Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) DOI: 10.17581/bp.2015.04103
Chromosome numbers in some species of Poaceae from Russia
Nina S. Probatova 1*, Vitaly P. Seledets 2 & Vyacheslav Yu. Barkalov1
Nina S. Probatova 1 * ABSTRACT e-mail: [email protected] Chromosome numbers (CN) for 35 species and one hybrid in Poaceae from 9 ge Vitaly P. Seledets 2 nera: Beckmannia, Brachypodium, Bromopsis, Catabrosa, Elymus, Elytrigia, Festuca, Mi e-mail: [email protected] lium, Poa, from Russia are presented. For Bromopsis austrosibirica Peschkova, De Vyacheslav Yu. Barkalov 1 schampsia baicalensis Tzvelev, D. submutica (Trautv.) O.D. Nikif., Festuca chionobia e-mail: [email protected] T.V. Egorova et Sipliv., F. dahurica (St.-Yves) V.I. Krecz. et Bobrov, F. kolesnikovii Tzvelev, F. vylzaniae (E. Alexeev) Tzvelev, Poa khokhrjakovii Prob. and P. krasno 1 Institute of Biology & Soil Science FEB borovii Stepanov the CNs were revealed for the first time. The new CNs were RAS, Vladivostok, Russia obtained for Bromopsis heterophylla (Klokov) Holub, Deschamp sia obensis Roshev., 2 Elytrigia trichophora (Link) Nevski. Some species were studied first time: Elymus Pacific Institute of Geography FEB RAS, kamoji (Ohwi) S.L. Chen and Catabrosa minor (Bab.) Tzvelev – in Russia, P. glauca Vladivostok, Russia Vahl – in Amurskaya Oblast’, Festuca jacutica Drobow – in Khabarovskii Krai, Poa angustifolia L. – in the North Koryakia, Milium effusum L. and Deschampsia komarovii * author for correspondence V.N. Vassil. – in Kamchatka Peninsula. From the Commander Islands the hyb rid Poa alpigena (Blytt) Lindm. × P. arctica R. Br. was studied. Besides, in the “tuft Manuscript received: 20.03.2015 form” of Fes tuca rubra L., which is distributed along seacoast and islands of the Review completed: 29.03.2015 Primorskii Krai, the CN was firstly revealed. F. chionobia obviously is the first Accepted for publication: 01.04.2015 pseudoviviparous species with the diploid CN (2n = 14). Published online: 20.04.2015 Keywords: chromosome numbers, vascular plants, Poaceae, flora, Russia РЕЗЮМЕ Пробатова Н.С., Селедец В.П., Баркалов В.Ю. Числа хромосом неко торых видов злаков (Poaceae) России Приводятся числа хромосом (2n) для 35 видов и одного гибрида злаков из родов Beckmannia, Brachypodium, Bromopsis, Catabrosa, Elymus, Elytrigia, Festuca, Milium, Poa во флоре России. Впервые исследованы в кариологическом от ношении Bromopsis austrosibirica Peschkova, Deschampsia baicalensis Tzvelev, D. submutica (Trautv.) O.D. Nikif., Festuca chionobia T.V. Egorova et Sipliv., F. dahurica (St.-Yves) V.I. Krecz. et Bobrov, F. kolesnikovii Tzvelev , F. vylzaniae (E. Alexeev) Tzvelev, Poa khokhrjakovii Prob., P. krasnoborovii Stepanov. Новые (не известные ранее) числа хромосом установлены у Bromopsis heterophylla (Klokov) Holub, Deschampsia obensis Roshev., Elytrigia trichophora (Link) Nevski. Elymus kamoji (Ohwi) S.L. Chen и Catabrosa minor (Bab.) Tzvelev исследованы впервые для России, Poa glauca Vahl – впервые для Амурской обл., Festuca jacutica Drobow – в Хабаровском крае, Poa angustifolia L. – в Северной Корякии, Milium effusum L. и Deschampsia komarovii V.N. Vassil. впервые исследованы на Камчатке. На Командорских островах изучен гибрид Poa alpigena (Blytt) Lindm. × P. arctica R. Br. Впервые уста новлено число хромосом у дернистой формы Festuca rubra L., которая встречается на морском побережье и островах в Примор ском крае. Впервые выявлен псевдовивипарный таксон освяницы с дипло идным числом хромосом – Festuca chionobia (2n = 14). Ключевые слова: числа хромосом, сосудистые растения, злаки, Poaceae, флора, Россия
Here we present results of сhromosome number (CN) stu St. Petersburg). First CN data are indicated by asterisk (*). The dy of 35 Poaceae species from Russia, mainly from East Siberia number of the dot on the map follows the number of voucher and the Russian Far East (RFE), as well as from European Arc specimen. Brief information on the affinity and distribution of tic, North Caucasus and Crimea (Fig. 1). Chromosome coun the species studied is presented. tings in some species were made by E.G. Rudyka (indicated by ER), on squashed preparations of root tips fixed with Carnoy’s Beckmannia borealis (Tzvelev) Prob., 2n = 14 (ER) solution. We also include here some previous CN countings of Russia, the Far East, Kamchatka Peninsula, Bystrinskii A.P. Sokolovskaya (indicated by AS). The root tips were taken Raion, Anavgai village, water-logged ground at the rivulet, from living plants, or from seedlings obtained through herbari 322 m alt., 9 Aug 2013, coll. V.V. Buryi 12597: 10. Endemic um specimens, which were collected in the field. Preparations of Russia. This species is distributed in the north and cent were stained with iron hematoxylin. Voucher specimens are ral parts of European Russia, in Altai and Yakutia, and it is preserved in the Herbarium VLA, Vladivostok (some – in LE, a rare species in the Russian Far East (RFE), where it occurs
©Botanical Garden-Institute FEB RAS. 2015 1 Probatova et al.
