal have followed a broad generic
concept. The Annotated checklist of the flowering plants of Nepal (Press et al. 2000,
http://www.efloras.org/flora_page.aspx?flora_id=110) and the Flora of Mustang, Nepal
(Ohba et al. 2008) recognized 11 species of Stipa: Stipa breviflora Griseb., S. capillata L.,
S. consanguinea Trin. & Rupr., S. duthiei Hook. f., S. koelzii R.R.Stewart, S. mongholica
Turcz. ex Trin., S. przewalskyi Roshev., S. roborowskyi Roshev., S. roylei (Nees) Mez., S.
sibirica (L.) Lam. and S. staintonii Bor and three species of Oryzopsis: O. gracilis (Mez)
Pilg., O. lateralis (Regel) Stapf ex Hook. f. and- O. munroi Stapf ex Hook.f. Two other
species have recently been reported from Nepal, Ptilagrostis milleri (Noltie) M.Nobis
& A.Nobis (=S. milleri Noltie), and S. krylovii Roshev. (Nobis et al. 2015; Gudkova et
al. 2017a). Unfortunately, identification of Nepalese feather grasses is difficult due to
the lack of recent, comprehensive, regional taxonomic studies. The main goal of this
paper is to provide an identification key and checklist including information on types,
nomenclature, distribution, and habitat for all Nepalese species of Stipeae.
Materials and methods
Our treatment is based on herbarium specimens deposited in BM, E, GOET, K, KATH,
KRA, KUN, LE, M, NY, P (Thiers 2018). Each species is listed with complete nomenclatural and type information (the type specimens examined has exclamation mark after a
herbarium code) synonyms, habitat requirements, and Nepalese and general distribution.
The distribution within Nepal is given by District (Fig. 1). Elevation ranges and habitat requirements have been determined from herbarium specimen labels and from the literature.
Morphological analyses
Nineteen morphological characters scored for each taxon were included in the analysis
(Table 1). Each species was treated as an Operational Taxonomic Unit (OTU) following Sokal and Sneath (1963). Cluster analysis was performed on all OTUs to estimate
morphological similarities among the species. The similarities among OTUs were calculated using Gower’s general similarity coefficient. A cluster analysis (UPGMA) was
carried out using PAST software (Hammer et al. 2001).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
99
Figure 1. Districts of Nepal.
Table 1. Morphological characters and character states.
Characters
States
Macromorphological characters:
Length of anthecium (lemma + callus) [mm] mean length
Length of callus [mm]
mean length
Ratio lemma / palea
subequal (1); lemma longer than palea (2)
No. of awn geniculations
without geniculations (0); unigeniculate (1); bigeniculate (2)
Length of awn [mm]
mean length
Hairs on column [mm]
mean length
Hairs on seta [mm]
mean length
Length of glumes [mm]
mean length
Apex of glumes
twisted (1), straight (2)
Ligules of vegetative leaves [mm]
mean length
Ratio lower glume / upper glume
subequal (1); lower longer than upper (2)
Hard prickles at lemma apex
absent (1); present (2)
Micromorphological characters of the lemma epidermis:
Length of long cells
1–3(–5) times as long as wide (1); (4–)5–9(–11) times as long as wide
(2), as wide as long (3)
Side walls of long cells
not thickened (1), thickened (2)
Presence of hooks
frequent (more than 12 on area of 0.015 mm2) (1); sparse (less than
12 on area of 0.015 mm2) (2), absent (3)
Presence of silica cells
frequent (more than 20 per area of 0.015 mm2) (1); sparse (less
than 20 per area of 0.015 mm2) (2); rare (less than 5 per area of
0.015 mm2) (3)
Constriction of silica cells
with constrictions (1), without constrictions (2)
Shape of silica cells
ovate (1); elongated to ovate (2), elliptic or reniform (3)
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Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
Stipeae Dumort., Observ. Gramin. Belg. 83, 88, 134 (1824)
Plants perennial, usually cespitose, occasionally rhizomatous. Culms erect, unbranched.
Leaf blades flat or convolute, abaxial surface smooth, scabrous or pubescent, adaxial
surface prominently ribbed, with 0.05–1 mm long hairs. Ligules membranous. Inflorescence a dense or open panicle. Spikelets with one bisexual floret. Glumes clearly unequal to subequal, membranous, obtuse or acute, tapering into a long tip. Awns scabrid
to plumose, straight, uni- or bi-geniculate. Lemmas narrowly lanceolate, terete, usually
leathery, usually hairy. Callus rounded or acute to sharply pointed.
Results and discussion
Detailed analyses of macro- and micromorphological structures of the lemma epidermis of Nepalese species of Stipeae confirmed that they form two main clusters, one
with three subclusters (Fig. 2). The clusters correspond to the four lemma epidermal
patterns (LEP): Stipa-like, Ptilagrostis-like, Piptatherum-like and Achnatherum-like (Fig.
3). The taxa from cluster I belonging to Stipa have long cells and hooks on the lemma
epidermis in an ordered saw-like pattern (Romaschenko et al. 2012; Fig. 3a). Within
subcluster A of cluster II, there are three genera, Ptilagrostis, Trikeraia and Orthoraphium (Fig. 2), that have LEPs dominated by elongated basal cells, frequent silica bodies
and cork cells (Fig. 3b–d, h). However, the presence of deflexed, hard prickles in the
case of Orthoraphium roylei (Fig. 3d), as well as 2–3 mm long awn-like lemma lobes
Figure 2. Cluster analysis (UPGMA method of classification and Gower’s general similarity coefficient)
performed on 16 qualitative morphological characters for all of the Nepalese members from the tribe Stipeae.
Synopsis of the tribe Stipeae (Poaceae) in Nepal
101
Figure 3. Lemma epidermal patterns (LEPs) of Old World Stipeae: a Stipa breviflora [Kyrgyzstan, near
Issyk-Kul Lake, M. Nobis (KRA)] b Ptilagrostis concinna [India, Ladakh, Himalayas, L. Klimeš (KRA)]
c Ptilagrostis duthiei [India, Himalayas, J.F. Duthie 3585 (LE) d Orthoraphium roylei [India, Himalayas,
J.F. Duthie 3568 (LE)] e Trikeraia hookeri [China, Tibet (PE 718306)] f Piptatherum munroi [Nepal,
Solukhumbu, M.F. Watson et al. DNEP3 AX33 (E)] g Achnatherum brandisii [India, Kashmir, R.R. Stewart
18120 (NY)] h Achnatherum staintonii [Nepal, Mustang, M.A. Farille 81-340 (E)]. Annotations: b – basal
cells, c – cork cells, sb – silica bodies, m – macro-hairs.
Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
102
in the case of Trikeraia hookeri (Fig. 3e) are unique characters which distinguish them
from other members of the subcluster. Subcluster B comprises species from the genus
Piptatherum. These species differ from those in subcluster A in their extremely short
callus, less numerous and rounded silica bodies on the lemma surface (Fig. 3f ). Taxa in
subcluster C, all of which belong to the genus Achnatherum, have a maize-like type of
LEP (Romaschenko et al. 2012), characterized by numerous silica bodies and very short
basal cells (Fig. 3g–h). All of these species have lemmas distinctly longer than paleas.
