A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia Author Yurtseva, Olga V. 0000-0002-6770-4358 Department of Higher Plants, Faculty of Biology, M. V. Lomonosov Moscow State University, 1 - 12, Leninskie Gory, 119234, Moscow, Russia; e-mail: olgayurtseva @ yandex. ru & olgayurtseva @ yandex. ru; https: // orcid. org / 0000 - 0002 - 6770 - 4358 olgayurtseva@yandex.ru Author Vasilieva, Natalia V. 0000-0002-4060-7988 N. V. Tsitsin’s Main Botanical Garden of the Russian Academy of Sciences, 4 Botanicheskaya Ulitsa, 127276, Moscow, Russia & dekanat. vas @ mail. ru; https: // orcid. org / 0000 - 0002 - 4060 - 7988 dekanat.vas@mail.ru Author Kostikova, Vera A. 0000-0002-5213-2166 Central Siberian Botanical Garden SB RAS, 101, Zolotodolinskaya Str., 630090, Novosibirsk, Russia & serebryakova-va @ yandex. ru; https: // orcid. org / 0000 - 0002 - 5213 - 2166 serebryakova-va@yandex.ru Author Samigullin, Tahir H. 0000-0002-9135-3367 A. N. Belozersky Research Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 1 - 40, Leninskie Gory, 119992, Moscow, Russia & samigullin. t. h @ gmail. com; https: // orcid. org / 0000 - 0002 - 9135 - 3367 samigullin.t.h@gmail.com text Phytotaxa 2022 2022-09-22 566 1 13 63 http://dx.doi.org/10.11646/phytotaxa.566.1.2 journal article 148711 10.11646/phytotaxa.566.1.2 1442bcf8-8360-4373-a7dc-d34e82d487ba 1179-3163 7103418 Atraphaxis pungens , A. ledebourii sp. nov. and A. × uyukensis nom. provis. Our current results have shown that “ Atraphaxis pungens s.l. ” combines plants from Southern Siberia and Mongolia with three different plastid haplotypes, which replace each other in the area ( Figs 2 and 8 ). All the plants demonstrate a great similarity in the appearance and morphology of shoots, inflorescences, perianths and fruits, so it is impossible to discriminate them. They are shrubs 15–50 cm high ( Figs 9–11 , Table 1 ) with vigorous vegetative shoots, often sylleptically branching. Second-year shoots become durable, long lasting, perennial axes bear prickly branchlets. The covering of second-year shoots is dark-gray, almost black, the cork peels off, exposing the reddish bark. Wood is pale pinkish. Second-year shoots and their branchlets bear short ( 5–10 cm long) axillary pale-green or creamy annual shoots. These shoots are leafy and some terminate with thyrses. Annual short generative and vegetative shoots bear small oblanceolate or spathulate leaf blades with acute or obtuse apex, gradually narrowed to the base ( Fig. 9A–C ; Fig. 10A, C, D ; Fig. 11 ; Fig. 12A–B ; Fig. 13A , Fig. 14D, E ). Annual long vegetative shoots bear large obovate, oblongelliptic, broadly elliptic or rhomboid-elliptic leaf blades with acute or mucronate apex ( Fig. 9D–F ; Fig. 10B, E, F ; Fig. 12C ; Fig. 13B ; Fig. 14A ). The thyrses are usually compact, frondo-bracteose, consist of 4–7 cymes with 2–3 flowers. Perianth is yellowish-white or white, sometimes pinkish, with a very long perianth tube articulated to as long as pedicel ( Fig. 12H, I ; Fig. 13C ; Fig. 14B, F ). The epidermal cells of tepals and tube are similar ( Fig. 6C–F ). Atraphaxis pungens has black glossy fruits twice shorter than inner tepals of the perianth ( Fig. 12J–L ). Atraphaxis ledebourii has dark-brown to black fruits, 1.5 times shorter of inner tepals ( Fig. 13F, G, I ). In plants from Altai and Tuva , the ovary and stamens are often underdeveloped. In both species three styles are fused at the very base ( Fig. 6J, K , Fig. 12D–G , Fig. 13D, E, H ). In A. pungens and A. × uyukensis , stigmas are either flat and rounded, or elongated and bent out, later bevealed outward ( Fig. 12G, L ; Fig. 14C, G–I ). In A. ledebourii from Altai Republic , the stygmas are flat and rounded ( Fig. 13D ), in plants from Tuva Basin and Ubsunur Basin the stygmas are globular ( Fig. 6K ; Fig. 13E, H ). FIGURE 6. Epidermal cells of perianth tube (A, C, E), tepals of perianth (B, D, F), fruits (G, L), styles (H, J, K), and fruit surface (I). A, B— A. kuvaevii (154), RUSSIA. Krasnoyarskii Krai: Kuvaev & Sonnikova 1015-13 (MW0061320, holotype). C, D— A. pungens (255), RUSSIA. Khakassia: Askiz Raion, Kazanovka, Skvortsov (MW0061302). E, F, K— A. ledebourii (59 ) , MONGOLIA. Ubsunur: S of Altan-Els, 70 km NE of Baruun-Turuun, 5 July 1988, Kamelin et al. 867 (LE). G–I— A. kuvaevii , RUSSIA, Tuva: Ulug-Khem Raion, West Sayan Mts, vill. Bayan-Kol, 8 July 1974, Lomonosova & Iskakova 2709 (MHA, NS). J— A. pungens (192), RUSSIA . Tuva : West Sayan, Kartushibinskii ridge, Semiozersk Slaughterhouse, 13 June 1980 , Shaulo & Danilov 3867 (MHA). L— A. decipiens × A. ledebourii (238), RUSSIA . Tuva : Ulug-Khem Raion, West Sayan Mts, the Uyuk ridge, vill. Bayan-Kol, 6 August 1976 , Lomonosova & Ivanova 2408 (MHA). Scale Bar = 10 µm (A-F); = 300 µm (G), =100 µm (H); = 30 µm (I); =500 µm (J); =3 µm (K, L). FIGURE 7. Pollen of A. kuvaevii (A, B), A. pungens (C), A. decipiens (D, E), A. decipiens × A. ledebourii (F). A, B—tricolporate pollen grain of A. kuvaevii , RUSSIA. Tuva: Ulug-Khem Raion, West Sayan Mts, the Uyuk ridge, vill. Bayan-Kol, 8 July 1974, Lomonosova & Iskakova 2709 (MHA, NS). C—tricolporate pollen grain of A. pungens (255), RUSSIA. Khakassia: Askiz Raion, the river Askiz, Kazanovka, 23 June 2001, Skvortsov (MW0061302). D, E—loxocolporate, syncolporate pollen grains and pollen with ring-like form of ectoapertures of A. decipiens , KAZAKHSTAN. Aktiubinsk reg.: Temir Raion, 1950, Proskuriakova (MHA). F—tricolporate pollen grain of A. decipines × A. ledebourii (238), RUSSIA. Tuva: Bayan-Kol vill., Lomonosova & Ivanova 2408 (MHA). Scale Bar = 3 µm (A, B); = 10 µm (C, E), =30 µm (D); = 5 µm (F). FIGURE 8. Distributional map of Atraphaxis species in RUSSIA: Tuva, based on plastid haplotypes: 1— A. pungens . 2— A. ledebourii . 3— A . × uyukensis. 4— A. decipiens . 5— A. decipiens × A. pungens . 6— A. kuvaevii . 