Figure 1 Study area. Dots with numbers from 1 to 44 are the sampling plot locations in wet meadows of riversides in Kamchatka (mainly cent alien in the RFE). Both species have variable ploidy: 2n = ral) and Okhotia (near Magadan). This species is probably 28, 42, 56 – many CN counts for B. inermis, but less – for arised from hybridization of B. syzigachne (Steud.) Fern. × B. australis: 2n = 28, 56 (Tzvelev & Probatova 2010 a; Pro B. eruciformis (L.) Host, but now the last species is absent in batova et al. 2010). In the past these forest and steppe spe the RFE. The RFE populations of B. borealis possibly are cies probably were more isolated, but now B. inermis, which relict. The CN of B. borealis was studied in Altai, Yakutia is younger and more active than B. australis, owing to human and Kamchatka (Probatova & Sokolovskaya 1980, 1984; impact, widely extended its area of distribution to the south Probatova & Seledets 2008). All the species of the small and they easily hybridize (Tzvelev & Probatova 2010 a). 2n genus Beckmannia are diploids. 2n = 2x. = 4x, 6x. Brachypodium spryginii (Tzvelev) Tzvelev (B. sylvaticum * Bromopsis austrosibirica Peschkova, 2n = 28 (AS) (Huds.) P. Beauv. subsp. spryginii Tzvelev), 2n = 18 (ER) Russia, East Siberia, Republic Sakha-Yakutia, Bulunskii Russia, North Caucasus, Krasnodarskii Krai, Abinskii Raion, left riverside of the Lena River, 30 km above Kiu Raion, 18 km S of Abinsk town, near Shapsugskaya village, ssiur settlement, near the mouth of the Tegeliakh River, the riverside forest along the Abin River (the Kuban’ Riv steep slope, 4 Aug 1973, coll. N.S. Probatova & V.P. Sele er basin), 1 Sep 2009, coll. N.S. Probatova & V.P. Seledets dets 3771: 3. Occurs on forest edges and clearings in the 11652: 44. This species was described from Crimea and it is river valleys. The Altai – East Siberian endemic species. It distributed also in the North Caucasus (west part, including belongs to the group of В. aggr. pumpelliana (Scribn.) Holub. the Kuban' River basin and the coast of the Black Sea). It This is the first CN counting forB. austrosibirica. Most likely is related to B. sylvaticum (Huds.) P. Beauv., where 2n = 18 it has variable ploidy, like the species of this affinity. Further is also known by multiple authors. We present here the se studies are needed. 2n = 4x. cond CN counting for B. spryginii (the first was in Probato Bromopsis benekenii (Lange) Holub, 2n = 28 (AS) va, Kazanovsky, Rudyka et al. 2012). 2n = 2x. The CN of Russia, North Caucasus, Republic of Daghestan, Bui species related to B. sylvaticum is constant (x = 9), however nakskii Raion, 16 km W of Buinaksk town, near Manas-aul, Brachypodium is polybasic genus. 960 m alt., the foot of Gimrinskii mountain ridge, the tourist Bromopsis australis (Zherebina) Tzvelev et Prob., 2n = 28 (AS) centre “Termenlik”, beech forest, 18 Jun 1979, coll. N.S. Russia, East Siberia, Republic of Buryatia, outskirts of Probatova 5410: 42. This forest species is distributed from Ulan-Ude city, meadow with steppe vegetation near airport, Europe and Caucasus to West Siberia. The same CN 2n = 12 Sep 1989, coll. N.S. Probatova & V.P. Seledets 6934: 34. 28 was earlier revealed from Azerbaidzhan (Sokolovskaya & Earlier the specimen № 6934 was misidentified as «Bromop Probatova 1979 – as B. ramosa ssp. benekenii), but the report sis inermis” by Probatova, Sokolovskaya & Rudyka (1991). of 2n = 42 also exists in the literature (see in Agapova et al. This steppe species was described from the south of Ural 1993). 2n = 4x. and was revealed in European Russia, North Caucasus, West Bromopsis heterophylla (Klokov) Holub, *2n = 56 (AS) and East (w) Siberia. It can be found in the RFE as well (as Russia, Republic of Crimea, Ai-Petrinskaya yaila, on adventive), just like its closely relative B. inermis (Leyss.) Ho the slope, 13 Aug 1974, coll. V.V. Fedyaeva 3958: . The lub (which is considered as forest species everywhere, but 46
2 Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) Chromosome numbers of Poaceae from Russia steppe species, distributed in the south-east of Europe, but 1973; Zhukova & Petrovsky 1976; Yurtzev & Zhukova described from Crimea. Probably it shows variable ploidy: 1978; Petrovsky & Zhukova 1981: Chukotka, Wrangel before the newly revealed CN 2n = 56, the diploid CN 2n Isl.). In the RFE this species becomes rare, the CN reports = 14 was known from Crimea (see Prokudin et al. 1977 – for southern parts of RFE probably belong to D. paramu as “B. cappadocica”); however these cytotypes may belong to shirensis. 2n = 2x, 4x. different taxa. 2n = 8x. Deschampsia komarovii V.N. Vassil., 2n = 26 (ER) Bromopsis variegata (M. Bieb.) Holub, 2n = 14 (AS) Russia, the Far East, Kamchatskii Krai, Kamchatka Pe Russia, North Caucasus, Republic of Daghestan, Gu ninsula, Bystrinskii Raion, the bottom of temporary lake, nibskii Raion, near Verkhnii Gunib village, 1800 m alt., san 884 m alt., 19 Aug 2013, V.V. Buryi 12598: 11; Russia, Far dy-pebbly bank of the Gunibka River, 26 Jun 1979, coll. East, Kamchatskii Krai, Kamchatka Peninsula, Elizovskii N.S. Probatova 5420: 41. The species is distributed in the Raion, Mutnovskii Volcano, poorly matted slope, above the Russian Caucasus, in high mountains. The CN 2n = 14 was geysers plot “Dachnyi”, 20 Aug 2004, coll. A.N. Berkutenko known from Kabardino-Balkaria Republic (Sokolovskaya & & M.A. Polezhaeva 9670: 13. Riverside and coastal Probatova 1979). 