Key to genera
1
–
2
–
3
–
4
–
5
–
Lemma with deflexed (retrorse), apical prickles .................... Orthoraphium
Lemma lacking deflexed, apical prickles ......................................................2
Lemma lobes awn-like, 2–3 mm long, setaceous ............................Trikeraia
Lemma without awn-like lobes, lobes (if present) flat and less than 1 mm
long ............................................................................................................3
Awns straight, scabrous. Anthecium usually dorsally compressed. Callus up
to 0.3 mm long ........................................................................ Piptatherum
Awns geniculate, scabrous or variously pilose. Anthecium not compressed or
laterally compressed. Callus longer than 0.3 mm ........................................4
Callus longer than 0.9 mm. Lemma epidermis with numerous minute hooks
(visible under high magnification) ....................................................... Stipa
Callus up to 0.8 mm long. Lemma epidermis smooth or rarely with infrequent minute hooks ....................................................................................5
Lower segment of awn pilose, with hairs over 0.3 mm long. Surface of lemma
epidermis covered with elongated basal cells (4–11 times longer than wider)
and occasional, 1–3-constricted silica bodies ...............................Ptilagrostis
Lower segment of awn scabrous, with hairs up to 0.1 mm long. Surface of
lemma epidermis covered with rounded or once-constricted silica, underlying
cells as wide as long or wider than longer ................................ Achnatherum
Orthoraphium Nees, Proc. Linn. Soc. Lond. 1: 94 (1841)
Type. Orthoraphium roylei Nees.
Orthoraphium roylei Nees, Proc. Linn. Soc. Lond. 1: 94 (1841).
≡ Stipa orthoraphium Steudel, Syn. Pl. Glumac. 1: 131 (1855) nom. superfl.;
≡ Stipa roylei (Nees) Duthie, Grasses North-Western India 27 (1883);
≡ Stipa roylei (Nees) Mez, Repert. Spec. Nov. Regni Veg. 17: 207 (1921).
Type. (India, W Himalaya) Kadarkanal, Royle (holotype: LIV).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
103
General distribution. Himalayas: Bhutan, S China, Ladakh, N Myanmar, N India, Nepal (Bor 1960; Freitag 1985; Wu and Phillips 2006).
Distribution in Nepal. Baglung, Bajura, Darchula, Dolkha, Humla, Jumla, Lalitpur, Mugu, Myagdi, Ramechhap, Rasuwa, Rukum, Sankhuwasabha, Solukhumbu,
Taplejung.
Habitat. Alpine meadows, Rhododendron scrub, oak-laurel forests.
Altitudinal range. 2200–4000 m.
Selected Nepalese specimens studied. Baglung: Dhorpatan, Vallee de Dhorpatan du cote est, 28°29'18"N, 83°4'1"E, 2800–3350 m, 16 Aug. 1981, M. Farille
81-168 (E). Bajura: Birseni – Porakya, 2250 m, 12 Aug. 1991, K.R. Rajbhandari
14829 (KATH). Darchula: Dopakhe, Dandar – Dopakhe, rocky slope, 2200 m, 28
Aug. 1980, K.R. Rajbhandari & K.J. Malla 5684 (KATH). Dolkha: Gyalche Kharka – Thang Dingma, 3100 m, 1 Sep. 1983, K.R. Rajbhandari 9744 (KATH); Bhitte
Kharka – Patlo Pokhari, shady place in forest, 3700 m, 12 Sep. 1983, K.R. Rajbhandari
10123 (KATH); Bhitte Kharka – Patlo Pokhari, 3800 m, 12 Sep. 1983, K.R. Rajbhandari 10164 (KATH). Humla: Tambe Danda, 2750 m, Q. semecarpifolia forest,
15 Aug. 1977, K.R. Rajbhandari & B. Roy 2268 (KATH). Jumla: Jumla, 29°16'31"N,
82°11'0"E, H. Tabata, K.R. Rajbhandari & K. Tsuchiya 9327 (KATH). Lalitpur:
Phulchoki, 27°34'14"N, 85°24'4"E, 2600 m, 14 Oct. 1990, K.R. Rajbhandari 4122
(KATH). Mugu: Chankheli Lagna, 29°38'21"N, 82°6'52"E, 3500 m, 9 Aug. 1979,
K.R. Rajbhandari & B. Roy 4445 (KATH). Myagdi: Gurjakhani, North-West of Gurjakhani, 28°36'N, 83°13'E, 3480–3490 m, 31 Jul. 1954, J.D.A. Stainton, W.R. Sykes
& L.H.J. Williams 3685 (BM, E, K). Ramechhap: Khola Kharka – Thare, 3600 m,
22 Jul. 1985, H. Ohba et al. 60583 (KATH); Bhandar – Deorali – Khasrubus – Shivalaya, 2600 m, H. Ohba et al. 62276 (KATH). Rasuwa: Rupchet Kharka - Balchagam,
3200 m, 16 Aug. 1994, K.R. Rajbhandari 17910 (KATH); Laurebina Yak, 28°5'32"N,
85°22'52"E, 3450 m, 30 Jul. 1995, T. Hoshino et al. 9537289 (KATH). Rukum: Sing
Khola – Farkama 3636 m, 18 Sep. 1976, H. Tabata, K.R. Rajbhandari & K. Tsuchiya
3720 (KATH). Sankhuwasabha: Hile Ghot, 27°24'N, 87°26'E, 3500 m, 20 Aug.
1972, J.F. Dobremez 1594 (BM, E). Solu Khumbu: Imja Khola Valley, Omoga, sandy
clay, floor of north-west/south-east river valley, west facing slope, mossy slope in shade,
27°50'38"N, 86°47'10"E, 3600–2300 m, 26 Sep. 2005, DNEP3 AX131 (E, KATH);
Chaurikharka, Q.semecarpifolia forest, mossy slope, 27°41'46"N, 86°43'31"E, 2729
m, 30 Sep. 2005, DNEP3 BY229 (E, KATH); Dudh Kund – Samakang, 27°41'N,
86°50'E, 3500 m, 24 Aug. 1995, F. Miyamoto et al. 9592410 (E); Samakang Kharka
27°41'N, 86°50'E, 3500 m, 24 Aug. 1995, F. Miyamoto et al. 9592410 (KATH); Beni
Kharka, 3100 m, 3 Sep. 1985, H. Ohba et al. 62035 (KATH); Beni Kharka, 3600–
2300 m, 2 Sep. 1985, H. Ohba et al. 61907 (KATH); Loding, 27°32'N, 86°32'E,
2700 m, 5 Sep. 1985, H. Ohba et al. 62108 (KATH); 62155; Pike Bhanjyang, 3700 m,
6 Sep. 1985, H. Ohba et al. 62155 (KATH); Pike Dongshar, Rhododendron campanulatum thicket, 27°30'N, 86°27'E, 2300–3600 m, 9 Sep. 1985, H. Ohba et al. 62192
(KATH); Rangdu Kharka, 27°8'N, 86°48'E 3550 m, 9 Aug. 1997, K.R. Rajbhandari
9740203 (KATH); Tangnag – Mosom Kharka, in forest on mossy ground, 3700 m, 21
Aug. 1997, K.R. Rajbhandari 9740472 (KATH); Beni – Tokchardingma, Basa Valley,
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27°32'59"N, 86°35'13"E, 3750 m, 2 Aug. 1995, K. Tsuchiya 40841 (KATH); Sengephuk, Beni VDC, 3580 m, 29 Aug. 1995, K. Tsuchiya 41782 (KATH); Luminasa,
Basa Valley, Beni, 3780 m, 8 Aug. 1995, K. Tsuchiya 42188 (KATH). Taplejung: forest ridge et Manedhanjang, N end of Milke Danda Ridge, Rhododendron / Bamboo
forest, caespitose grass, 27°27'N, 87°28'E, 3340 m, 22 Oct. 1991, D.G. Long et al.
