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Pollen morphology and taxonomy of Atraphaxis (Polygoneae, Polygonaceae)

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Abstract

Pollen grains of 32 species of Atraphaxis L. including six species previously attributed to Polygonum L. s.str. were investigated with LM (acetolysed pollen) and SEM (dry pollen). Pollen grains of all studied species are spheroidal to oblong-spheroidal, mostly tricolporate, rarely 4-6-loxocolporate, ellipsoidal in equatorial view, rounded-trilobed or trilobed in polar view, with striate or striato-perforate, reticulato-foveolate, or reticulato-perforate sporoderm ornamentation. Exine sculpture is the most variable feature, differing in width, distinctness, and spacing of the striae, which are divided by grooves with small pits or perforations in rows. The most distinct variants of sporoderm ornamentation are connected by a full range of transitions. Striato-perforate ornamentation is common in species with the perianth typical for Atraphaxis, whereas reticulato-foveolate or reticulato-perforate sporoderm ornamentation was found in species with the perianth of Polygonum-type.

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Acknowledgments

The work was performed at the User Facilities Center of M.V.Lomonosov Moscow State University with the financial support from the Ministry of Education and Science of the Russian Federation. We thank the staff of the User Facilities Center for support and assistance. We are grateful to the curators of the Herbaria LE, MW, MHA, and TAD for allowing us to examine the specimens and take pollen samples. We are grateful to O. A. Volkova, N. I. Filina, A. G. Bogdanov, and S. V. Polevova for technical assistance; to G. Yu. Klinkova, N. A. Suprun, and V. G. Onipchenko who collected some specimens for this study. We should like to thank Dr. S. B. Yazvenko (LGL Limited, Canada) for reviewing the English, and two anonymous reviewers for their critical reading of the manuscript and helpful suggestions. The study was supported by the Russian Foundation for Basic Research (project No. 11-04-01300-a).

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  1. Department of Higher Plants, Faculty of Biology, Lomonosov Moscow State University , Leninskie Gory, 1-12, 119 991, Moscow, Russia

    O. V. Yurtseva, E. E. Severova & I. Yu. Bovina

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  1. O. V. Yurtseva
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Atraphaxis angustigolia Jaub. et Spach.—Armenia, inter Ordubad et Akilis. 3.06.1914. Woronov (MW); Armenia., Megri d., Agarak, slope of Zangezur Ridge. 1.06.1973. Shvedchikova (MW) [SEM]; Caucasus, Ordubad. 13.05.1934. Ter-Minosyan (LE) [LM].

Atraphaxis ariana (Grigorj.) T.M.Schust. & Reveal (= Polygonum arianum Grigorj.)—Turkmenskaya SSR, vill. Morgunovsky, 25.04.1988. Gorelova (LE) [SEM]; Turkmenistan, [Badghyz, Kushka d.,] Morgunovsky, 730 m a.s.l., 05.1964. n 800. Chukavina (LE) [LM].

Atraphaxis atraphaxiformis (Botsch.) T.M.Schust. & Reveal (= Polygonum atraphaxiforme Botsch.)—Tajikistan, Turkestan ridge, confl. of the Kshemysh and the Bel-Su. 03.07.1963. n 65. Abdusaljamova (LE) [SEM, LM].

Atraphaxis avenia Botsch.—South Tajikistan, Bag on the river Pjandzh, right bank of the river Shpilyau. 1.06.1960. n 779. Botschantzev and Egorova (LE) [SEM, LM].

Atraphaxis aucherii Jaub. et Spach—West Persia, Iran, Mazanderan, Bakhtiar Mts, Arabistan, Mulla-Ili, Gilang. 15.05.1904. n 341. Gadd (LE) [SEM]; Persia, n 3271. Aucher-Eloy (LE) [LM].

Atraphaxis badghysi Kult.—Turkmenistan, Maryiskaya reg. 35 km S of Shor-Tepe. 4.04.1952. Zhudova and Gruzdeva (MW); Turkmenistan, Badghys, S of saline Namaksaar. 25.04.1976. n 259. Botschantzev (LE) [SEM]; Turkmenia, in cliff Kzyl-Dzhar, NE of lake Er-Oylan-duz. 16.04.1964. n 281. Botschntsev (LE) [LM].

Atraphaxis billardierei Jaub. et Spach—Turkey, Prov. Nevsehir, Ürgüp. 1300 m. 24.05.1960. n 8469. Stainton and Henderson (LE) [SEM, LM].