2n = 2x. meadows, sands and pebbles. RFE (Arctic, Kamchatka, Okhotia). Endemic. 2n = 26, 38, 42 (Zhukova et al. 1973; Catabrosa minor (Bab.) Tzvelev (C. aquatica var. minor Bab.), 2n = 20 (AS) Yurtzev & Zhukova 1978: Chukotka - east, Wrangel Isl.). 2n = 2x and aneuploids. Russia, European Arctic, Komi Republic, in vicinity of Vorkuta city, the farm (sovkhoz) “Tsentral'nyi”, weedy Deschampsia macrothyrsa (Tatew. et Ohwi) Kawano, 2n place, 17 Jul 1967, coll. N.N. Tzvelev 63 (LE); A.P. Sokolov = 26 (ER) skaya 124: 2. This specimen was misidentified as “Catabrosa Russia, the Far East, Sakhalin (north), Ekhabi Bay, SE aquatica s. str.” (Sokolovskaya 1970; Sokolovskaya & Proba of Vostochnyi settlement, on the bank of a brackish lake, 3 tova 1975). C. minor was collected by Tzvelev in the same Aug 2001, coll. V.Yu. Barkalov 8606: 16. Coastal boggy mea expedition with Sokolovskaya, and she counted the CN on dows, swamps. RFE: Sakhalin, the Kurils (south), reported this specimen (but with her number 124). The correct name from Okhotia (south); East Asia (Hokkaido). Described of this plant was determined much later (Tzvelev 2013). from Hokkaido. 2n = 26 (Probatova 1984; Probatova et al. C. minor occurs near seacoasts in European Arctic, but in 2009: Sakhalin). 2n = 2x. Komi Republic, near Vorkuta, it is adventive. The related widespread species C. aquatica (L.) P. Beauv. does not occur Deschampsia obensis Roshev., * 2n = 26 (AS) in the Arctic. 2n = 4x. Russia, Komi Republic, Intinskii Raion, the farm (sov khoz) “Gornyak”, the farm Yun’-Yaga, the Yun’-Yaga Ri * Deschampsia baicalensis Tzvelev, 2n = 26 (ER) ver, along sandbanks, 12 Jul 1967, coll. A.P. Sokolovskaya Russia, East Siberia, Irkutskaya Oblast’, Ol’khonskii 77: 1. Earlier this specimen was misidentified as “D. caespi Raion, W lakeside of Baikal Lake, the Khokhe-Nokhoitui tosa subsp. orientalis” (Sokolovskaya & Probatova 1975). Cape, 496 m alt., on bank of the swampy lakelet behind the Sandy and pebble banks of river valleys in lower courses, lakeside barrage, 28 Jul 2005, coll. S.G. Kazanovsky 11600: poorly fixed and mobile sands of rivers and near seacoast. 30; Russia, East Siberia, Irkutskaya Oblast’, Ol’khonskii East Europe (Arctic, Kola Peninsula, Pechora River, Ural Raion, near Chernorud settlement, the Kuchulga River, – north); West Sibe ria (Arctic and lower course of the Ob’ 464 m alt., former river-bed, the lowland sedge marsh, 1 River); East Sibe ria (Arctic); RFE (Arctic – west). Endemic. Aug 2004, coll. S.G. Kazanovsky 11321: 31. Sandy and 2n = >42, >44, 52 (Zhukova & Petrovsky 1975; Zhukova muddy banks of river deltas. East Siberia (the Baikal Lake). & Petrovsky 1980: Chukotka). 2n = 2x, 4x. Endemic. Probably the species was originated from hybri dization between D. turczaninowii (Litv.) Roshev. and D. suka Deschampsia pamirica Roshev., 2n = 26 (AS) tschewii (Popl.) Roshev. This is the first chromosome count Russia, West Siberia, Altai, Chuyskaya steppe, Kosh- for the species. 2n = 2x. Agach, 1835 m alt., along the riverside, 25 Jul 1937, coll. A.P. Sokolovskaya & O.S. Strelkova 26: 38; Russia, West Deschampsia glauca Hartm., 2n = c. 26 (AS) Siberia, Altai, the Kuraiskii mountain ridge, Tabozhek ca Russia, East Siberia, Republic of Sakha-Yakutia, lower nyon, 2300 m alt., on placer deposits, 30 Jul 1937, coll. course of the Lena River, in vicinity of the Tiksi Bay, A.P. Sokolovskaya & O.S. Strelkova 73: 37; Russia, West on stony plots of riverside of a small river, 1 Aug 1948, Siberia, Altai, the Chegan-Uzun River, 2000 m alt., pebbles coll. A.P. Sokolovskaya 129: 4. Earlier this specimen was on the slope, 5 Aug 1937, coll. A.P. Sokolovskaya & O.S. misidentified as “D. caespitosa subsp. borealis” by Sokolovskaya Strelkova 125: 39; Russia, West Siberia, Altai, Katunskie & Probatova (1975). In tundras, meadows, on pebbles and Belki, the Kazinikha Pass, 2400 m alt., 20 Aug 1937, coll. stony slopes, by the streams. East Europe (from Arctic to A.P. Sokolovskaya & O.S. Strelkova 165: 40. Earlier all these Ural – north and middle), West Siberia (Arctic, Ob’ River specimens were misidentified as D.“ caespitosa subsp. koeleri – as adventive), East Siberia (Arctic, Putorana Mts., Lena oides” in Sokolovskaya & Probatova (1975). Saline meadows, and Kolyma – north), RFE (Arctic, Anadyr’, Penzhina); sands and pebbles; in the middle and upper mountain belts. Europe (Scandinavia), North America (Alaska). (Tzvelev & West Siberia (Altai), East Siberia (West Sayan); Middle and Probatova 2012). – 2n = 26, 48, 52 (Zhukova & Tikhonova Central Asia. 2n = 2x.
Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) 3 Probatova et al.