966 (E, KATH); Topke Gola, Arun-Tamur watershed S of Topke Gola, on slopes,
27°38'25"N, 87°34'59"E, 3940 m, 13 Sep. 1956, J.D.A. Stainton 1728 (BM, E).
Stipa L., Sp. Pl. 1: 78 (1753)
Type. Stipa pennata L.
Key to the genus Stipa
1
–
2
–
3
–
Upper part of awn (seta) scabrous, with hairs up to 0.4 mm long ...............2
Upper part of awn (seta) pilose, with hairs over 0.4 mm long .....................3
Upper part of awn and tips of glumes spirally twisted ...............S. capillacea
Upper part of awn and tips of glumes not twisted ........................ S. krylovii
Ligules of vegetative shoots up to 0.3 mm long. Awn column with 0.6–
0.7 mm long hairs..................................................................... S. breviflora
Ligules of vegetative shoots over 2 mm long. Awn column with 1.5–2.5 mm
long hairs ..................................................................................... S. klimesii
Stipa breviflora Griseb., Nachr. Ges. Wiss. Göttingen, Math.-Phys. Kl. 3:82
(1868).
= S. aliciae Kanitz, Növényt. Gyujtesek Eredm. Grof Szechenyi Bela Keletazsiai Utjabol 61, t. 7 (1891).
Type. (China) Tibet, Gnari (Nari) Khorsum, Schlagentweit 7105 (holotype GOET!,
isotype LE!).
General distribution. China, Kyrgyzstan, N India, Mongolia, Nepal (Grubov
1982; Tzvelev 1968, 1976; Freitag 1985; Wu and Phillips 2006).
Distribution in Nepal. Mustang.
Habitat. High altitude steppes, scree.
Altitudinal range. 2750–3600 m.
Selected specimens studied. Mustang: Entre Jomsom et Kagbheni, dans la
steppe aride a Caragana gerardiana et C. brevispina (limite), 28°46'51"N, 83°43'27"E,
2750 m, 17 Sep. 1981, M.A. Farille 81-362 (E); Muktinath, on open slopes near cultivations, 28°48'58"N, 83°51'47"E, 3640 m, 8 Jun 1954, J.D.A. Stainton, W.R. Sykes
& L.H.J Williams 5647 (E, K).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
105
Stipa klimesii M.Nobis, Phytotaxa 174(3): 166–168 [174–176] (2014).
= Stipa basiplumosa Munro ex Hook. f., Fl. Brit. India 7(22): 229 (1896) var. longearistata Munro ex Hook. f., Fl. Brit. India 7(22): 229 (1896).
Type. India, NW India, Jammu and Kashmir State, Ladakh, Indus Vy: Zhung (Leh),
Ganglas – upper part, springs with drinking water, 3880–4000 m, 30 Jul. 2001,
34°12.3'N, 77°36.8'E, L. Klimeš 1155, 1156 (holotype KRA!, isotype PRA!).
General distribution. Bhutan, China (Tibet), India (Ladakh, Sikkim), Nepal, Pakistan, (Nobis et al. 2014, 2015).
Distribution in Nepal. Mustang.
Habitat. High mountain steppes and alpine mats, among subalpine shrubs and
on rocky ledges.
Altitudinal range. 3500–5000 m.
Notes. These specimens were previously identified as S. roborowskyi, but this species does not occur in Nepal. This species differs from S. klimesii in having shorter
ligules on the vegetative shoots [0.5–1.5(–2) vs. (2–)3.5–7.5(–9) mm], shorter anthecium [(6–)6.5–7.5(–7.7) vs. (7–)8.3–9.5(–10) mm] and shorter hairs on seta
[(0.3)0.5–1.1(–1.4) vs. (1–)1.3–2(–2.3) mm long, respectively].
Selected specimens studied. Mustang: Damodar, on dry sandy slope, 29°11'N,
83°58'E, 28 Jul. 1979, P.R. Shakya, S.R. Adhikari & K.R. Amatya 5111 (KATH); Kunda, 28°59'9"N, 84°9'26"E, 4720–4740 m, 11 Aug. 2001, S. Noshiro, M. Amano & T.
Kurosawa 20104179 (KATH).
Stipa krylovii Roshev., Izv. Glavn. Bot. Sada S.S.S.R. 28: 379 (1929).
≡ S. sareptana subsp. krylovii (Roshev.) Cui, Fl. Xinjiang. 6: 299 (1996);
≡ S. sareptana var. krylovii (Roshev.) Kuo & Sun, Fl. Reipubl. Popularis Sin. 9(3): 275,
pl. 65, f. 37–41 (1987).
= S. capillata var. coronata Roshev., Fl. Aziat. Ross. 1(12): 168, pl. 8, 8b (1916);
= S. densiflora P.A.Smirn., Repert. Spec. Nov. Regni Veg. 26: 265 (1929) hom. illeg.
non Hughes;
= S. densa P.A.Smirn., Del. sem. Hort. Bot. Univ. Mosquensis 15 (1930);
= S. decipiens P.A.Smirn., Ucen. Zap. Moskovsk. Gosud. Univ. 2: 338 (1934).
Type. Selenginskaya Dauriya, gory mezhdu Temnikom i Dzhidoi, yugo-zapadnaya
chast khr. Borgoiskogo, na sklonakh so stepnoi rastitelnostyu, 28 Jul. 1912, V. Smirnov
524 (lectotype: LE!, designated by Tzvelev 1976).
General distribution. Widely distributed throughout Central Asia (Eastern Kazakhstan, Russia (Siberia: Altai, Khakasiya, Tuva, South Krasnoyarsk, Irkutsk, Buryatiya,
Chita, South Yakutia); China (Gansu, Hebei, Nei Mongol, Ningxia, Qinghai, Shanxi,
Xinjiang, Xizang), Mongolia, eastern Kyrgyzstan, Tajikistan (Pamir), North India, Nepal.
(Tzvelev 1968, 1976; Freitag 1985; Wu and Phillips 2006; Gudkova et al. 2017a, 2017b).
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Distribution in Nepal. Mustang (Gudkova et al. 2017a).
Habitat. High mountain semi-desert.
Altitude range. 3900–4000 m.
Notes. These specimens were previously identified as S. capillata, but this species
does not occur in Nepal. Stipa krylovii differs from S. capillata mainly in having a ring
of hairs at the top of the lemma.
Selected specimens studied. Mustang: s.loc., on dry sandy ground, 29°14'N,
83°52'E, 13000 ft, 3 Aug. 1954, Stainton, Sykes, Williams 2161 (E, K, BM).
Stipa capillacea Keng, Sunyatsenia 6(2): 100, pl. 15 (1941).
= S. koelzii R.R.Stewart, Brittonia 5: 441 (1945).
Type. Open grass land in rear of Shaowusze Agricultural Station, Taining district, Sikang province, 22 Jul. 1940, K.L. Chü 7449 (holotype: N, isotype: PE!).
General distribution. Bhutan, S China, N India, Nepal, Pakistan, (Freitag 1985;
Noltie 2000; Wu and Phillips 2006).
