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

OLGA V. YURTSEVA; NATALIA V. VASILIEVA; VERA A. KOSTIKOVA; TAHIR H. SAMIGULLIN

doi:10.11646/phytotaxa.566.1.2

Issue: Vol. 566 No. 1: 22 September 2022

Type: Article

Published: 2022-09-22

DOI: 10.11646/phytotaxa.566.1.2

Page range: 13-63

Abstract views: 50

PDF downloaded: 1

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

OLGA V. YURTSEVA⁺⁻
NATALIA V. VASILIEVA⁺⁻
VERA A. KOSTIKOVA⁺⁻
TAHIR H. SAMIGULLIN⁺⁻

Department of Higher Plants, Faculty of Biology, M.V. Lomonosov Moscow State University, 1–12, Leninskie Gory, 119234, Moscow, Russia

N.V.Tsitsin’s Main Botanical Garden of the Russian Academy of Sciences, 4 Botanicheskaya Ulitsa, 127276, Moscow, Russia

Central Siberian Botanical Garden SB RAS, 101, Zolotodolinskaya Str., 630090, Novosibirsk, Russia

A.N.Belozersky Research Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 1–40, Leninskie Gory, 119992, Moscow, Russia

endemic hybridization ILS molecular phylogeny taxonomy pollen Siberia Eudicots

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Abstract

Maximum Likelihood (ML) and Bayesian analyses (BI) applied for combined plastid matrix (cpDNA trnL^(UAA) intron, trnL–trnF IGS, and rpl32–trnL^(UAG) IGS regions) of 124 accessions of 36 species of the genus Atraphaxis have shown new species from southern Siberia. The analysis of 112 accessions of 36 species of nrDNA ITS1–5.8S–ITS2 regions showed less clear topology. A new species A. kuvaevii endemic for Yenisey Valley in Tuva was revealed with molecular and morphological data. The plants attributed previously to A. pungens split into three cryptic species with different plastid regions. A. pungens grows in the south of Krasnoyarskii Krai, Khakassia, Tuva and Mongolia. A. ledebourii is distributed in Altai Mts, Tuva and Mongolia. It combines a functional ITS ribotype and a psedogenized ribotype, what agrees with high chromosome numbers. The third pseudogenized ribotype shared with A. ariana was found in a single sample from SE Tuva. A. ×uyukensis is a species of hybrid origin from the Uyuk ridge with the ITS ribotype of A. pungens, which probably captured cpDNA of unknown species related to A. selengensis and A. davurica by maternal lineage. Despite the differences in plastid regions, these cryptic species have no clear morphoplogical distinctions. The hybrid A. pungens × A. ledebourii from Tuva and the hybrid A. ledebourii × A. pungens from Hangay Mts in Mongolia do not differ from the parents. The eastern race of A. frutescens s.l. sub A. decipiens from Tuva and Altai Republics differs from the western race of A. frutescens with a single nucleotide substitution in cpDNA trnL–trnF IGS, a distinct ITS ribotype, a short perianth tube equal to outer tepals, herbaceous tubular ochreolas and linear-lanceolate leaf blades. The hybrids A. pungens × A. decipiens and A. decipiens × A. pungens were detected in three locations in Tuva. Cryptic hybrids A. ledebourii × A. decipiens, A. ledebourii × A. frutescens, and A. decipiens × A. ledebourii were detected in Altai and Tuva Republics. Two hybrids A. decipiens × A. ledebourii have shown the capture of cpDNA of A. decipiens. Atraphaxis laetevirens is limited by the territory of Kazakhstan, Altai Republic and Dzungaria and does not grow in Tuva and Khakassia.