Deschampsia paramushirensis Honda (D. cespitosa (L.) south of continental Far East (the Amur River basin). The P. Beauv. subsp. orientalis Hultén), 2n = 26 (AS, ER) revision of the whole material studied as to CNs is needed. Russia, the Far East, Sakhalin (north), near the former 2n = 2x, 4x. settlement Muz’ma, N of Pomr’ Bay, 10 Aug 2001, coll. Elymus kamoji (Ohwi) S.L. Chen (Agropyron tsukushiense V.Yu. Barkalov 8613: 18; Russia, the Far East, the North (Honda) Ohwi var. transiens (Hack.) Ohwi), 2n = 42 (ER) Kurils, Paramushir Island, outskirts of Severo-Kuril’sk set Russia, the Far East, Primorskii Krai, Khassanskii Rai tlement, on pasture, 20 Aug 1971, coll. N.S. Probatova 3351: on, Slavyanka town, a meadow near the sea front, 19 Aug 16; Russia, the Far East, Khabarovskii Krai, Tuguro-Chumi 2012, coll. A.V. Agafonov 12719: 28. East Asian species, kanskii Raion, the Shantarskie Islands, Feklistova Island, N described from the North-East China. It was revealed re coast, Lissya Bay, Lissje Lake, at the rivulet, 20 Aug 2010, cently in the RFE (Primorskii Krai, Khassanskii Raion), coll. V.V. Bogatov 12326: 20; Russia, the Far East, Sakha firstly – from Ryazanovka settlement (Tzvelev & Probatova lin (north), outskirts of Moskalvo settlement, by the lake, 2010 b). This is the second record of the species for Rus Aug 2001, coll. V.Yu. Barkalov 8601: 17. Earlier the speci sia. The hexaploid CN 2n = 42 is typical for two closely men № 3351 was misidentified as “D. cespitosa s. str.” (So related East Asian species – E. kamoji (Nishikawa 2008) kolovskaya & Probatova 1975) and the specimen № 12326 and E. tsukushiensis Honda, but uncommon in the section as “D. glauca” (Probatova et al. 2014). Riverside and seaside Goulardia. In the RFE E. tsukushiensis was found in Kunashir meadows, sands and pebbles. RFE (Anadyr’ River basin, Isl., South Kurils (2n = 42 – Probatova et al. 2000). 2n = 6x. North Korya kia, Kamchatka, Okhotia, Sakhalin – north, Elytrigia gracillima 2n = 28 North Kurils); East Asia (Hokkaido). Some reports of vari (Nevski) Nevski, (AS) ous CNs for D. paramushirensis (also under other names, e.g., Russia, North Caucasus, Republic of Daghestan, Gunib its synonym D. cespitosa subsp. orientalis – see in Agapova et skii Raion, near Verkhnii Gunib village, Khotochinskii al. 1993) hardly belong to this species, which obviously oc mountain ridge, ca. 2000 m alt., on the rocks, 26 Jun 1979, curs in oceanic (coastal) regions of NE Asia. The revision coll. N.S. Probatova 5424: 43. On limestone rocks and of the whole material studied is needed. 2n = 2x. stony slopes, in lower and middle mountain belts. Caucasian species; described from Daghestan. This is the second CN *Deschampsia submutica (Trautv.) O.D. Nikif., 2n = 26 counting from Russia, also from Daghestan (2n = 28 – Pro (P.G. Zhukova – LE, VLA) batova & Sokolovskaya 1978). 2n = 4x. Russia, Far East, Chukotskii natsionalnyi okrug, E coast Elytrigia repens (L.) Nevski, 2n = 42 (ER) of the Chaunskaya Guba [Gulf], Apapelkhino settlement, 2 Sep 1965, V.V. Petrovsky & P.G. Zhukova 65–436: 5. Sands Russia, the Far East, Primorskii Krai, Mikhailovskii Rai and pebbles on riversides, wet meadows, among shrubs. on, outskirts of Novoshakhtinskii town, as a weed in ve East Europe (Arctic, Ural – north), West Siberia (Arctic, getable garden, 27 Jul 2004, coll. V.T. Lapenko 11676: 27. Ob' River basin – north), East Siberia (Arctic – south, Yeni Meadows, forest clearings, on sands and pebbles, often as ssei north, Lena and Kolyma), RFE (Chukotka – west). En a weed. Holarctic. Everywhere in Russia, but alien in the demic. 2n = 2x. RFE. Multiple CN data published, almost all of them – 2n = 42. 2n = 6x. Deschampsia sukatschewii (Popl.) Roshev., 2n = 26 (AS, ER) Elytrigia stipifolia (Czern. ex Nevski) Nevski, 2n = 28 (AS, ER) Russia, East Siberia, Irkutskaya Oblast', Irkutskii Rai Russia, North Caucasus, Republic of Daghestan, Gu on, Baikal Lake, the mouth of the Bol'shaya Goloustnaya nibskii Raion, near Kharti-Kuni village, on the slope with River, along the lakeside barrage at the lagoon lake, 4 Sep Pinus kochiana, at the pass, 16 Jul 1975, coll. N.S. Probatova 2003, coll. V.V. Chepinoga & al. 9098: 32; Russia, the Far & V.P. Seledets 4058: 39. Earlier this specimen was misiden East, North Koryakia, Olyutorskii Raion, 2.5 km S of tified asElytrigia “ tauri subsp. pertenuis“ in Probatova & Khailino settlement, on pebbles of a rivulet, 16 Aug 1970, Sokolovskaya (1978). On exposed chalk and limestone, in coll. N.S. Proba tova & V.P. Seledets 2655: 7; Russia, the Far steppes, up to the lower mountain belt. East Europe, Cau East, Sakhalin, Makarovskii Raion, near the railway station casus. 2n = 28 (see Prokudin et al. 1977). 2n = 4x. Pugachevo, on the railroad embankment, abundant, 15 Sep 1982, coll. N.S. Probatova, V.P. Seledets & al. 6270: 19. The Elytrigia trichophora (Link) Nevski, * 2n = 28 (ER) specimen № 2655 earlier was misidentified as “D. caespitosa Russia, North Caucasus, Krasnodarskii Krai, Novoros subsp. orientalis” in Sokolovskaya & Probatova (1975). Sands siyskii Raion, in vicinity of Raevskaya settlement, Volchye and pebbles on riversides, forest clearings, forest edges; up Vorota Pass, rubbly marl placer, 3 Sep 2009, coll. N.S. Pro to the middle mountain belt. East Europe (Arctic, Pechora batova & V.P. Seledets 11612: 45. In steppes, stony slopes, and Mezen’ rivers); West Siberia (Arctic, the Ob’ River, Al among shrubs, up to lower mountain belt. East Europe, tai), East Siberia, RFE; Asia (Central – north) and E (north Caucasus, Central Asia. 2n = 42 (see Prokudin et al. 1977; west). 2n = 26, 28, 42, 42+1B, 52 (see Agapova et al. 1993; Sokolovskaya & Probatova 1978). New cytotype for the Zhukova et al. 2009; Chepinoga et al. 2010; Probatova et al. species. 2n = 4x, 6x. 2014). Many of the multiple reports with various CN for * Festuca chionobia T.V. Egorova et Sipliv. (F. auriculata subsp. D. sukatschewii (including under other names, e.g., D. cespito chionobia (T.V. Egorova et Sipliv.) Tzvelev), 2n = 14 (ER) sa subsp. orientalis) hardly belong to this continental forest Russia, the Far East, Khabarovskii Krai, Nanaiskii Raion, species, which occurs for certain in East Siberia and in the Sikhote-Alin’ mountain range, the Tardoki-Yani Mt., upper