Distribution in Nepal. Mustang, Rasuwa, Solukhumbu.
Habitat. Alpine meadows.
Altitudinal range. 2800–4100 m.
Note. These specimens were previously identified as Stipa consanguinea, but this
species does not occur in Nepal. Stipa capillacea differs from other species of the genus
in awns twisted together at top of panicle.
Selected specimens studied. Mustang: Kali Gandaki, Thulo Bugin, ESE Lete, S- facing steep slope, 28°38'4"N, 83°36'20"E, 2820 m, 10 Oct. 1977, G. Miehe 762b (BM); at
edge of field, 13000 ft, 3 Aug. 1954, J.D.A. Stainton, W.R. Sykes & L.H.J. Williams 2150
(BM, K); Rasuwa: Langtang, 28°13'N, 85°3'E, 3900 m, 7 Aug. 1970 J.F. Dobremez 522
(BM, E); Kyanjin Gompa, 28°12'42"N, 85°34'1"E 11500–12500 ft, 10 Aug. 1969, A.
Richard 103 (BM); Ganesh Himal, 28°20'N, 85°10'E, Apr. 1975 - May 1975, B. Yon 252
(E). Solukhumbu: Orsho, east facing slope open ground, grazed area with scattered Juniperus indica, 27°52'15"N, 86°48'44"E, 4100 m, 19 Sep. 2005, DNEP3 AX83 (E, KATH).
Ptilagrostis Griseb., Fl. Ross. 4(13): 447 (1852)
Type. Ptilagrostis mongholica (Turcz. ex Trin.) Griseb.
Key to the genus Ptilagrostis
1
–
Awn with 0.3–0.5 mm long hairs on column. Seta scabrous .........P. duthiei
Awn variously pilose on column, with hairs over 1 mm long. Seta with 0.5–
1.5 mm long hairs.......................................................................................2
Synopsis of the tribe Stipeae (Poaceae) in Nepal
2
–
3
–
107
Glumes, lemma and palea distinctly unequal (lower glume 1.5–3.5 mm longer
than the upper and lemma 1–2.5 mm longer than palea) .........P. yadongensis
Glumes, lemma and palea equal or only slightly unequal ............................3
Panicles open, 3–5 cm wide, branches up to 6 cm long, spreading ................
................................................................................................ P. dichotoma
Panicles compressed, 0.7–2 cm wide, branches 0.3–2.8 cm long, suberect or
narrowly ascending .....................................................................P. concinna
Ptilagrostis duthiei (Hook.f.) M.Nobis & P.D.Gudkova, comb. nov.
urn:lsid:ipni.org:names:77200949-1
Stipa duthiei Hook.f., Fl. Brit. India 7: 232, 1896. (Basionym).
≡ Achnatherum duthiei (Hook.f.) Kuo & Lu, Fl. Reipubl. Popularis Sin. 9(3): 322, pl.
80, f. 9–14 (1987).
Type. [India] Tehri Garwhal, Lekhus, below Srikanta, 12000–13000ft, 11 Aug. 1853,
Duthie 273 (holotype K 32097!).
General distribution. China, N India, Kashmir, Nepal (Freitag 1985; Wu and
Phillips 2006).
Distribution in Nepal. Myagdi.
Habitat. Mountain shrublands.
Altitudinal range. 3400–3800 m.
Selected specimens studied. Myagdi: North of Barse, among dwarf Rhododendron, 3940 m, [28°35'N, 83°11'E], 14 Aug. 1954, J.D.A. Stainton, W.R. Sykes & L.H.J.
Williams 3844 (E 690624).
Ptilagrostis dichotoma Keng ex Tzvelev, Rast. Tsentr. Azii 4: 43 (1968).
Type. China, Kansu and Tsinghai border [in regione opp. Labrang], I.C. Wu 478 (holotype N, isotype LE!).
General distribution. Bhutan, Birma, China (Tibet), N India, Nepal (Wu and
Phillips 2006; Nobis et al. unpbl).
Distribution in Nepal. Bajhang, Bajura, Dolakha, Jumla, Mustang, Myagdi, Ramechhap, Rasuwa, Rukum, Solukhumbu.
Habitat. Alpine meadows, grassy mountain slopes.
Altitudinal range. 3300–5000 m.
Note. These specimens were previously identified as Ptilagrostis mongholica [=Stipa
mongholica] (Bor 1960; Freitag 1985), but the two species are easily distinguished as
P. dichotoma has a tuft of short hairs at the apex of anthers (glabrous in P. mongholica).
They are disjunctly distributed with P. dichotoma found in the mountains of southerncentral Asia while P. mongholica occurs mainly in the mountains of northern-central
Asia; (Tzvelev 1968; Wu and Phillips 2006).
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Selected specimens studied. Bajhang: Manane Lekh 29°36'45"N, 80°59'35"E,
3830 m, 14 Jul. 2009, Bajhang 09 20917078 (E, KATH); Saipal, 29°57'51"N,
81°13'6"E, 3909 m, H. Tabata, K.R. Rajbhandari & K. Tsuchiya 1808 (KATH); Saipal,
29°57'51"N, 81°13'6"E, 30 Jul. 1976, H. Tabata, K.R. Rajbhandari & K. Tsuchiya 4024
(KATH); Saipal, 29°57'51"N, 81°13'6"E, 31 Jul. 1976, H. Tabata, K.R. Rajbhandari
& K. Tsuchiya 4028 (KATH). Bajura: Chauki Lekh, 29°35'34"N, 81°38'5"E, 4427 m,
15 Aug. 2017, BSH C42; Chauki Lekh, 29°37'16"N, 81°34'30"E, 4427 m, 16 Aug.
2017, BSH C52. Dolakha: Dudh Kunda, 4550 m, 5 Sep. 1983, K.R. Rajbhandari
9915 (KATH). Dolpa, Nahure, 14,000 ft, 24 Jun. 1952, Polunin, O.V., Sykes, W.R.
& Williams, L.H.J. 1434. Jumla: Maharigaon, 15,000 ft, 20 Jul. 1952, Polunin, O.V.,
Sykes, W.R. & Williams, L.H.J. 226. Mustang: Muktinath Himal, Muktinath Range,
28°44'37"N, 83°53'14"E, 480–5000 m, 18 Sep. 1981, M.A. Farille 81-415 (E); Muktinath Himal, Muktinath Range, hab dans la praire alpine, 28°44'37"N, 83°53'14"E,
4800–5000 m, 18 Sep. 1981, M.A. Farille 81-415 (E); Kaisang – Omang Kharka
3800 m, 1 Aug. 1996, K.R. Rajbhandari 9672251 (KATH); Muktinath, 28°48'58"N,
83°51'47"E, 4180 m, 12 Oct. 1976, H. Tabata, K.R. Rajbhandari, K. Tsuchiya & Y. Konno 6310 (KATH); Muktinath, alpine grassland, 28°48'58"N, 83°51'47"E, 4120 m, 12
Oct. 1976, H. Tabata, K.R. Rajbhandari, K. Tsuchiya & Y. Konno 6342 (KATH). Myagdi: 28°32'0"N, 83°13'0"E, 3360 m, 20 Sep. 1996, M. Mikage, R. Hirano, A. Takahashi & K. Yonekura 9682900 (KATH). Ramechhap: Thare Og, 27°45'N, 86°28'E
24 Jul. 1985, H. Ohba, M. Wakabayashi, M. Suzuki, N. Kurosaki, K.R. Rajbhandari &
S.K. Wu 60657 (KATH). Rasuwa: Gosainkund, 28°4'58"N, 85°24'51"E, 4300 m, 27
Jul. 1995, T. Hoshino, K. Dan, H. Koba, Y. Omori, C.P. Rauniyar, M. Sato, P. Shrestha
& S. Takatsuki 9537190 (KATH); Gosainkund, 4350 m, 28°4'58"N, 85°24'51"E, 27
Jul. 1995, T. Hoshino, K. Dan, H. Koba, Y. Omori, C.P. Rauniyar, M. Sato, P. Shrestha
& S. Takatsuki 9537218 (KATH); Gosainkund, 28°4'58"N, 85°24'51"E, 4300 m, 28
Jul. 1995, T. Hoshino, K. Dan, H. Koba, Y. Omori, C.P. Rauniyar, M. Sato, P. Shrestha
& S. Takatsuki 9537221, 9537–222 (KATH). Rukum: Chalikhe Pahar, near Chalike
Pahar, exposed south facing slopes, 28°40'N, 83°4'E, 4240 m, 17 Jun 1954, J.D.A.
