Corresponding author: Polina D. Gudkova (
Academic editor: Maria Vorontsova
In Nepal the
Nobis M, Gudkova PD, Pendry CA (2019) Synopsis of the tribe Stipeae (Poaceae) in Nepal. PhytoKeys 128: 97–119.
The tribe
Although agrostologists have maintained a broad concept of the genus
All previous treatments of the
Our treatment is based on herbarium specimens deposited in BM, E, GOET, K, KATH, KRA, KUN, LE, M, NY, P (
Districts of Nepal.
Nineteen morphological characters scored for each taxon were included in the analysis (Table
Morphological characters and character states.
Characters | States |
---|---|
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) |
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) |
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.
Detailed analyses of macro- and micromorphological structures of the lemma epidermis of Nepalese species of
Cluster analysis (
Lemma epidermal patterns (LEPs) of Old World
1 | Lemma with deflexed (retrorse), apical prickles |
|
– | Lemma lacking deflexed, apical prickles |
|
2 | Lemma lobes awn-like, 2–3 mm long, setaceous |
|
– | 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 |
|
– | 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) |
|
– | 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 |
|
– | 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 |
|
(India, W Himalaya) Kadarkanal,
Himalayas: Bhutan, S China, Ladakh, N Myanmar, N India, Nepal (
Baglung, Bajura, Darchula, Dolkha, Humla, Jumla, Lalitpur, Mugu, Myagdi, Ramechhap, Rasuwa, Rukum, Sankhuwasabha, Solukhumbu, Taplejung.
Alpine meadows,
2200–4000 m.
1 | Upper part of awn (seta) scabrous, with hairs up to 0.4 mm long |
|
– | Upper part of awn (seta) pilose, with hairs over 0.4 mm long |
|
2 | Upper part of awn and tips of glumes spirally twisted |
|
– | Upper part of awn and tips of glumes not twisted |
|
3 | Ligules of vegetative shoots up to 0.3 mm long. Awn column with 0.6–0.7 mm long hairs |
|
– | Ligules of vegetative shoots over 2 mm long. Awn column with 1.5–2.5 mm long hairs |
|
(China) Tibet, Gnari (Nari) Khorsum,
China, Kyrgyzstan, N India, Mongolia, Nepal (
Mustang.
High altitude steppes, scree.
2750–3600 m.
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,
Bhutan, China (Tibet), India (Ladakh, Sikkim), Nepal, Pakistan, (
Mustang.
High mountain steppes and alpine mats, among subalpine shrubs and on rocky ledges.
3500–5000 m.
These specimens were previously identified as
Selenginskaya Dauriya, gory mezhdu Temnikom i Dzhidoi, yugo-zapadnaya chast khr. Borgoiskogo, na sklonakh so stepnoi rastitelnostyu, 28 Jul. 1912,
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. (
Mustang (
High mountain semi-desert.
3900–4000 m.
These specimens were previously identified as
Open grass land in rear of Shaowusze Agricultural Station, Taining district, Sikang province, 22 Jul. 1940,
Bhutan, S China, N India, Nepal, Pakistan, (
Mustang, Rasuwa, Solukhumbu.
Alpine meadows.
2800–4100 m.
These specimens were previously identified as
1 | Awn with 0.3–0.5 mm long hairs on column. Seta scabrous |
|
– | Awn variously pilose on column, with hairs over 1 mm long. Seta with 0.5–1.5 mm long hairs |
|
2 | 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) |
|
– | Glumes, lemma and palea equal or only slightly unequal |
|
3 | Panicles open, 3–5 cm wide, branches up to 6 cm long, spreading |
|
– | Panicles compressed, 0.7–2 cm wide, branches 0.3–2.8 cm long, suberect or narrowly ascending |
|
[India] Tehri Garwhal, Lekhus, below Srikanta, 12000–13000ft, 11 Aug. 1853,
China, N India, Kashmir, Nepal (
Myagdi.
Mountain shrublands.
3400–3800 m.
China, Kansu and Tsinghai border [in regione opp. Labrang],
Bhutan, Birma, China (Tibet), N India, Nepal (
Bajhang, Bajura, Dolakha, Jumla, Mustang, Myagdi, Ramechhap, Rasuwa, Rukum, Solukhumbu.
Alpine meadows, grassy mountain slopes.
3300–5000 m.
These specimens were previously identified as
China. Xizang: Yadong, 14 Sep. 1974,
Bhutan, China (Tibet), Nepal (
Bajhang, Dolkha, Rasuwa, Solukhumbu.
Alpine meadows, moist grassy places, under shrubs, swampy places,
3600–4600 m.
These specimens were previously identified as
Sikkim-Himalaya, Tibetan region, 14000–16000ft, 1861,
Himalayas: China (Tibet), India (Ladakh and Sikkim), Nepal (
Solukhumbu, Mustang.
Alpine meadows, moist grassy places, under shrubs, swampy places,
4400–5300 m.
1 | Glumes distinctly unequal. Lemma apex with ring of hairs over 3 mm long. Callus 0.5–0.7 mm long, acute at the apex |
|
– | 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 |
|
– | Lemma and palea subequal. Leaves flat. Culms over 60 cm long. Panicle with widely spreading branches |
|
[India] N. W. Himalaya, Kulla, Oct. 1876,
Afghanistan, Bhutan, China, NW India, Nepal, Pakistan (
Dolpa, Manang, Mustang.
Open dry slopes, among shrubs and in Bamboo (
2400–4000 m.
[India] ad ruped in excelsis Emodi Cashemyrianim 2750 m, 1831,
E Afghanistan, Bhutan, China (Tibet), NW India, Nepal, Pakistan (
Mustang.
Dry mountain slopes, especially in rock crevices.
2500–3000.
Nepal, near Seng Khola, 12500 ft [3810 m.], exposed cliffs, 4 Okt 1954,
Nepal (endemic;
Baglung, Dolpa, Manang, Mustang, Rukum.
open rocky or stony sandy slopes and scrublands.
3000–4200 m.
Although
1 | Panicle compressed |
|
– | Panicle lax |
|
2 | Awn terminal |
|
– | Awn subterminal |
|
3 | Lemma almost equal to glumes, apical part gradually narrowed into a persistent awn |
|
– | Lemma much shorter than glumes, apical part abruptly contracted into a slender, caducous awn |
|
India: distr. Jansar, Gamble 15143 (lectotype K! designated by
Afghanistan, Bhutan, S China, NW India, Nepal, Pakistan (
Rukum.
Moist mesophytic forests.
3500–4000 m.
Tibet occ., 3900–4000 m.,
Afghanistan, China, N India, Nepal, Pakistan, Tajikistan (
Mustang.
alpine steppes and meadows.
2500–4000 m.
NW India, Chenab Himalayas, 1852, Thomson (lectotype E 360583!, designated by
China, N India, Kashmir, Nepal (
Dolpa, Jumla, Mustang, Sankhuwasabha, Solukhumbu.
Among dwarf
3490–4500.
Afghanistan, Kurram valley, Sikarm, common at 3650 m., dry localities, 1879,
widely distributed species, occurring from Turkey up to Bhutan and S China (
Although we did not find any specimens of
Tibet, 4500 m, sheltered nullahs near water, Jul-Sep. 1891, Thorold 124 (holotype K!, isotype C).
China (Tibet), India (Sikkim, Ladakh), Pakistan (
Although we did not find any specimens of
Scrublands, alpine mats.
4000–4300 m.
The 2005 DNEP3 expedition to Solukhumbu collected several specimens which were identified as
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