Stipa tremula (Rupr.) M. Nobis, 2022

Nobis, Marcin, Krzempek, Marta, Nowak, Arkadiusz, Gudkova, Polina D. & Klichowska, Ewelina, 2022, Resurrection of Stipa tremula and taxonomy of the high-alpine species from the Stipa purpurea complex (Poaceae, Pooideae), PhytoKeys 196, pp. 21-47 : 21

Stipa tremula (Rupr.) M. Nobis comb. nov.

Ptilagrostis semenovii = Ptilagrostis semenovii Krasn. [originally P. semenovi Krassn.], Spisok rastenii sobrannykh v vostochnom Tyan-Shane, letom 1886 goda, 125, 1887. Type protologue: Prope fl. sary-Jassy. Type: Ptilagrostis tianschanica Krassn., Ad flumen Sary-Jassy, 1 Aug 1886, Krassnow s.n. (lectotype, distinguished here, LE 01009431!, isolectotype, LE 01009430!);

Stipa semenowii ≡ Stipa semenowii Krasn. [originally S. semenowi Krassn.], Scripta Botanica Horti Universitatis Imperialis Petropolitanae, Botanicheskiia Zapiski 2(1): 22. 1887. Type protologue: In valle fluminis Sary-Jassy in montibus Thian-Schan non procul ab alpe Chan-tengri et in trajectu Turguen-Aksu non rara;

Stipa semenovii ≡ Stipa semenovii Krasn. [originally S. semenovi Krassn.], Zapiski Imperatorkago Russkago Geograficheskago Obschestva, Opyt’ istorii rasvitya flory yuzhnoi chasti vostochnago Tyan-Shanya 19: 341-342. 1888. Type protologue: In valle fluminis Sary-Jassy in montibus Thian-Schan non procul ab alpe Chan-tengri et in trajectu Turguen-Aksu non rara.

Stipa pilgeriana = Stipa pilgeriana K.S. Hao, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 68(5): 583-584. 1938. Type protologue: China: Kokonor [Qinghai]: Ming-ke-Shan, Tsi-gi-gen-ba-Gebiete, 3900 m (Nr. 1009 - am 25 August). Type: Kokonor [Qinghai], Mingke, Tsigigenpa, alt. 3900 m, 25 Aug 1930, Hopkinson 1009 (holotype, PE 707247 [additional no. on the sheet 01940135], [label 2: S. pilgeriana Hao sp. nov.]).

Basionym.

Lasiagrostis (Leptanthele) tremula Rupr., in Ost.-Sack. & Rupr., Sertum Tianschanicum, Mémories de L’Académie Impériale des Sciences de St.- Pétersbourg, Sér. 7, 14(4): 35. 1869.

Type protologue.

Die Gegend des Sarymeki-Flüsschens, 28 Jul 1867, F. Osten-Sacken s.n. Type. [China] In der Gegend des Flusses Sarymeki, Südlicher Abhang des Tian-Schan, in regione subalpina jugi Thian-Schan, 28/9 Julio 1867, Lib. Baro Fr. Osten-Sacken s.n. (lectotype, designated by Tzvelev 1976: 583, LE 01009426!, isolectotypes, P 02241060!, K 000587435!).

Description.