Stainton, W.R. Sykes & L.H.J. Williams 3163 (E). Solukhumbu: Seto Pokhari (4810m)
– Chhomalang Base Camp (4495), 27°47'N, 86°57'E, 4810 m, 17 Aug. 1995, F.
Miyamoto, M. Amano, H. Ikeda, C.M. Joshi, K. Arai & T. Komatsu 9592313 (E); Beni,
alpine meadow, 27°32'59"N, 86°35'13"E 4600 m, K. Tsuchiya 42693 (KATH).
Ptilagrostis yadongensis Keng & Tang, J. SouthW. Agric. Univ. 4: 44 (1985).
≡ Ptilagrostis macrospicula Cai, Acta Bot. Boreal.-Occid. Sin. 23(11): 2018 (2003).
superfl. name.
= Stipa milleri Noltie, Edinburgh J. Bot. 56(2): 288 (1999); ≡ Ptilagrostis milleri (Noltie) M.Nobis & A.Nobis, Nordic J. Bot. 31: 623 (2013).
Type. China. Xizang: Yadong, 14 Sep. 1974, Qinghai-Xizang Exped. 74–2496 (lapsus
calami as 74–2469; holotype HNWP, isotype PE).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
109
General distribution. Bhutan, China (Tibet), Nepal (Noltie 1999; Nobis and
Nobis 2013; Nobis et al. 2015; Zhang et al. 2016).
Distribution in Nepal. Bajhang, Dolkha, Rasuwa, Solukhumbu.
Habitat. Alpine meadows, moist grassy places, under shrubs, swampy places, Kobresia moors.
Altitudinal range. 3600–4600 m.
Note. These specimens were previously identified as P. concinna which also occurs
in Nepal and which can be distinguished by its subequal glumes, lemma and palea. It
has also been confused with Ptilagrostis bhutanica (Noltie) M.Nobis (basionym: Stipa
bhutanica Noltie 1999: 289; Nobis et al. 2016), from Bhutan and China. However,
these two taxa differ in the upper part of the awn which is scabrous in P. bhutanica and
shortly pilose (with hairs over 0.5 mm long) in P. yadongensis.
Selected specimens studied. Bajhang: Saipal Aletsoura, 29°57'51"N, 81°13'6"E,
4333 m, 31 Jul. 1976, H. Tabata, K.R. Rajbhandari & K. Tsuchiya 1941 (KATH).
Dolkha: Rolwaling Dudh Kunda, 4520 m, 15 Jul. 1975, P.R. Shakya, K.R. Rajbhandari & H.K. Saiju 75/2978 (KATH). Rasuwa: Ya La, 3600 m, 29 July 1972,
A.Maire, AMA 250 (E); Upper Langtang, 4600 m, 30 Sep. 1986, G. Miehe 13090
(KATH). Solukhumbu, Chola Tsho, north side of lake, SE facing slope, rocks and
sand near lakeside, Juniperus indica dwarf scrubland with Rhododedron setosum
and Potentilla fruticosa, 27°55'18"N, 86°47'50"E, 4500 m, 21 Sep. 2005, DNEP3
AX98 (E, KATH).
Ptilagrostis concinna (Hook. f.) Roshev., Fl. URSS 2: 75 (1934).
Stipa concinna Hook. f., Fl. Brit. India 7(22): 230 (1897) (Basionym).
Type. Sikkim-Himalaya, Tibetan region, 14000–16000ft, 1861, Hooker (holotype K!,
isotypes G, GOET!, LE 9267!).
General distribution. Himalayas: China (Tibet), India (Ladakh and Sikkim), Nepal (Freitag 1985; Wu and Phillips 2006).
Distribution in Nepal. Solukhumbu, Mustang.
Habitat. Alpine meadows, moist grassy places, under shrubs, swampy places, Kobresia moors.
Altitudinal range. 4400–5300 m.
Selected specimens studied. Solukhumbu: Seto Pokhari, 27°47'N, 86°55'E,
4495–4810 m, 17 Aug. 1995, F. Miyamoto, M. Amano, H. Ikeda, C.M. Joshi, K. Arai &
T. Komatsu 9592313 (KATH). Mustang: Thorung La, 5200–5300 m, 19 Sep. 1981,
M.A. Farille 81-434 (E 189114).
Achnatherum P.Beauv., Ess. Agrostogr.: 19, 146, pl. 6, f. 7 (1812)
Type. Achnatherum calamagrostis (L.) P.Beauv.
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Key to the genus Achnatherum
1
–
2
–
Glumes distinctly unequal. Lemma apex with ring of hairs over 3 mm long.
Callus 0.5–0.7 mm long, acute at the apex ...............................A. staintonii
Glumes equal or almost so. Lemma apex with ring of hairs up to 2 mm long.
Callus up to 0.5 mm long, rounded at the apex ..........................................2
Lemma and palea clearly unequal. Leaves filiform, inrolled. Culms up to 45
cm long. Panicle with very short branches.............................A. jaquemontii
Lemma and palea subequal. Leaves flat. Culms over 60 cm long. Panicle with
widely spreading branches..........................................................A. brandisii
Achnatherum brandisii (Mez) Z.L.Wu, Acta Phytotax. Sin. 34: 154 (1996).
Stipa brandisii Mez, Repert. Spec. Nov. Regni Veg. 17(13–18): 207 (1921) (Basionym).
= Stipa subeffusa Ohwi, Acta Phytotax. Geobot. 17: 15 (1957).
Type. [India] N. W. Himalaya, Kulla, Oct. 1876, Brandis 1005 (lectotype K 32092!
selected and labeled by H. Freitag on 15 March 1984 but designated here).
General distribution. Afghanistan, Bhutan, China, NW India, Nepal, Pakistan
(Freitag 1985; Wu and Phillips 2006).
Distribution in Nepal. Dolpa, Manang, Mustang.
Habitat. Open dry slopes, among shrubs and in Bamboo (Sinarundinaria sp.) thicket.
Altitudinal range. 2400–4000 m.