4 Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) Chromosome numbers of Poaceae from Russia course of Sukhaya Pad’ stream (the Anjui River basin), 1900 (east); Korean Peninsula. 2n = 14 (Probatova & Sokolovs m alt., alpine belt, stony dwarfshrub-mossy tundra, 29 Aug kaya 1984; Alexeev et al. 1987). 2n = 2x. 2012, coll. V.Yu. Barkalov 12189: 22. Mountain tundras and Festuca rubra L., 2n = 42 (ER) meadows. Baikal Siberia, RFE. Endemic. Described from Russia, the Far East, Primorskii Krai, Peter the Great the Baikal’skii mountain ridge. In RFE it occurs mainly in Bay, Rechnoi Island (Amurskii Gulf), on coastal rocks, 14 high altitudes of Sikhote-Alin’ and also in the mountains Aug 2010, coll. V.A. Nechaev 11659: . Extremely poly of the Upper Amur River basin. The species belongs to 26 morphous species, widely distributed throughout the world, F. aggr. auriculata Drobow. For F. auriculata there are many with multiple CN data. In F. rubra most common is the CN CN reports of 2n = 14 only, from Chukotka, Wrangel Isl. 2n = 42 (6x), which was revealed throughout Russia as well, and East Siberia (see Agapova et al., 1993). Another related including the RFE; rarely 2n = 56 is reported. The speci species is F. mollissima V.I. Krecz. et Bobrov, also with 2n = men from Rechnoi Island represents the “tuft form” of 14 (many CN counts from Primorskii Krai; see in Probatova, Festu ca rubra L. which is common on seacoastal rocks and 2014). F. chionobia is a pseudoviviparous species. (Fig. 2). stony slopes of marine terraces in continental part and in This is the first CN counting forF. chionobia. Obviously it is the islands of the Primorskii Krai (s.); its CN was firstly re the first pseudoviviparous species with diploid CN 2n = 14. vealed. 2n = 6x. Other festucoid grasses with pseudoviviparity (Deschampsia, Festuca, Poa) are polyploids and aneuploids: see the review * Festuca vylzaniae (E. Alexeev) Tzvelev, 2n = 28 (ER) of Sarapul’tsev (2001). Russia, East Siberia, Republic of Buryatia, Pribaikal’skii * Festuca dahurica (St.-Yves) V.I. Krecz. et Bobrov, 2n = Raion, coast of Baikal Lake, in vicinity of Turka settlement, 14 (ER) on sands, 3 Jul 2010, coll. A.A. Gnutikov 12236: 33. Endemic Russia, East Siberia, Republic of Buryatia, Mukhorshi of the Baikal Lake region. Sandy-steppe species. This is the birskii Raion, near Olon-Sheber Lake, on sands, 3 Jul 2010, first CN count for the species. 2n = 4x. coll. A.A. Gnutikov & Yu.A. Gnutikova 12259: 35. Mostly Milium effusum L., 2n = 28 (ER) Chinese species; occurs on rocks and steppes. In Russia it is Russia, the Far East, Kamchatskii Krai, Kamchatka Pe distributed in the south of Transbaikalia. Its CN was revea ninsula, Verkhne-Opal‘skie hot springs, t = 42°C, on the led for the first time. 2n = 2x. slope, in Filipendula kamtschatica community, 3 Aug 2013, Festuca jacutica Drobow, 2n = 14 (ER) coll. O.A. Chernyagina & L. Shtreker 12457: 14. In forest Russia, the Far East, Khabarovskii Krai, Poliny Ossi edges and clearings, tall herbs communities, among shrubs. penko Raion, the valley of the Nylan River (left tributary European Russia, Caucasus, West and East Siberia, RFE of the Amgun’ River), middle course, stony outcrops (up to Kamchatka in the north). Almost Holarctic. 2n = 28 with limestone “Mramornye skaly” on the right riverside, (multiple CN reports from throughout Russia, and from va fissures in the rocks, 25 Jul 2012, coll. M.V. Kriukova & rious subregions of the RFE). First CN count from Kam L.A. Antonova 12680: 21. In light forests, among shrubs, chatka. 2n = 4x. on stony slopes and rocks. East Siberia, RFE (Kolyma, Poa alpigena (Blytt) Lindm. × P. arctica R. Br., 2n = 42 (ER) Okhotia, the Amur River basin); Northeast China. De Russia, the Far East, Kamchatskii Krai, Aleutskii Raion, scribed from Yakutia. 2n = 14 (Yurtzev & Zhukova 1982; Commander Islands, Bering Island, upper course of the Probatova & Sokolovskaya 1984). First CN data from Vodopadnaya River, wet forb meadow along the lakeside, Khabarovskii Krai. 2n = 2x. 15 Aug 1999, coll. O.A. Mochalova 8085: 9. Both widely *Festuca kolesnikovii Tzvelev, 2n = 14 (ER) disributed Holarctic species P. alpigena and P. arctica are com Russia, the Far East, Primorskii Krai, Terneiskii Raion, mon plants in the Kamchatskii Krai, they occur in mea Sikhote-Alinskii biosphere reserve, locality Blagodatnoe, dows, among riverside shrubs, on sands and pebbles of ri marine terrace, dry meadow with steppe features, 25 Jun ver valleys, on roadsides. Such hybrid is not rare, the plants 2003, coll. I.A. Nesterova 9023: 23; Russia, the Far East, have some pubescence in lower part of the lemma between Primorskii Krai, Peter the Great Bay, Lissyi Island (opposite veins or along veins, but at the same time they look like Nakhodka city), 11 Sep 2002, coll. V.A. Nechaev 8901: 25. P. alpigena. 2n = 6x. On costal rocks, stony slopes and pebbles. RFE: Primorskii Poa angustifolia L., 2n = 70 (AS) Krai (east macroslope of Sikhote-Alin’ mountain ridge). Russia, the Far East, Kamchatskii Krai, Koryakskii Endemic. Most probably this species has hybrid origin: Natsional'nyi Okrug, Oliutorskii Raion, near Korf settle F. aggr. ovina L. × F. aggr. auriculata Drobow. First CN data ment, marine spit, middle part, grass community on sands, for the species. 2n = 2x. 17 Sep 1970, coll. N.S. Probatova & V.P. Seledets 3100: 8. Festuca mollissima V.I. Krecz. et Bobrov, 2n = 14 (ER) Earlier this specimen was misidentified as “Poa alpigena” in Russia, the Far East, Primorskii Krai, Partizanskii Raion, Sokolovskaya & Probatova (1973). Dry meadows, riverside Alexeevskii mountain ridge, Olkhovaya Mt., on the spots of and coastal sands and pebbles, roadsides. European Rus fine-grain soil, 1600 m alt., 10 Sep 2010, coll. V. Yu. Bar sia, Caucasus, West and East Siberia, RFE (but alien on the kalov 11667: 24. On stony slopes and rocks, sometimes north, in Sakhalin and in the Kurils); Eurasian. In the North on coastal sands and pebbles, up to the middle mountain Koryakia P. angustifolia is alien. 2n = 56, 63–64, 70–72 (from belt. RFE (Okhotia – Amgun’, Amur (east), Primorskii Krai some regions of Russia, including RFE). 2n = 8x, 9x, 10x.
Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) 5 Probatova et al.
Figure 2 Specimen of pseudoviviparous Festuca chionobia with diploid CN 2n = 14, collected in study area 22 (see Fig. 1).
6 Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) Chromosome numbers of Poaceae from Russia
Poa glauca Vahl, 2n = 42 (ER) phenomenon obviously indicates progressive species. The Russia, the Far East, Amurskaya Oblast’, Zeyskii nature tetra ploid CNs (2n = 4x) show the evolutionary optimal reserve, E part of Tukuringra mountain ridge, alpine zone, ploidy level for species in their taxonomic groups. The he 15 Jul 2011, coll. T.N. Veklich 12102: 29. Mountain tundras, xaploid CN (2n = 6x) may indicate the hybrid origin of rocks, screes, dry pebbles in the Arctic and alpine zone of the taxa. 9 species were studied caryologically for the first mountains. European Russia, Caucasus, West Siberia, East time, besides for 3 species the new CN data were obtained. Siberia, RFE; Holarctic. 2n = 42, с.48, 48, 49, 50, 56, 62, 63, Further CN studies on the Poaceae family in Russia are in 70 (multiple CN reports from Russia and elsewhere). The process. CN of the species is studied in Amurskaya Oblast' (the Up ACKNOWLEDGEMENTS per Amur River basin) for the first time. 2n = 6x, etc. The study was supported by Russian Foundation for Poa kamczatensis ., 2n = 56 Prob (ER) Basic Research (RFBR): grants 04-04-49750, 07-04-00610, Russia, the Far East, Kamchatskii Krai, Kamchatka Pe 11-04-00240 to N.S. Probatova and grant 12-04-01586 to ninsula, near Petropavlovsk-Kamchatskii city, Bab‘ya Bay, A.V. Verkhozina. The authors are grateful to E.G. Rudyka the slope of marine terrace, on rocks, 1 Aug 2014, coll. for chromosome countings in several species, to N.N. Tzve O.A. Chernyagina & V.Yu. Neshataeva 12713: 12. On rubb lev – for his help in some species identifications, also to ly slopes of marine terraces, on rocks and screes usually of A.V. Agafonov, L.A. Anto nova, V.V. Buryi, V.V. Chepinoga, coastal hills, up to the lower mountain belt. RFE (Kamchat O.A. Chernyagina, A.A. Gnutikov, S.G. Kazanovsky, M.V. ka – south, Okhotia (near Magadan). Endemic. 2n = 49, 56 Krjukova, O.A. Mochalova, V.A. Nechaev, I.A. Nesterova, (Sokolovskaya et al. 1985). P. kamczatensis is relative to P. al T.N. Veklich – for providing plant material for our study. masovii Golub: most probably, these species were originated from the same (or close) species of sections Malacanthae and LITERATURE CITED Stenopoa. Now they both belong to section Poastena. 2n = Agapova, N.D., K.B. Arkharova, L.I. Vakhtina, E.A. Zems 7x, 8x. kova & L.V. Tarvis 1993. Chromosome numbers in flowering * Poa khokhrjakovii Prob., 2n = c. 70 (AS) plants of the flora of the USSR: Moraceae – Zygophyllaceae. Na uka, St.-Petersburg, 430 pp. (in Russian). [Агапова Н.Д., Russia, the Far East, Kamchatskii Krai, Koryakskii na Архарова К.Б., Вахтина Л.И., Земскова Е.А., Тарвис tsio nal’nyi okrug, Penzhinskii Raion, 3 km S of Kamenskoe Л.В. 1993. Числа хромосом цветковых растений фло settlement, rubbly slope of a hill, the upper one third, 22 ры СССР: cемейства Moraceae–Zygophyllaceae. СПб: Нау Jul 1971, coll. N.S. Probatova & V.P. Seledets 3291: 6. Dry ка. 430 с.]. stony slopes, in the middle and upper mountain belts. RFE Alexeev, E.B., A.P. Sokolovskaya & N.S. Probatova 1987. Taxonomy, distribution and chromosome numbers in (the basins of Penzhina, Omolon, Kolyma rivers). Ende Festu ca L. (Gramineae) of the flora of USSR. 2. Sec mic. Closely relative to P. malacantha Kom., but differs by tion Festu ca: F. djimilensis – F. lenensis. Bulleten’ Moskovskogo narrow leaf blades, loose tufts with acuate subterranean Obshchestva Ispytatelei Prirody. Otd. biol. 92(5): 122–132 (in shoots, the sheats of old leaves thin, light (and not thick, Russian). [Алексеев Е.Б., Соколовская А.П., Пробато brownish), panicle branches very thin, smooth, lemmas al ва Н.С. 1987. Таксономия, распространение и числа хромосом овсяниц ( L., Gramineae) флоры most glabrous between veins. First CN count for the spe Festuca CCCР. 2. Cекция Festuca: F. djimilensis – F. lenensis // cies. 2n = 10x. Бюл. Моск. общ. испыт. прир. Отд. биол. Т. 92, вып. * Poa krasnoborovii Stepanov, 2n = 42 (ER) 5. С. 122–132]. Russia, East Siberia, Krasnoyarskii Krai, Ermakovskii Chepinoga, V.V., A.A. Gnutikov & I.V. Enushchenko 2010. Chromosome numbers of some plant species from the Raion, the Us River above the mouth of the Tsap River and south part of the East Siberia. Botanicheskii Zhurnal 95(1): below the Verkhn’aya Buyba River, the Pinus and Picea fo 129–130 (in Russian). [Чепинога В.В., Гнутиков А.А., rest plots on the riverside, 19 Aug 2010, coll. N.V. Stepanov Енущенко И.