Perennial plants, densely tufted, with a few culms and numerous vegetative shoots; culms (11-)20-30(-45) cm tall, 1-2(-3)-noded, nodes distributed close together and only in the lowermost part of the culm. Leaves of vegetative shoots: sheaths glabrous and smooth rarely sparsely and shortly pubescent; ligules truncate, ciliate on the apex, on the external sheaths (0.3-)0.7-1.1(-1.6) mm long, whereas on the internal sheaths, (0.5-)1.0-2.0(-3.0) mm long; blades convolute, green, pale green to greyish, (3-)4-12(-18) cm long, 0.3-0.5(-0.7) mm in diameter, adaxial surface densely covered by 0.15-0.25 mm long hairs, abaxial surface glabrous or scabrous to shortly pilose and grading to almost smooth towards the apex. Cauline leaves: sheaths glabrous and smooth; ligules on the lower sheaths truncate or acute (0.4-)1.0-1.5(-2.1) mm long, on the middle and upper sheaths acute (1.0-)1.5-2.7(-4.0) and 2.0-3.1(-3.6), respectively; blades convolute, green, pale green or greyish (2.3-)2.0-3.0(-8.9) cm long, adaxial surface covered with short hairs, abaxial surface glabrous, scabrous to shortly pubescent. Panicle (7-)11-18(-26) cm long, open, with (3-)6-15(-18) spikelets, at base enclosed by the sheath of the uppermost leaf or rarely exerted; branches ascending, flexuous, setulose, slightly scabrous to glabrous and smooth, single or paired, 2-6 cm long. Glumes subequal (the lower slightly longer than the upper), purplish, (13.0-)14-17.0(-19.5) mm long, lanceolate, with hyaline margins and long tip. Floret (lemma + callus) (7.5-)8.4-9.5(-11.0) mm long and (0.7-)0.8-1.0 mm wide. Callus (1.5-)1.6-2.0(-2.3) mm long, densely pilose, on ventral part with hairs (0.4-)0.5-0.8(-1.0) mm long, on dorsal with (0.3-)0.4-0.6(-0.8) mm long hairs; callus base 0.5-0.8 mm long and 0.15-0.25 mm in diameter, sharply pointed, scar narrowly elliptic. Lemma coriaceous, pale-green, purplish or brownish, covered throughout, from the bottom up to 0.5-2.3 mm to the top, by dense ascending to appressed hairs 0.3-0.8 mm long, above being glabrous (hairless) or with scattered hairs just below the apex; top of lemma glabrous or surpassed by a poorly- to well-developed ring of unequal hairs 0.2-0.5 mm long and with (or without) two minute apical lobes. Awn (44-)60-74(-90) mm long, bigeniculate; the lower segment of the awn (column) (6-)11-15(-20) mm long, twisted, 0.3-0.4 mm wide at the base, with (1.3-)1.7-2.0(-2.3) mm long hairs, the middle segment of the awn (5-)6.5-10(-13) mm long, twisted, with (1.6-)2.0-2.3(-2.6) mm long hairs; terminal segment of the awn (seta) slightly arcuate or flexuous (26-)39-53(-73) mm long with hairs longer than those on columns, (1.8-)2.2-2.6(-3.1) mm long, gradually decreasing in length towards the apex. Palea equalling lemma in length. Ovary with two styles.

Habitat.

High mountain steppes, dry grasslands, mats, screes, semi-deserts, stony slopes, gravelly or sandy flats and valleys (1900-)3000-4800(-5100) m a.s.l.

Distribution.

Tian Shan, Pamir, Karakorum, eastern Himalayas, Kun-lun, Qilan-Shan, eastern Tibetan Plateau (Fig. 3 ); Kazakhstan, Kyrgyzstan, Tajikistan, north Pakistan, India (Ladakh), China (Xinjiang, Gansu, Qinghai, Sichuan, western and eastern Xizang) ( Pavlov 1956; Ovchinnikov and Chukavina 1957; Lazkov and Sultanova 2014).

Note.

Lasiagrostis tremula was described from Tian-Shan Mts. by Ruprecht (in Osten-Sacken and Ruprecht 1869), based on the collection of Fr. Osten-Sacken (without indication of type, place of its preservation and the collection number). During the revision of the herbarium material of Stipa purpurea s.l., we found three sheets with specimens of Lasiagrostis tremula collected by Osten-Sacken, of which the one preserved in LE, previously regarded as type by Tzvelev 1976, Freitag 1985 and Nobis et al. 2020) is here corrected to lectotype. The duplicates preserved in K and P are isolectotypes.

Selected specimens examined.