Note. Stipa brandisii was described by Mez (1921) based on a specimen housed at
B but destroyed during the Second World War. In his original description of the species
Mez (1921) reported that the species had been collected in ‘Western-Himalaya, Kulla
([by] Brandis)’, but did not provide further information about the date of specimen collection, number and place where it was housed. In the absence of the original material, the
specimen at K (http://apps.kew.org/herbcat/getImage.do?imageBarcode=K000032092)
was selected as lectotype by H. Freitag in 1984 and subsequently cited as the holotype
(Freitag 1985). We designate this here as the lectotype.
Selected specimens studied. Dolpa: Above Chong, near Tibrikot Growing,
among shrubs on open slopes, 29°1'40"N, 82°46'22"E, 2580 m, 11 Sep. 1952,
O.V. Polunin, W.R. Sykes & L.H.J. Williams 3314 (E). Manang: Humde, Bhraka,
open place, 3400 m, Aug. 1983, K.R. Rajbhandari 8786 (KATH); Boraga, Entre
Braga et Ghyaru, 28°39'24"N, 84°2'22"E, 3500 m, 22 Sep. 1981, M.A. Farille
81-486 (E); Boraga, Entre Braga et Ghyaru, bois ouvert xerophile, 28°39'24"N,
84°2'22"E 3500 m, 22 Sep. 1981, M.A. Farille 81-491 (E); Boraga Annapurna
Himal, Manang, Annapurna III, north-slope above Braga, on pastures, 28°39'24"N,
84°2'22"E, 3850 m, 14 Oct. 1969, T. Wraber, 36404(502) (BM). Mustang: Versant
de Muktinath Range, faisant face a Jomsom, 50 m au dessus du village Rochers [on
Synopsis of the tribe Stipeae (Poaceae) in Nepal
111
rocks], environnement rocheux, mais dans les touffes d’epineux oub d’Artemisia,
28°46'51"N, 83°43'27"E, 2750 m, 16 Sep. 1981, M.A. Farille 81-358 (E); Ommang 28°44'N, 83°45'E, 3600 m, 31 Jul. 1996, T. Hoshino, M. Amano, H. Koba, N.
Miyoshi, K.R. Rajbhandari, M. Sato, P. Shrestha & S. Takatsuki 9662100 (KATH);
Jomsom, 28°46'51"N, 83°43'27"E, 3350 m, 31 Jul. 1996, T. Hoshino, M. Amano, H. Koba, N. Miyoshi, K.R. Rajbhandari, M. Sato, P. Shrestha & S. Takatsuki
9670079 (KATH); Kali Gandaki, Tangdung-Khola, S-facing, wind-blown slope,
2490–2480 m, Aug. 1977, G. Miehe (BM); Cha Lungpa, NE-facing slope, alpine
pastures 3940 m, 27 July 1977, G. Miehe 351b (BM); NW of Tukche, valley of
Yamkin Khola, in Bamboo (Sinarundinaria sp.) thicket, 28°41'15"N, 83°37'35"E,
2840 m, 20 Sep. 1995, M. Mikage & K. Yonekura 9552331 (KATH); Tukucha (Kali
Gandaki), 3180 m, [28°42'33"N, 83°38'37"E], 21 Aug. 1954, J.D.A. Stainton, W.R.
Sykes & L.H.J. Williams 7363 (BM, E 619028); Tukucha, Kali Gandaki, amongst
hillside shrubs, 28°42'33"N, 83°38'37"E, 3030 m, 12 Sep. 1954, J.D.A. Stainton,
W.R. Sykes & L.H.J. Williams 7813 (E); Chimgaon (N of Tukucha) Kali Gandaki, on
dry slopes, 28°43'38"N, 83°40'45"E, 2880 m, 14 Sep. 1954, J.D.A. Stainton, W.R.
Sykes & L.H.J. Williams 9887 (BM).
Achnatherum jacquemontii (Jaub. & Spach) P.C.Kuo & S.L.Lu, Fl. Reipubl. Popularis Sin. 9(3): 323, pl. 80, f. 15–19 (1987)
Stipa jacquemontii Jaub. & Spach, Ill. Pl. Orient. 4: 60, pl. 339 (1851) (Basionym).
≡ Lasiagrostis jacquemontii (Jaubert & Spach) Munro ex Boiss., Fl. Orient. 5:
506 (1884);
≡ Lasiagrostis jacquemontii (Jaub. & Spach) Munro ex Aitch., J. Linn. Soc., Bot. 18:
107 (1880).
= Stipa jacquemontii Jaub. & Spach subsp. chuzomica Noltie, Edinburgh J. Bot. 56(2):
290, f. 1Q–U (1999).
Type. [India] ad ruped in excelsis Emodi Cashemyrianim 2750 m, 1831, Jacquemont
994, (holotype P, isotype K!).
General distribution. E Afghanistan, Bhutan, China (Tibet), NW India, Nepal,
Pakistan (Freitag 1985; Noltie 2000; Wu and Phillips 2006).
Distribution in Nepal. Mustang.
Habitat. Dry mountain slopes, especially in rock crevices.
Altitudinal range. 2500–3000.
Selected specimens studied. Mustang: Marpha, pentes rocheues arides,
28°45'11"N, 83°41'28"E, 2650 m, 16 Sep. 1981, M. Farille 81-340 (E); Versant
de Muktinath Range, faisant face a Jamson, 100 m au dessus du Village Rochers,
2800 m, 16 Sep. 1981, M.A. Farille 81-347 (E 188712); Barsumg Khola, on a dry
cliff, 28°52'N, 83°16'E, 10000 ft, 18 Jul. 1963, J.D.A. Stainton 4417 (E 00619022).
112
Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
Achnatherum staintonii (Bor) M.Nobis & P.D.Gudkova, comb. nov.
urn:lsid:ipni.org:names:77200950-1
Stipa staintonii Bor, Bull. Bot. Surv. India 7: 133 (1965) (Basionym).
≡ Stipella staintonii (Bor) Röser & Hamasha, Pl. Syst. Evol. 298: 365 (2012), nom. inval.;
≡ Stipellula staintonii (Bor) Röser & H.R. Hamasha, Schlechtendalia 24: 92 (2012).
Type. Nepal, near Seng Khola, 12500 ft [3810 m.], exposed cliffs, 4 Okt 1954, Stainton, Sykes & Williams 4677 (holotype K!, isotype BM!).
General distribution. Nepal (endemic; Bor 1965; Freitag 1985).
Distribution in Nepal. Baglung, Dolpa, Manang, Mustang, Rukum.
Habitat. open rocky or stony sandy slopes and scrublands.
Altitude. 3000–4200 m.
Note. Although Achnatherum staintonii has been confused with Stipa przewalskyi,
the latter species does not occur in Nepal. Achnatherum staintonii is easily distinguished
from Stipa przewalskyi by having maize-like vs. saw-like LEPs and in having unequal
glumes and distinctly longer lemma than palea vs. glumes as well as lemma and palea
subequal, respectively. Röser (2012) transferred five species of Stipa, including Stipa
staintonii, into their new genus Stipellula on the basis of his earlier molecular analysis
(Hamasha et al. 2012). Stipellula is characterized by its maize-like lemma epidermal
pattern which clearly distinguishes it from Stipa and confirms that these species belong
to the achnatheroid group of grasses within the Stipeae. However, there are no unique,
diagnostic morphological characters to separate Stipellula from Achnatherum which is
itself polymorphic and highly polyphyletic (Romaschenko et al. 2012; Hamasha et al.