В. Числа хромосом некоторых видов 12490: 36. This species, perhaps endemic, is concerned with растений из южной части Восточной Сибири // Бо coniferous forests of the West Sayan Mts, where it occurs танический журнал. Т. 95, № 1. С. 129–130]. in tallgrass clearings. The species belongs to the Poa pratensis Nishikawa, T. (ed.) 2008. Chromosome atlas of flowering plants in Japan. Tokyo, 706 pp. group, plants are tall, leaf blades up to 8 mm width, more or Petrovsky, V.V. & P.G. Zhukova 1981. Chromosome num less rigid, ligules up to 5 mm, panicles up to 25 sm, oblong, bers and taxonomy of some plant species of the Wrangel their branches rather long, spikelets up to 8.5 mm. This is Island. Botanicheskii Zhurnal 66(3):380–387 (in Russian). the first CN count for the species. 2n = 6x. [Петров ский В.В., Жукова П.Г. 1981. Хромосомные числа и таксономия некоторых видов растений ост CONCLUSION рова Врангеля // Ботанический журнал. Т. 66, № 3. С. 380–387]. The 35 species presented here demonstrate the diversity of CNs: different ploidy levels, a series of basic CN numbers Probatova, N.S. 1984. New taxa of Poaceae from the Far East of the USSR. Botanicheskii Zhurnal 69(2):251–259 (in (2x – 10x; x = 7, 9, 13). The diploids prevail (14 species, ca. Russian). [Пробатова Н.С. 1984. Новые таксоны сем. 40 %), but only 7 species among them are primary diploids Poaceae с Дальнего Востока СССР // Ботанический (2n = 14, x = 7), namely in Festuca, Bromopsis, Beckmannia. журнал. 1984. Т. 69, № 2. С. 251–259]. Also there are species with variable ploidy (sometimes – Probatova, N.S. 2014. Chromosome numbers in vascular plants expected), especially in Poa and in some Deschampsia. This of the Primorskii Krai (Russian Far East). Dal’nauka, Vladi
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vostok, 343 pp. (in Russian). [Пробатова Н.С. 2014. Sokolovskaya, A.P. 1970. Karyological study on the flora of Хромосомные числа сосудистых растений Приморс the Ussa River basin (Komi ASSR). Vestnik Leningradskogo кого края (Дальний Восток России). Владивосток: Universiteta, Biol. 2(9):106–114 (in Russian). [Соколовская Дальнаука, 343 с.]. А.П. 1970. Кариологическое исследование флоры Probatova, N.S., V.Yu. Barkalov, E.G. Rudyka & Z.V. Ko бассейна р. Усы (Коми АССР) // Вестник Ленингр. zhevnikova 2009. Additions to chromosome numbers for унив., Биол. Вып. 2, № 9. С. 106–114]. vascular plants from Sakhalin and the Kurile Islands (1). Sokolovskaya, A.P. & N.S. Probatova 1973. Karyotaxono In: Biodiversity and Biogeography of the Kuril Islands and Sakha mic study on the Far East species of Poa L. II. Botanicheskii lin (H. Takahashi & M. Ohara, eds.), issue 3, pp. 35–47, Zhurnal 58 (1): 89–96 (in Russian). [Соколовская А.П., Hokkaido University Museum, Sapporo, Japan. Пробатова Н.С. 1973. Кариосистематическое иссле Probatova, N.S., V.Yu. Barkalov, E.G. Rudyka & S.A. Shatalo дование дальневосточных видов Poa L. II // Бота ни va 2000. Chromosome study on vascular plants of the Ku ческий журнал. Т. 58, № 1. С. 89–96]. rile islands. Natural History Research 7:21–38. Sokolovskaya, A.P. & N.S. Probatova 1975. Chromo Probatova, N.S., S.G. Kazanovsky, E.G. Rudyka, V.P. Seledets some numbers of some grasses (Poaceae) of the flora of & V.A. Nechaev 2012. IAPT/IOPB chromosome data 13 USSR. Botanicheskii Zhurnal 60(5):667–678 (in Russian). (K. Marhold, ed.). Taxon 61(4):899–902, E 34-42. [Соколовская А.П., Пробатова Н.С. 1975. Хромосом ные числа некоторых злаков (Poaceae) флоры СССР // Probatova, N.S., A.A. Korobkov, A.A. Gnutikov, E.G. Ru Ботанический журнал. Т. 60, № 5. С. 667–678]. dyka, V.V. Kotseruba & V.P. Seledets. 2010 IAPT/IOPB chromosome data 10 (K. Marhold, ed.). Taxon 59(6): Sokolovskaya, A.P. & N.S. Probatova 1978. Chromosome 1934–1937, E 6–10. numbers of some grasses (Poaceae) of the flora of USSR. II. Botanicheskii Zhurnal 63(9):1247–1257 (in Russian). Probatova, N.S., E.G. Rudyka, V.P. Seledets & T.N. Moto [Соколовская А.П., Пробатова Н.С. 1978. Ботани чес rykina 2014. Chromosome numbers in vascular plants кий журнал. Т. 63, № 9. С. 1247–1257]. from the Russian Far East: Amurskaya Oblast’, Khabarov skii Krai, Primorskii Krai. Botanica Pacifica 3(2):129–134. Sokolovskaya, A.P. & N.S. Probatova 1979. Chromosome numbers of some grasses (Poaceae) of the flora of USSR. Probatova, N.S. & V.P. Seledets 2008. IAPT / IOPB chro III. Botanicheskii Zhurnal 64(9):1245–1258 (in Russian). mosome data 5 (K. Marhold, ed.). Taxon 57(2):555–558, [Соколовская А.П., Пробатова Н.С. 1979. Ботаничес E 7–16. кий журнал. Т. 64, № 9. С. 1245–1258]. Probatova, N.S. & A.P. Sokolovskaya 1978. Chromosome Sokolovskaya, A.P., N.S. Probatova & E.G. Rudyka 1985. numbers and taxonomy of some grasses from Cauca Chromosome numbers in species of the families Astera sus. Botanicheskii Zhurnal 63(8):1121–1131 (in Russi an). ceae, Poaceae, Rosaceae from Primorskii Krai, Kamchatka and [Пробатова Н.С., Соколовская А.П. 1978. Хромосом Sakhalin. Botanicheskii Zhurnal 70(1):126–128 (in Russian). ные числа и таксономия некоторых злаков Кавказа // [Соколовская А.П., Пробатова Н.С., Рудыка Э.