Kyrgyzstan: Tsentralnyi Tyan’-Shan’, dolina r. Kuilyu bliz ust’ya r. Oroi-su, na rechnoi terrase, alt. 2100 m, 10 Aug 1956, I.S. Pushkin s.n. ( MW 803233 , 803232); Tian-Schan centr., in valle flum. Sary-tschat, prope glaciem Kolpakovskyi, orientem versus a statione meteorologica, steppum frigidum, alt. ca. 3400 m, D. Wyschiwkin s.n. ( MW 803231 , US 04002929, P 02662772); Narynskii raion, bassein r. Aksai, khr. Kok-kiya, kobrezovyi lug, 15 Aug 1926, M. Sovetkina, M. Uspenskaya 1745 (MW 803 229); Issykulskie syrty, kovyl’. step, alt. ca. 4000 m, 16 Aug 1953, N. Trulevich s.n. ( MW 803234 ); Issykulskie syrty, kovyl’. step, alt. ca. 3000 m, 16 Aug 1953, N. Trulevich s.n. ( MW 803228 ); Semirech. obl. Przhevalsk. u. dolina r. Naryna mezhdu r. Bashka-Su i r. Ulan’, kamenistye sklony, 20 Jul 1913, V. Sapozhnikov 252 (LE); Semirech. obl. Przhevalsk. u. r. Ak-tash’, plato, alpiiskaya pustyn. step’, 28 Jul 1913, V. Sapozhnikov 250 (LE, TK); Issyk-Kulskaya obl., Ak-Sulskii raion, khr. Teskei Ala-Too, pravyi bereg r. Sary-Dzhaz, 10 Aug 1988, Pimenov, Kmaishov, Kamsharaeva s.n. (KRA, FRU); Naryn prov., At-Bashii reg., Ken-Suu r., 15 Aug 1976, Abgarova s.n. (KRA); Tian’-Shan’, dolina r. Kuelyu, prav. prit. Sary-Dzhasa, 20-30 Jul 1902, V. Sapozhnikov s.n. (LE, TK); Dzhety-Oguzovskii region, Akshiryak, 6-7 km NE of Uzun River, terrace of Cholak River, alt. 3800 m, high mountain steppe, 26 Jul 1935, I Sanych, G. Sabardina 121 (BM 1191534); Dolina Aksya, Poima r. Terek, Ken-Suu, 1960, Makarenko s.n. (FRU, KRA); Semirech. obl. Przhev. u r. Inyl’chek, v 10 v. ot’ lednika, 1 Aug 1912, V. Sapozhnikov, B. Shishkin s.n. (LE); C Tian’-Shan’, verkhie Narynskie syrty, ur. Kum-tala, 23 Aug 1926, R.I. Abolin 1034 (LE); Semirech. obl. Przhev. u r. Sarydzhas’, pri usti r. Myntur’, syrty, 28 Jul 1912, V. Sapozhnikov, B. Shishkin s.n. (LE, TK, AA); Tian’ -shanskaya obl., Terskei khr., pravyi bereg r. Sary-dzhaz, uste r. Kuisyu, 19 Jul 1967, Pavlova s.n. (LE); Issykulskaya obl. Pokrovskie syrty, Aug 1957, Moldoyarov s.n. (LE); Semirech. obl. Przhev. u r. Dzhangart’ srednee techenie, travyanistye sklony, 31 Jul 1913, B. Shishkin s.n. (TK); Semirech. obl. Przhev. u Kara-Archa, kamenistye sklony, 6 Aug 1912, V. Sapozhnikov, B. Shishkin s.n. (TK); C Tian’-Shan’, Khr. Bornokoi, basein r. Karakol, alt. 3700 m, 1949, M. Tlo[..] s.n. (AA); Tyan’ -shanskaya obl. At-bashinsii raion, bass. r. Ak-saya, stats. Cykanova, Jul 1972, Nemetskaya s.n. (FRU); Atbashinskii raion, dolina Ak-sai, pologie sklony, 31 Jul 1939, Gusarova s.n. (FRU). Tajikistan: Pamir, Kara-kul, dol. Kara-Art, v 5 km ot ust’ya na terrase, alt. 4050 m, 28 Aug 1962, S.S. Ikonnikov 14896 (LE, KRA 479099); Pamir, ad lacum Kara-kul, alt. 13500 ft, 5 Jul 1901, Alexeenko 1392 (LE); Pamir, Shad-Put, peski nizovev dol. r. Up-turukh, alt. 4000 m, 19 Jul 1945, I. Raikova 159 (LE); Pamir, Rang-Kulskii c/c, 1952, Sudorov s.n. (LE); SE Pamir, zap. sklon Ak-tash, Shaimak settl., 29 Aug 1953, S. Ikonnikov s.n. (LE); Shot-put, Rang-Kulkii c/c, 20 Jul 1945, I. Raikova 161 (LE); Pamir, Kara-kul’, po krayu lugov u reki Kara-art, 20 Aug 1955, Sudorov s.n. (LE). Pakistan: S side of Pamir Pass, near Shuwart, alpine