2012). Thus, we prefer to treat Stipa staintonii as a member of Achnatherum.
Selected specimens studied. Baglung: Sing Khola, wet rocky cliff, 18 Sep. 1976,
H. Tabata, K.R. Rajbhandari & K. Tsuchiya 3711 (KATH). Dolpa: Ringmo, dry
hillslope, 29°10'20"N, 82°55'50"E, 3400 m, 2 Aug. 1973, S. Einarsson, L. Skärby &
B. Wetterhall 3128 (BM); Barbung Khola, 28°52'N, 83°15'E, 3030 m, 18 Jul. 1963,
J.D.A. Stainton 4417 (BM); Barbung Khola, 28°52'N, 83°18'E, 3030 m, 13 Jul. 1963,
J.D.A. Stainton 4417 (E); Suligad, Rhagaon, rocky slope, 29°28'N, 82°55'E, 2600 m,
25 Sep. 1982, K.R. Rajbhandari & K.J. Malla 6740 (KATH); Karnali, Ringmigaon,
on dry hilslope, 3400 m, 2 Aug. 1973, S. Einarsson, L. Skärby, B. Wetterhall 3126
(UPS); Karnali, Manang: Bhraka, Humde, open rocky slope, 28°38'24"N, 84°5'36"E,
3400 m, 3 Aug. 1983, K.R. Rajbhandari 8814 (KATH); Marsyandi valley, Tangi
above Manangbhot, on stony sandy places, 28°39'56"N, 84°1'33"E, 3800 m, 12
Oct. 1969, T. Wraber 36427 (E); Tangi, above Manangbhot, Tangje, on stony sandy
places, 28°39'22"N, 84°2'2"E, 3800 m, 12 Oct. 1969, T. Wraber 479 (BM). Mustang:
Entre Marpha et Syang; Syang, Marpha, 28°45'11"N, 83°41'28"E, 2650 m, 16 Sep.
1981, M.A. Farille 81-340 (E); Entre Larjung et Tukuche, 2550 m, 14 Sep. 1981,
M.A. Farille 81-313 (E); Cha Lungpa, in E-facing Cupressus forest, 3030 m, 3 Oct.
1977, G. Miehe 80 (BM); Phalyak, dry place 28°49'24"N, 83°44'23"E, 4110 m, 9
Synopsis of the tribe Stipeae (Poaceae) in Nepal
113
Aug. 2002, F. Miyamoto, N. Kurosaki, S. Akiyama, H. Ikeda, Y. Iokawa, Y. Takahashi,
M. Tsusaka & M.N. Subedi 20210022 (KATH); Phalyak to Pongio Kharka, 28°49'0"N,
83°45'0"E, 3800–4000 m, 9 Aug. 2002, F. Miyamoto, N. Kurosaki, S. Akiyama, H.
Ikeda, Y. Iokawa, Y. Takahashi, M. Tsusaka & M.N. Subedi 20220034 (KATH); Chele –
Samar, 28°57'43"N, 83°48'6"E, 3450-3670 m, 4 Aug. 2001, S. Noshiro, M. Amano, Y.
Iokawa, T. Kurosawa, & M.N. Subedi 20105005 (KATH); Tukuche, open rocky slope,
28°42'33"N, 83°38'37"E, 2630 m, 19 Jul. 1983, K.R. Rajbhandari 7991 (KATH);
Jomsom, open slope, 28°46'51"N, 83°43'27"E, 2760 m, 21 Jul. 1983, K.R. Rajbhandari
8072 (KATH); Jomsom to Sayang, cliffs in dry valley with scrub (Caragana, Clematis,
Rosa etc.), 28°46'51"N, 83°43'27"E, 2700 m, 19 Sep. 1999, Shrestha et al. 1014 (E);
Tukuche, 28°42'33"N, 83°38'37"E, 3180 m, 21 Aug. 1954, J.D.A. Stainton, W.R.
Sykes & L.H.J. Williams 7352 (BM). Rukum: Sen Khola, 28°42'54"N, 82°57'21"E,
3790 m, 4 Oct. 1954, J.D.A. Stainton, W.R. Sykes & L.H.J. Williams 4677 (BM).
Piptatherum P.Beauv., Ess. Agrostogr.: 17: 173 (1812)
Type. Piptatherum coerulescens (Desf ) P. Beauv.
Key to the genus Piptatherum
1
–
2
–
3
–
Panicle compressed .....................................................................................2
Panicle lax ...................................................................................................3
Awn terminal ................................................................................ P. laterale
Awn subterminal.............................................................................P. gracile
Lemma almost equal to glumes, apical part gradually narrowed into a persistent awn................................................................................. P. aequiglume
Lemma much shorter than glumes, apical part abruptly contracted into a
slender, caducous awn ................................................................... P. munroi
Piptatherum aequiglume (Duthie ex Hook.f.) Roshev., Bot. Mater. Gerb. Bot.
Inst. Komarova Akad. Nauk SSSR 14: 113 (1951).
Oryzopsis aequiglumis Duthie ex Hook. f., Fl. Brit. India 7(22): 234 (1896) (Basionym).
= Piptatherum sinense Mez, Repert. Spec. Nov. Regni Veg. 17(486–491): 211 (1921).
Type. India: distr. Jansar, Gamble 15143 (lectotype K! designated by Bor 1970).
General distribution. Afghanistan, Bhutan, S China, NW India, Nepal, Pakistan
(Freitag 1975; Noltie 2000; Wu and Phillips 2006).
Distribution in Nepal. Rukum.
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Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
Habitat. Moist mesophytic forests.
Altitude range. 3500–4000 m.
Selected specimens studied. Rukum: near Dogadi Khola, 3660 m, 8 Aug. 1954,
J.D.A. Stainton, W.R. Sykes & L.H.J. Williams 3801 (E 814753); nr. Dogadi Khola,
3790 m, 8 Aug. 1954, J.D.A. Stainton, W.R. Sykes & L.H.J. Williams 3794 (E 814768).
Piptatherum gracile Mez, Repert. Spec. Nov. Regni Veg. 17(486–491): 211 (1921).
≡ Oryzopsis gracilis (Mez) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 14: 347 (1939:).
Type. Tibet occ., 3900–4000 m., Thomson s.n. (lectotype W designated by Freitag
1975, isolectotype K!).
General distribution. Afghanistan, China, N India, Nepal, Pakistan, Tajikistan
(Freitag 1975; Wu and Phillips 2006).
Distribution in Nepal. Mustang.
Habitat. alpine steppes and meadows.
Altitude range. 2500–4000 m.
Selected specimens studied. Mustang: Marpha, 28°45'11"N, 83°41'28"E,
2670 m, 16 Sep. 1981, M.A. Farille 81-336 (E); Jomsom, 28°46'51"N, 83°43'27"E,
3200 m, Versant de Muksant Range, faisant face a Jomsom, 500 m au dessus du village, on rocks, 16 Sep. 1981, M.A. Farille 81-352 (E); Jharkot – Kagbeni, 28°50'17"N,
83°47'3"E, 2800–3550 m, 17 Sep. 1981, M.A. Farille 81-359 (E); 10 Jul. 2000, Y.
Iokawa, M.N. Subedi, Y. Takahashi & K. Kano 20020054 (E); Dzong Pura (Muktinath),
28°49'41"N, 83°51'19"E 3640 m, 29 July 1954, J.D.A. Stainton, W.R. Sykes & L.H.J.