Atraphaxis bracteata Losinsk.—Mongolia, E of Altay Goby, ridge Hurh-Ula, South Goby aimag. 20-30 km S of Nomgon. 30.07.1989. n 89. Gubanov and Grubov (MW) [SEM, LM].

Atraphaxis caucasica (Hoffm.) Pavlov—Georgia, Akhaltsikhe d., cl. Akhaltsikhe, 1080 m. a.s.l. 26.05.1973. Schvedchikova (MW) [SEM]; Georgia, Gory distr., Gorisdzhvary. 19.05.1949. Lorkipanidze (LE) [LM].

Atraphaxis canescens Bunge—Kazakhstan, Dzhambul reg., Mts Chu-Ili, top of Sunhar-Tyube (Han-Tau). 14.05.1951. n 167. Pavlov (MW) [SEM]; East Kazakhstan, N of Izbaksan, outflow of Cherny Irtysh, foothills of Mt. Chakylmaak. 10.05.1863. n 370. Potanin (LE) [LM].

Atraphaxis compacta Ledeb.—Kazakhstan, Syr-Darya reg. Mts Kara-Tau, Kos-Degeres. 11.05.1936. n 46. Chilikina (MW) [SEM]; Uzbekistan, Gyssar ridge, Mt. Baysun, Myn-Chukur, 2800 m a.s.l. 1935. Gordienko (MHA) [LM].

Atraphaxis daghestanica (Lovelius) Lovelius—Azerbajan, Apscheron d., Ambaky. Volkano Gaildag. 3.05.1974. Mailov (MHA) [SEM, LM]; Daghestan, Untsukul, 1970 m a.s.l. 7-8.07.1967. Bekova (MHA) [SEM, LM]; Daghestan, Gergebil distr., Avarskoye koysu, between the rivers Irganay and Untsukul. 10.06.1961. n 723, Guseinov and Tzvelev (LE) [LM].

Atraphaxis decipiens Jaub. et Spach—West Kazakhstan, Aktiubinsk reg. Temir distr. 1950. Proskuriakova (MHA) [SEM]; Kazakhstan, Karaganda reg., Central Betpak-Dala. 9.08.1955. Nikiforov (MHA) [LM].

Atraphaxis frutescens (L.) K.Koch—Mongolia, Mongol Altay, Goby Altay aimag, Ridge Huvchin-Nuru, 25 km E of Hayrhan. 9.08.1978. n 7772. Ogureeva (MW) [SEM]; Tuva ASSR, Kyzyl reg., 20 km S of Cherby. 13.07.1980. Kuminova and Kuprianov (MW) [LM].

Atraphaxis karataviensis Pavlov & Lipsch.—West Tien Shan, Ridge Kara-Tau, Dzhyren-Tura. 12.06.1931. Pavlov (MW) [SEM]; Uzbekistan, Ridge Nuratau, Gurdara, Sarymsak. 23.06.1971. n 392, Botschantzev and Kamelin (LE) [LM].

Atraphaxis kopetdagensis Kovalevsk.—Turkmenistan, Central Kopet-Dagh, Geok-Tepe d., Kara-Agach. 30.05.1972. Mescheryakov (LE) [SEM]; 05.1886. n 527. Germ. (LE) [LM].

Atraphaxis laetevirens (Ledeb.) Jaub. et Spach—Dzungar Alatau, West Ridges, Begash. 4.06.1959. Goloskokov (MW) [SEM]; Tadjikistan. 1988. Ryabkova (TAD) [LM].

Atraphaxis lanceolata (Bieb.) Meisn.—Altay, Semipalat. d. Ust-Kamenogorsk. 1912. n 145. Resnichenko (MW) [SEM]; Russia, Orenburg. reg., Buguruslan, Abdulinsky distr., Proletarsky. 06.1930. Kurnaev (MW) [LM].

Atraphaxis muschketowi Krasn.—Kazakhstan, Tien Shan, Zaily Alatau, Alma-Ata. 24.05.1998. n 98-24a. Majorov (MW) [SEM]; Moscow, Lomonosov Moscow State University, in culture. 22.05.1964. Trubacheva [LM].

Atraphaxis ovczinnikovii (Czukav.) O.V.Yurtseva (= Polygonum ovczinnikovii Czukav.)—South Tajikistan, Bag on the river Pyandzh. Right bank of the Shpilau. 1.06.1960. Nepli (LE) [SEM, LM].