7— A. decipiens × A. ledebourii . FIGURE 9. Atraphaxis pungens (A–E) and A. × uyukensis (F): A–C—short generative shoots with small leaf blades; D–F—long vegetative shoots with large leaf blades. A—(184) RUSSIA. Tuva: Pii-Khem Raion, Kartushibinskii ridge, vill. Shivilig, 16 June 1987, Shaulo & Krasnikov 84 (NSK). B—RUSSIA. Tuva: the West Sayan, Kartushibinskii ridge, environs of Semiozersk Slaughterhouse, 13 June 1980, Shaulo & Danilov 3867 (NSK). C—(304) RUSSIA. Tuva: Pii-Khem Raion, vicinity of Shivilig vill., 923 m, 52°13.434’N, 93°44.148’E, 26 Jul 2015, Kostikov & Kostikova (MW). D—(253) RUSSIA. Khakassia: Shira Raion, lake shore Fyrkal, 05 July 2014, Kostikov & Kostikova (NSK). E—(115) MONGOLIA. Uver-Khangay: S of Tugrek, Bajangyin-Nuru (spurs of Dzegest-Ula), 27 July 1983, Gubanov 7471 (MW). F—(301) RUSSIA. Tuva: Kyzyl Raion, Kara-Khaak vill., 655 m, 27 July 2015, Kostikov & Kostikova (MW). FIGURE 10. Atraphaxis ledebourii (A–E) and A. pungens × A. ledebourii (F). A–D—short generative shoots with small leaf blades; B, E, F—long vegetive shoots with large leaf blades. A—(233) RUSSIA. Altai Rep.: Kosh-Agach Raion, the Chuya river, the mouth of the Chagan-Uzun river, 23 July 1995, Kamelin et al. (LE) B—(234) RUSSIA. Altai Rep.: Kosh-Agach Raion, the Kyzylshin river above Kokorya vill., 19 Aug 2000, Kamein et al. 2227 (ALTB). C—(296) RUSSIA. Tuva: Chaa-Khol’ Raion, the Sayano-Shushenskii reservoir, steppe slope, 12 July 2014, Kostikov & Kostikova (MW). D—(191) RUSSIA. Tuva: Chedy-Khol’ Raion, the Elegest river, 10 July 2014, Kostikov & Kostikova (MW, NSK). E—(298) RUSSIA. Tuva: Erzin d., 17 km N of vill. Erzin, river bank Tarlashkyn, 29 July 2015, Kostikov & Kostikova (MW, NSK). F—(243) RUSSIA. Tuva: Kyzyl Raion, Kaa-Khem vill., 30 July 2002, Nikitin et al. 98 (LE). FIGURE 11. A. × uyukensis Yurtseva, nom. provis . A, D—(241) RUSSIA. Tuva: Kyzyl Raion, spurs of the Uyuk ridge, the right bank of the Big-Khem (Big Yenisey) river above the mouth of the Seserlig river, opposite the vill. Kara-Khaak. nanophyton steppe, 600 m, 51°54’14’’N, 94°25’30”E, 1 June 2013, Doronkin. T 212 (NSK). B, E—(311) RUSSIA. Tuva: Pii-Khem Raion, the West Sayan, mouth of the Uyuk river, rocks, 52°07’N, 94°20’E, 31 May 1988, Shaulo & Shulo NS-11 / 560 (NSK). C, D—RUSSIA. Tuva: Pii-Khem Raion, the West Sayan, the Uyuk ridge, mouth of the Uyuk river, 9 August 1986, Shaulo & Zhdanova 312 (NSK). FIGURE 12. Atraphaxis pungens (M.Bieb.) Jaub. & Spach. : leaf blades of short (A–B) and long (C) shoots, flowers (D–F, H–I), styles (G), and fruits (J–L). A, D—(192) RUSSIA. Tuva: the West Sayan, Kartushibinskii ridge, Semiozersk Slaughterhouse, 13 June 1980, Shaulo & Danilov 3845 (MHA). B, E–G—the same, 3867 (NSK). C, H, K, L—(303) RUSSIA . Tuva : Ordzhonikidze Raion , Iyus vill., Mt Sunduk , 05 July 2014 , Kostikov & Kostikova ( MW ). I, J—(304) RUSSIA . Tuva , Pii-Khem Raion , Shivilig vill., 26 July 2015 , Kostikov & Kostikova ( MW ). Scale Bar = 1 µm (A, B, I); = 2 µm (C, H), = 0.2 µm (D, G, L); = 0.5 µm (E, F, J, K). FIGURE 13 . Atraphaxis ledebourii Yurtseva sp. nov . : leaf blades at short shoots (A) and long shoots (B), flowers (C, G), styles (D, E, H), and fruits (F, I). A—(233) RUSSIA. Altai Rep.: Kosh-Agach Raion, the Chuya river, 5 km above the mouth of the Chagan-Uzun river, 23 July 1995, Kamelin (LE). B–F—(234) RUSSIA. Altai Rep.: Kosh-Agach Raion, the Kyzylshin river above Kokorya