Williams 2087 (E); Tange, 29°0'38"N, 83°56'45"E, 3640 m, 1 Aug. 1954, J.D.A.
Stainton, W.R. Sykes & L.H.J. Williams 2125 (E); Kagbeni, 28°50'17"N, 83°47'3"E,
3030 m, 8 Jun 1954, J.D.A. Stainton, W.R. Sykes & L.H.J. Williams 5659 (E); Ekle
Bhatti, on sunny rocky steep slope at pathside, 2270 m, 22 Sep. 1995, M. Mikage et
al. 9552384 (E 224287).
Piptatherum munroi (Stapf ex Hook.f.) Mez, Repert. Spec. Nov. Regni Veg. 17:
212 (1921).
Oryzopsis munroi Stapf ex Hooker, Fl. Brit. India 7(22): 234 (1897) (Basionym).
= Oryzopsis stewartiana Bor, Kew Bull., 272 (1953);
= Oryzopsis geminiramula Ohwi, Acta Phytotax. Geobot. 17: 14 (1957).
Type. NW India, Chenab Himalayas, 1852, Thomson (lectotype E 360583!, designated by Freitag 1975).
General distribution. China, N India, Kashmir, Nepal (Freitag 1975).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
115
Distribution in Nepal. Dolpa, Jumla, Mustang, Sankhuwasabha, Solukhumbu.
Habitat. Among dwarf Rhododendron shrubland and in coniferous forest.
Altitude range. 3490–4500.
Selected specimens studied. Dolpa: Sangdan, 28°55'N, 83°41'E, 4550 m, 21
Jul. 1963, J.D.A. Stainton 4443 (E). Jumla: Maharigaon, grassy slope, 29°19'50"N,
82°22'15"E, 4090 m, 18 Jul. 1952, O.V. Polunin, W.R. Sykes, & L.H.J. Williams
219 (E). Mustang: 2800 m, 16 Sep. 1981, M. Farille, 81-348 (E); Lo Tsho Dhyum,
Nr. Kali Grandaki River, Dhi (Dhee) area, stony river bank, scattered open vegetation, 22 Jul. 1998, W.R. Sykes 285/98 (E). Sankhuwasabha: Thudam, 27°45'31"N,
87°32'59"E, 3490–3480 m, 2 Nov 1971, L.W. Beer, C.R. Lancaster & D. Morris 10679
(E). Solukhumbu: Namche Bazar, along the trail to Phurte, South east facing grassy
slopes, open grassy slopes with bushes of Juniperus, 27°48'24"N, 86°42'46"E, 3420
m, 13 Sep. 2005, F.M. Watson et al. DNEP3 AX33 (E, KATH).
Piptatherum laterale (Regel) Munro ex Nevski, Trudy Bot. Inst. Akad. Nauk SSSR
Ser. 1, Fl. Sist. Vyssh. Rast. 4: 217 (1937).
Milium laterale Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 7: 645 (1881) (Basionym).
≡ Oryzopsis lateralis (Regel) Stapf ex Hook. f., Fl. Brit. India 7(22): 234 (1896);
≡ Piptatherum laterale (Regel) Roshev., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Kazahsk. SSR 14: 117 (1951).
= Oryzopsis pubiflora Hack., Denkschr. Kaiserl. Akad. Wiss., Wien Math.-Naturwiss.
Kl. 50(2): 8 (1885); ≡ Piptatherum pubiflorum (Hack.) Roshev., Bot. Mater. Gerb.
Bot. Inst. Komarova Akad. Nauk SSSR 14: 111 (1951).
= Oryzopsis vavilovii Roshev., Trudy Prikl. Bot. Selekts. 19(1): 123 (1928); ≡ Piptatherum vavilovii (Roshev.) Roshev., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk
Kazahsk. SSR 14: 118 (1951).
Type. Afghanistan, Kurram valley, Sikarm, common at 3650 m., dry localities, 1879,
Aitchison (holotype LE!, isotype K).
General distribution. widely distributed species, occurring from Turkey up to
Bhutan and S China (Freitag 1975, Tzvelev 1976).
Distribution in Nepal. Although we did not find any specimens of Piptatherum
laterale during this study, it is known from nearby regions of Bhutan, China, India
(Freitag 1975, Noltie 2000, Wu and Phillips 2006), so it is very likely also to be present in Nepal.
Trikeraia Bor, Kew Bull. 9(4): 555, f. s.n. (1954)
Type. Trikeraia hookeri (Stapf ) Bor.
116
Marcin Nobis et al. / PhytoKeys 128: 97–119 (2019)
Trikeraia hookeri (Stapf ) Bor, Kew Bull. 9(4): 555–556 (1954).
Stipa hookeri Stapf, J. Linn. Soc., Bot. 30: 120 (1894) (Basionym).
≡ Achnatherum hookeri (Stapf ) Keng, Claves Gen. Sp. Gram. Prim. Sinic. 106, 213
(1957).
= Timouria aurita Hitchc., J. Wash. Acad. Sci. 23: 134 (1933).
Type. Tibet, 4500 m, sheltered nullahs near water, Jul-Sep. 1891, Thorold 124 (holotype K!, isotype C).
General distribution. China (Tibet), India (Sikkim, Ladakh), Pakistan (Freitag
1985; Wu and Phillips 2006).
Distribution in Nepal. Although we did not find any specimens of Trikeraia hookeri during this study we include it here because it is known from nearby regions (Freitag 1975, Wu and Phillips 2006), so it is very likely to be present in Nepal. Freitag
(1985) reported this species from Nepal based on Sufed 104 (K), Mt. Everest, Tinkye
palin, 4270 m, but this specimen appears to have been collected on the Tibetan side
of Mt Everest.
Habitat. Scrublands, alpine mats.
Altitude range. 4000–4300 m.
Note. The 2005 DNEP3 expedition to Solukhumbu collected several specimens
which were identified as Trikeraia oreophila Cope by H. Noltie [Dingboche, on trail
south to the Lobuche Khola bridge, east facing valley side, 27°52'50"N, 86°49'7"E,
4230 m, 23 Sep. 2005, M.F. Watson et al. DNEP3 AX107 (E, KATH)]. These specimens are characterized by having 4–6 mm long lemma lobes, the awn arising below
the middle of the lemma and ovary with two stigmas. Recently, Trikeraia oreophila was
found to be conspecific with Sinochasea trigyna Keng (WCSP 2019). Morphological
and molecular studies have shown that the genus Sinochasea is distinct not only from
Trikeraia, but also from all the other genera of the tribe Stipeae, and therefore it was
transferred to the tribe Phaenospermateae Renvoize & Clayton (Schneider et al. 2011;
Romaschenko et al. 2012; Kellogg 2015).
Acknowledgements
We would like to express our gratitude to Paul Peterson and Mary Barkworth for their
valuable comments and improvements to the manuscript of the paper as well as to the
curators of BM, E, GOET, K, KATH, KRA, KUN, LE, NY, P, for their kind hospitality and assistance during visits and for making specimens of Stipa available on loan. We
thank The Sibbald Trust, the Davis Expedition Fund and the Russian Foundation for
Basic Research (RFBR) research project no. 18-34-20112). Financial support for M.
Nobis research came from the National Science Center, Poland (grant no. 2013/09/B/
NZ8/03287).
Synopsis of the tribe Stipeae (Poaceae) in Nepal
117
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