Atraphaxis popovii (Borodina) O.V.Yurtseva (= Polygonum popovii Borodina)—NW China, Xinjiang, from Urumchi to Karashar. 22.07.1958. n 6223. Popov (LE) [SEM]

Atraphaxis pungens (Bieb.) Jaub. et Spach—Siberia, Khakasia, Askyz. d., the river Askyz, opposite Kazanovka, the road to Bolganov (Anchil-Chon). 23.06.2001. Skvortsov (MW) [SEM]; Tuva ASSR, Tes-Khem d., Ridge Tannu-Ola, 1200 m. a.s.l. Kharalyg-Khem. 3.06.1972. n 4821. Khanminchun (MW) [LM].

Atraphaxis pyrifolia Bunge—Uzbekistan, Tashkent reg., inter Gazalkent and Chimgan. 29.06.1976. Baranova and Kluykov (MW); Kyrgyzstan, Panfilov d. Cholok-Kaindy, 25.05.1986. n 430-7. Kuvaev (MW) [SEM]; Uzbekistan, Chatkal Mt. Reserv. Bash-Kyzyl-Say. 26.04.1975. Alferova and Udintseva (MHA) [LM].

Atraphaxis replicata Lam.—Daghestan, Buinaksk d., Kumtorkale. 29.09.63. Rotov (MW) [SEM]; Astrakhan reg., Narimanov distr., Kucherganovka. 23.06.1972. Safonov (MHA) [LM].

Atraphaxis rodinii Botsch.—Turkmenistan, Badghys, Akar-Cheschme. 11.05.1976. n 679. Botschantzev (LE) [SEM]; Turkmenia, Badghys, Pynkhan-Cheshme. 27.04.1978. n 2198. Gorelova (LE) [LM].

Atraphaxis seravschanica Pavlov—Uzbekistan, West Tien Shan, Chatkal Ridge, 40 km NE of Mt Angren. 16.05.1965. n 43. Boryaev and Gubanov (MW) [SEM]; Tajikistan, Turkestan Ridge. Ryabkova (TAD) [LM].

Atraphaxis spinosa L.—Russia, Volgograd reg., Kirov. d., Bolshaya Otrada. 07.05.2009. Klinkova and Suprun (MW) [SEM, LM]; Kazakhstan, Guriev (Atyrau) d. lake Inder. 4.05.2011. Onipchenko (MW) [SEM, LM]; Daghestan, Usuh-Chay. 9.07.1940. Nenevsky (MW) [SEM]; Armenia, Ararat. d. Kyarky. 1600 m a.s.l. 19.07.1972. Shvedchikova (MW) [LM]; Russia, Volgograd, Nizhnya Dobrinka. 27.07.1974. Artemenko (MW) [LM].

Atraphaxis teretifolia (Popov) Komarov ex Pavlov (= Atraphaxis jrtyschensis Chang Y.Yang & Y.L.Han)—Kazakhstan, Karaganda reg., Pribalkhashye, Sar-Tchagan. 26.05.1951. n 333. Pavlov (MW) [SEM, LM].

Atraphaxis toktogulica (Lazkov) T.M.Schust. & Reveal (= Polygonum toktogulicum Lazkov)—Kyrgyzstan, Toktogul d., Karajigach, Tor-Kolot. 5.07.1973. Aidarova et al. (LE) [SEM, LM]

Atraphaxis tortuosa Losinsk. (= Polygonum intramongolicum Fu & Zhao; = Polygonum tortuosum (Losinsk.) Lovelius)—Mongolia occ., the river Huang-He, Muni-Ula. ½.05.1872. Przhewalsky. (LE) [SEM, LM]

Atraphaxi s tournefortii Jaub. et Spach—Persia, Azerbaijan, 26 km W of Rezaiyeh, to Serow. 14.06.1971. n 4137. Lamond (LE) [SEM].

Atraphaxis virgata (Regel.) Krasn.—Mongolia, Altay Goby, foothills of Atas-Bogdo, 15 km N of Den-Ula. 3.08.1978. n 171. Ogureeva (MW) [SEM]; Tadjikistan, Turkestan Ridge. 06.1967. Soskov (TAD) [LM].

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Yurtseva, O.V., Severova, E.E. & Bovina, I.Y. Pollen morphology and taxonomy of Atraphaxis (Polygoneae, Polygonaceae). Plant Syst Evol 300, 749–766 (2014). https://doi.org/10.1007/s00606-013-0917-4

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