steppe, dominated by Gramineae and flat cushions of Oxytropis spp., pastures and flushes, grazed, 36°23-25'N, 75°41-43'E, alt. 4420-4520 m, 16 Aug 1991, G. & S. Miehe 6173 (GOET); Upper Braldu tributary, above Chikor, alpine steppe, dominated by Gramineae and flat cushion of Oxytropis spp., 36°22-24'N, 75°22-24'E, alt. 4220 m, 17 Aug 1991, G. & S. Miehe 6209 (GOET); Khunjerab-pass, alpine steppe, 36°50'N, 75°25'E, alt. 4460 m, 18 Aug 1990, G. & S. Miehe 2495 (MSB 154127). India: NW India, Jammu and Kashmir State, Ladakh, Zanskar, Markha, Hankar Village to Zalung Karpo La, W slopes of Kyangze, alt. 4700 m, 20 Aug1998, 33°44,5'N, 77°29'E, L. Klimeš 86 (KRA 479097). China: Xinjiang province, Qira Xian, Nor, Yamei, on slope grassland, alt. ca. 3100 m, 3 Jul 1988, S. Wu, H. Ohba, Y. Wu, Y. Fei 2532 (MOIS 4374351); Qiemo Xian, Kongqibulaker, on desert grassland, alt. ca. 3200 m, 19 Jul 1988, S. Wu, H. Ohba, Y. Wu, Y. Fei 2591 (MOIS 5660759); alt. 4050 m, 8 Aug 1959, no. 1685 ( PE 707259 ); 26 Aug 1965, no. 39181 ( PE 707280 ); Qiemo Xian, Kongqibuaker, on arid soil slope, alt. ca. 4000 m, 26 Jul 1988, S. Wu, H. Ohba, Y. Wu, Y. Fei 2103 (MOIS 4364181); Qiemo Xian, Kongqibulaker, on sparsely grass-covered hill slope, alt. ca. 3150 m, 19 Jul 1988, S. Wu, H. Ohba, Y. Wu, Y. Fei 2056 (MOIS 4373368); SW Xinjiang, Karakoram, Aghil Shan northern declivity, ca. 7 km northwest of Aghil Pass, gravelly slope, limestone, 36°14'N, 76°34'E, alt. 4200 m, 30 Aug 1986, B. Dickoré 487 (GOET); SW Xinjiang, Karakoram, Aghil Shan northern declivity, Aghil Valley, at the Kirghiz summer settlement, ca. 19 km SW of Ylik (Yarkand), north-facing limestone cliff, in fissures, 36°15'N, 76°33'E, alt. 4150 m, 29 Aug 1986, B. Dickoré 472 (GOET); SW Xinjiang, Karakoram, Aghil Shan northen declivity, ca. 3 km NWW of Aghil Pass, gravelly slope, between granite boulders, dry alpine “turf”, 36°13'N, 76°35'E, alt. 4520 m, 30 Aug 1986, B. Dickoré 513 (GOET); Tibet borealis, ozero Orich-nor, yuzhn. bereg, alt. 13500 ft, 18 Jul 1884, N.M. Przwalski 338 (LE), Kam (Tibet), basein Yan-tszy-tszyan’a (r. Goluboi), 1 Aug 1900, V.O. Ladygin 434 (LE); Kun-lun, Kashgaria, verkhov’e r. Lapet, 20 Jul 1942, V.I. Serpukhov 474 (LE); Kun-lun, Kashgaria, r. Kara-dshilga, levyi pritok r. Gon-arek, alt. 4000-4500 m, 22 Jul 1942, V.I. Serpukhov 508 (LE); Gansu Province, Yumu Mountain, Dahe District, Sunan County, dry hillside, 5 Aug 1967, Hexi Team 165 (PE); Kansu, alt. 2600 m, 25 Aug 1967, no. 274 ( PE 707218 ); Kansu, Richthofen (Nan-Shan), Hung-Shui-Pa-Shang-Ho, alt. ca. 3500 m, 28 Aug 1931, B. Fries-Johansen 2878 (BM 1031149, 1191537); Qinghai Province, Haixi Mongol and Tibetan Autonomous Prefecture, stone gap, alt. 3000 m, 28 Jul 1975, W. Wong, B. Guo 11742 (PE); near the road G 109, 18 km SW of the lake, NE of Dashi Bridge, 3617 m, 3 6°43'34"N, 99°34'51"E, 24 Jul 2010, B. Paszko s.n. (KRA); Mengnan County, Qingshui Town, dry slope, alt. 3100 m, 26 Aug 1975, W. Wang, Be-z. Guo 12192 (PE); Qaidam, kumirnya Dulan’ -Khiti, alt. 10100 ft, 12 Aug 1901, V.O. Ladygin s.n. (LE); Yeningou North Mountain, Qilian County, meadow, alt. 3400 m, 5 Sep 1975, W. Wang, B-z. Guo 