vill., 19 August 2000, Kamelin et al. 2227 (ALTB). G, H—(300) RUSSIA. Altai Rep.: Onguday Raion, between the vill. Kupchegen and Inya, 8 July 2015, Kostikov & Kostikova (NSK). I—(296) RUSSIA. Tuva: Chaa-Khol’ Raion, Sayano-Shushenskii reservoir, steppe slope, 12 Jul 2014, Kostikov & Kostikova (MW). Scale Bar: = 1 µm (A, C, G, F); = 2 µm (B), = 0.2 µm (D, H); = 0.5 µm (E, I). Pollen of A. pungens and A. ledebourii is tricolporate, oblong-spheroidal, elliptical in equatorial view, roundedtrilobed in polar view, with distinct, deep, and long colpi (ectoapertures). Atraphaxis pungens from Askiz, Khakassia (255) has syncolporate pollen 29.3–32.4 × 20.5–24.5 µm, A. ledebourii (sub A. pungens ) from Kharalyg-Khem, Tuva (247) has the same pollen size (27.7 (25.8–31.8) × 26.3 (24.5–27.9) µm ( Yurtseva et al. 2014 ). The pollen surface of A. pungens has striato-perforate ornamentation with distinct, protruding, intensively branching and merging striae and grooves with 2–7 pits or perforations in a row ( Fig. 7C ). Atraphaxis pungens 304 from Kartushibinskii ridge has spherical pollen grains 30 µm in diameter with abnormal flat ridges and slit-like grooves (not shown). Thus, A. pungens and A. ledebourii do not show reliable morphological characteristics which let to clearly distinguish them. Due to high morphological similarity of A. pungens and A. ledebourii , their putative hybrid A. pungens × A. ledebourii (243) from Kaa-Khem village (Tuva, Kyzylskii Raion) does not differ from the parents. This hybrid has larger leaves at vegetative shoots, larger perianths and fruits ( Fig. 10F ) The reciprocal hybrid A. ledebourii × A. pungens 318 from Hangay, Mongolia , has black, shiny fruits common for A. pungens . FIGURE 14. Atraphaxis × uyukensis Yurtseva nom. provis .: leaf blade (A), perianth in fruiting (B), fruit (C). styles in young flower (D, E, F). A–C—(301) RUSSIA . Tuva : Kyzyl Raion , Kara-Khaak vill., 27 July 2015 , Kostikov & Kostikova ( MW ). D–G—(241) RUSSIA . Tuva : Kyzyl Raion , the Uyuk ridge, the right bank of the Big Yenisey opposite Kara-Khaak vill., 1 June 2013 , Doronkin T 212 ( NSK ). H, I—(311) RUSSIA . Tuva : Pii-Khem Raion, mouth of the Uyuk river, 31 May 1988 , Shaulo & Shaulo 11 / 560 (NSK) Scale Bar: = 2 µm (A); =1 µm (B, D–F, H); = 0.2 µm (C, G, I). Three specimens of A × uyukensis (241, 301, 311) from Kara-Khaak and the Uyuk ridge, Tuva , combine plastid haplotype U and ITS ribotypes P2 and P2del of A. pungens (303, 316, 317, 319) ( Figs 1 , 3 ). The sample (301) from Kara-Khaak, Tuva ( Fig. 9F ) resembles other specimens of A. pungens in the appearance, long vegetative shoots, broadly elliptical leaf blades, perianth, and black glossy fruits ( Fig. 14A–C ). The samples 241 and 311 ( Fig. 11 ) resemble the samples A. pungens (184, 192, 304) from Kartushibinskii ridge ( Fig. 9A–C ) in the appearance, oblongoblanceolate leaf blades of short generative and vegetative shoots, small thyrses, and yellowish perianths ( Fig. 14D–F ). Their stygmas are flat, which is a characteristic of A. pungens (compare Fig. 12D–F and Fig. 14G–I ).