Г. 1985. Ботанический журнал. Т. 63, № 8. С. 1121–1131]. Числа хромосом видов семейств Asteraceae, Poaceae, Ro Probatova, N.S. & A.P. Sokolovskaya 1980. Contribu saceae из Приморского края, Камчатки и Сахалина // tion to karyotaxonomic study on Poaceae of Gornyi Ботанический журнал. Т. 70, № 1. С. 126–128]. Altai. Botanicheskii Zhurnal 65(4):509–520 (in Russian). Tzvelev, N.N. 2013. Notes on some genera of the Family Poa [Пробатова Н.С., Соколовская А.П. 1980. К кариотак ceae. Novitates systematicae plantarum vascularium 44:26–38 (In сономическому изучению злаков Горного Алтая // Russian). [Цвелëв Н.Н. 2013. Заметки о некоторых родах Ботанический журнал. Т. 65, № 4. С. 509–520]. семейства злаков (Poaceae) // Новос ти систематики Probatova, N.S. & A.P. Sokolovskaya 1984. Chromosome высших растений. СПб. Т. 44. С. 26–38]. numbers in representatives of the families Butomaceae, Tzvelev, N.N. & N.S. Probatova 2010 a. New taxa of grasses Papaveraceae, Poaceae from the Far East of the USSR. Bo (Poaceae) of Russia. Botanicheskii Zhurnal 95(6):857–869 (in tanicheskii Zhurnal 69(3):410–412 (in Russian). [Пробато Russian). [Цвелëв Н.Н., Пробатова Н.С. 2010. Новые ва Н.С., Соколовская А.П. 1984. Числа хромосом таксоны злаков (Poaceae) России // Ботанический предста ви телей семейств Butomaceae, Papaveraceae, Poaceae журнал. Т. 95, № 6. С. 857–869]. c Дальнего Востока СССР // Ботанический журнал. Т. 69, № 3. С. 410–412]. Tzvelev, N.N. & N.S. Probatova 2010 b. The genera Ely mus L., Elytrigia Desv., Agropyron Gaertn., Psathyrostachys Probatova, N.S., A.P. Sokolovskaya & E.G. Rudyka 1991. Nevski and Leymus Hochst. (Poaceae: Triticeae) in the Chromosome numbers of some vascular plant species flora of Russia.Komarovskie Chtenya 57:5–102 (in Russian). from the Far East and other regions of the USSR. Botani [Цвелев Н.Н., Пробатова Н.С. 2010. Роды Elymus L., cheskii Zhurnal 76(8):1174–1178 (in Russian). [Пробатова Elytrigia Desv., Agropyron Gaertn., Psathyrostachys Nevski и Н.С., Соколовская А.П., Рудыка Э.Г. 1991. Числа хро Leymus Hochst. (Poaceae: Triticeae) во флоре России // мосом некоторых видов сосудистых растений Даль Комаровские чтения. Вып. 57. C. 5–102.]. него Востока и других регионов СССР // Ботаничес кий журнал. Т. 76, № 8. С. 1174–1178]. Tzvelev, N.N. & N.S. Probatova 2012. The survey of genera Deschampsia, Agrostis, Calamagrostis (Poaceae – Poeae) and the Prokudin, Yu.N., A.G. Vovk, O.A. Petrova, E.D. 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8 Botanica Pacifica. A journal of plant science and conservation. 2015. 4(1) Chromosome numbers of Poaceae from Russia
Zhukova, P.G. & V.V. Petrovsky 1975. Chromosome num которых видов растений Сибири и Дальнего Востока bers in some plant species from the West Chukotka. // Ботанический журнал. Т. 58, № 9. С. 1331–1342]. Botanicheskii Zhurnal 60(3):395–401 (in Russian). [Жуко Zhukova, P.G. & A.D. Tikhonova 1973. Chromosome num ва П.Г., Петровский В.В. 1975. Хромосомные числа bers in some plant species from Chukotka. II. Botanicheskii неко торых видов растений Западной Чукотки // Бо Zhurnal 58(3):395–402 (in Russian). [Жукова П.Г., Тихо танический журнал. Т. 60, № 3. С. 395–401]. нова А.Д. 1973. Хромосомные числа некоторых видов Zhukova, P.G. & V.V. Petrovsky 1976. Chromosome num растений Чукотки. II // Ботанический журнал. Т. 58, bers in some plant species from West Chukotka. II. Botani № 3. С. 395–402]. cheskii Zhurnal 61(7):963–969 (in Russian). [Жукова П.Г., Yurtzev, B.A. & P.G. Zhukova 1978. Cytotaxonomical sur Петровский В.В. 1976. Хромосомные числа некото vey of Monocots in the east of Chukotka Peninsula. рых видов растений Западной Чукотки. II // Бота ни Botanicheskii Zhurnal 63(8):1132–1144 (in Russian). [Юр ческий журнал. 1976. Т. 61, № 7. С. 963–969]. цев Б.А., Жукова П.Г. 1978. Цитотаксономический Zhukova, P.G. & V.V. Petrovsky 1980. Chromosome num обзор однодольных востока Чукотского полуострова bers and taxonomy of some species of the Anyui Mts. Bota // Ботанический журнал. Т. 63, № 8. С. 1132–1144]. nicheskii Zhurnal 65(5):651–659 (in Russian). [Жукова П.Г., Yurtzev, B.A. & P.G. Zhukova 1982. Chromosome numbers Петровский В.В. 1980. Хромосомные числа и таксо но in some plants from the north-east of Yakutia (the ba мия некоторых видов растений Анюйского нагорья // sin of middle course of the Indigirka River). Botanicheskii Ботанический журнал. Т. 65, № 5. С. 651–658]. Zhurnal 67(6):778–787 (in Russian). [Юрцев Б.А., Жу Zhukova, P.G., V.V. Petrovsky & T.V. Plieva 1973. Chromo кова П.Г. 1982. Хромосомные числа некоторых рас some numbers and taxonomy of some plant species from тений северо-восточной Якутии (бассейн среднего Siberia and the Far East. Botanicheskii Zhurnal 58(9):1331– тече ния реки Индигирки) // Ботанический журнал. 1342 (in Russian). [Жукова П.Г., Петровский В.В., Пли Т. 67, № 6. С. 778–787]. ева Т.В. 1973. Хромосомные числа и таксономия не
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