12473 ( PE 707250 ); alt. 3700 m, 4 Sep 1975, W. Wang, B-z. Guo 12411 ( PE 707242 ); alt. 3600 m, 10 Aug 1975, W. Wang, B-z. Guo 12053 ( PE 707240 ); Dari Xian: Jimai Xian, Huleanma, along Huang, flood plain of Hunag He, tussock grass, 33°43'40"N, 99°21'1"E, alt. 4030 m, 11 Aug 1993, T-n. Ho, B. Bartholomew 1158 ( PE 707267 , BM 573601 ); Shinaihaixiang, alt. 3230 m, 36°59'26,9"N, 99°36'03,0"E, 24 Jul 2010, B. Paszko s.n. (KRA); Tibetan Autonomous Prefecture of Haibei, hillside, alt. 3300 m, 29 Aug 1975, W. Wang, B-z. Guo 12262 ( PE 707241 ); Menyuan County, Gingshizui, dry hillside, alt. 3200 m, 24 Aug 1975, W. Wang, B-z. Guo 12176 (PE); Qinghai, C Tibet, Tangula Shan N, Upper Yangtse Basin, Bi Qu, Wenquan - Yanshiping (Lhasa - Golmud Rd.), 33°31'N, 91°58'E, alt. 4800 m, 18 Aug 1989, B. Dickoré 4207 (MSB 152888); Qinghai, C Tibet, Tangula Shan N, Upper Yangtse Basin, Gar Qu Vy. (Mt. Geladandong - Yanshiping), 33°36'N, 91°44'E, alt. 4850 m, 2 Sept 1989, B. Dickoré 4617 (MSB 152889); Heka area of Xinghai County in Qinghai, dry slope, 35.9 N, 99.9 E, alt. 3300 m, Jul 1965, P-c. Kuo & T-n. Ho 65-6111 (GOET); Gande ( Gadê) Xian, near Shanggongma Xiang, Gande ( Gadê) Shan, on road from Dari (Darlag) to Gande ( Gadê), flat-bottomed valley with moist alpine meadows on bottom, slopes with rocky outcrops and thick turf, on grassy slope, alt. 4150 m, 33°53'45"N, 99°40'50"E, 9 Aug 1993, T. Ho, B. Bartholomew & M. Gilbert 952 (E 690603, BM 000573603); Madoi Xian: just E of Malayiwan, on road between Gonghe and Madoi, open Stipa purpurea steppe on sandy soil, alt. 4050 m, 35°0'N, 98°30'E, 10 Aug 1996, T. Ho, B. Bartholomew, M. Watson & M. Gilbert 1585 (E 125866, PE 707271 , BM 573604 ); Maquin ( Maqên) Xian, Naheqingma, Youyun Xiang, between Dari (Darlag) and Huashuxua, consolidated sand dunes with disturbed flat areas, alt. 4190 m, 33°18'39"N, 99°10'53"E, 17 Aug 1993, T. Ho, B. Bartholomew & M. Gilbert 1342 (E 690703, BM 573602 ); Dari (Darlag) Xian: Huleanma, Jianshe Xiang, S side of the Huang He and SW of confluence with the Dari He (Dar Qu), flood plain of the Huang He, tussock grass, alt. 4030 m, 33°43'N, 99°21'E, 11 Aug 1993, T. Ho, B. Bartholomew & M. Gilbert 1158 (E 690604); Maquin ( Maqên) Xian, Dawu Xiang, along the Gequ He, N of Maquin ( Maqên), Jiangrang, side of valley with steep slope, mostly dry with large tussock grasses and bare soil, shallower slopes with alpine meadow, areas of deeper soil with shrubs, on slope, alt. 3500 m, 34°42'28"N, 100°14'39"E, 31 Jul 1993, T. Ho, B. Bartholomew & M. Gilbert 623 (E 690605, PE 707268 , BM 573600 ); Yushu Xian, Xiao Surmang Xiang, between Jerikug and the Xizang border, alt. 3550-3650 m, 32°6'N, 97°16'E, 24 Aug 1996, T. Ho, B. Bartholomew, M. Watson & M. Gilbert 2315 ( E 61899 ); Near the camp XLIV in E Tibet, 15 Jul 1901, Sv. Heidin 5127 (BM 001191536); Xizang Province, near Ranwu District, Basu County, alt. 4000 m, 18 Aug 1980, no. 1231 ( PE 707340 ); Tibet, alt. 5100 m, 27 Aug 1963, no 1989 ( PE 707466 ).

Figures

Figure 3. The distribution range of Stipa purpurea (purple backgrounds and points), S. tremula (blue backgrounds and points) and S. arenosa (yellow points).