References

Anisimova, M., Gil, M., Dufayard, J-F., Dessimoz, C. & Gascuel, O. (2011) Survey of branch support methods demonstrates accuracy, power, and robustness of fast likelihood-based approximation schemes. Systematic Biology 60 (5): 685–699. https://doi.org/10.1093/sysbio/syr041
Bajtenov, M.B. & Pavlov, N.V. (1960) Polygonum L. In: Pavlov, N.V. (Ed.) Flora of Kazakhstan, vol. 3. Akademia Nauk Kazahkskoj SSR, Alma-Ata, pp. 147–176. [in Russian]
Bao, B. & Grabovskaya-Borodina, A.E. (2003) Atraphaxis L. In: Wu, Z.Y., Raven, P.H. & Hong, D.Y. (Eds.) Flora of China vol. 5. Science Press, Beijing; Missouri Botanical Garden Press, St. Louis, pp. 328–332. [http://www.efloras.org/florataxon.aspx?flora_ id=2&taxon_id=103103]
Borodina, A.E. (1989) Polygonaceae. In: Grubov, V.I. (Ed.) Rastieniya Centralnoi Azii. Po materialam Botanicheskogo Instituta im.V.L. Komarova (Plantae Asiae Centralis, secus materies Instituti botanici nomine V.L. Komarovii). 9 (Salicaceae—Polygonaceae). Nauka, Leningrad, pp. 77–129. [in Russian]
Brandbyge, J. (1993) Polygonaceae. In: Kubitzki, K., Rohwer, J.G. & Bittrich, V. (Eds.) The families and genera of vascular plants, vol. 2. Springer, Berlin, pp. 531–544. http://dx.doi.org/10.1007/978-3-662-02899-5_63
Campbell, C.S., Wojciechowski, M.F., Baldwin, B.G., Alice, L.A. & Donoghue, M.J. (1997) Persistent nuclear ribosomal DNA sequence polymorphism in the Amelanchier agamic complex (Rosaceae). Molecular biology and evolution 14 (1): 81–90.
Cullen, J. (1967) Atraphaxis L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean islands, vol. 2. Edinbourgh University Press, Edinbourgh, pp. 266–267.
Doyle, J.J. (1992) Gene trees and species trees: molecular systematics as one-character taxonomy. Systematic Botany 17 (1): 144–163. https://doi.org/10.2307/2419070 
Drábková, L.Z., Kirschner, J., Štěpánek, J., Záveský, L. & Vlček, Č. (2009) Analysis of nrDNA polymorphism in closely related diploid sexual, tetraploid sexual and polyploid agamospermous species. Plant systematics and evolution 278 (1): 67–85. https://doi.org/10.1007/s00606-008-0134-8
Edman, G. (1931) Apomeiosis and Apomixis bei Atraphaxis frutescens C.Koch. Acta horti bergiani (Almqvist & Wiksell) 11 (2): 11–66.
Erdtman, G. (1971) Pollen morphology and plant taxonomy: Angiosperms. An Introduction to Palynology, vol. 1. Hafner Publishing Co, New York, 539 pp.
Fehrer, J., Gemeinholzer, B., Chrtek, Jr. J. & Bräutigam, S. (2007) Incongruent plastid and nuclear DNA phylogenies reveal ancient intergeneric hybridization in Pilosella hawkweeds (Hieracium, Cichorieae, Asteraceae). Molecular phylogenetics and evolution 42 (2): 347–361. https://doi.org/10.1016/j.ympev.2006.07.004
Gmelin, I.G. (1768) Flora Sibirica, Tome 3. Academiae Scientiarum, Petropoli, pp. 1–276. + 67 tables.
Grant, V. (1981) Plant speciation. 2nd ed. Columbia university press, New York, 563 pp.
Grubov, V.I. (1982) Opredelitel sosudistyh rasteniy Mongolii (Key to the vascular plants of Mongolia, with atlas). Nauka, Leningrad, 442 pp. [in Russian]
Gubanov, I.A. (1996) Conspectus of Flora of Outer Mongolia: Vascular Plants. Valang, Moscow, 136 pp. [in Russian]
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence editor alignment and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
Hesse, M., Halbritter, H., Zetter, R., Weber, M., Buchner, R., Frosch-Radivo, A. & Ulrich, S. (2009) Pollen terminology: An illustrated handbook. Springer Wien, New York, 264 pp.
Holder, M.T., Anderson, J.A. & Holloway, A.K. (2001) Difficulties in detecting hybridization. Systematic Biology 50 (6): 978–982.
Hong, S.-P., Ronse De Craene, L.-P. & Smets, E.F. (1998) Systematic significance of tepal surface morphology in tribes Persicarieae and Polygoneae (Polygonaceae). Botanical Journal of the Linnean Society 127 (2): 91–116. https://doi.org/10.1111/j.1095-8339.1998.tb02091.x
Ivanova, M.M. (1979) Atraphaxis L. In: Malyshev, L.I. & Peshkova, G.A. (Eds.) Flora Siberiae Centralis, vol. 1. Nauka, Sibir. otdelenie, Novosibirsk, p. 281. [In Russian]
Jaubert, H.-F.C. & Spach, E. (1844–1846) Illustrationes Plantarum Orientalium, vol. 2. Roret, Parisiis, iii+123 pp.; tabs. 101–200.
Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30 (4): 772–780. http://dx.doi.org/10.1093/molbev/mst010
Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30 (14): 3059–3066. http://dx.doi.org/10.1093/nar/gkf436 
Kashina, L.I. (1979) Atraphaxis L. In: Krasnoborov, I.M. & Kashina, L.I. (Eds.) Opredelitel’ rasteniy yuga Krasnoyarskogo Kraia (Key to plants of the south of the Krasnoyarsk Territory). Nauka, Novosibirsk, pp. 138. [in Russian] 
Kashina, L.I. (1992) Atraphaxis L. In: Krasnoborov, I.M. & Malyschev, L.I. (Eds.) Flora Sibiriae, vol. 5. Salicaceae—Amaranthaceae. Nauka, Div. Sibirica, Novosibirsk, pp. 108–109. [in Russian] 
Kongar, E.T. & Tupitsyna, N.N. (2007) Polygonaceae. In: Shaulo, D.N (Ed.) Opredelitel’ rasteniy Respubliki Tyvy (Key to plants of the Republic of Tuva), 2nd ed. rev. and add., Publishing house of the SB RAS, Novosibirsk, pp. 154–168. [in Russian] 
Koch, K. (1872) Dendrologie. Bäume, Sträucher und Halbsträucher, welche in Mittel- und Nord-Europa im Freien kultivirt werden. Kritisch beleuchtet von Karl Koch. Bd. 2 (1), F.Enke, Erlangen, 424 pp. [http://nrs.harvard.edu/urn-3:HUL.FIG:005788153] 
Koropachinsky, I.Yu. & Vstovskaya, T.N. (2002) Drevesnye rasteniia Aziatskoi Rossii (Woody Plants of Asian part of Russia). Publishing house of the SB RAS, Geo, Novosibirsk, 706 pp.
Kostikov, D.K. & Banaev, E.Y. (2016) Izmenchivost’ morfologicheskikh priznakov Atraphaxis frutescens (L.) C. Koch. (Variability of morphological features of Atraphaxis frutescens (L.) C. Koch. Izvestiya Samarskogo nauchnogo centra Rossijskoj akademii nauk 18 (2): 109–113. [in Russian]
Kovarik, A., Dadejova, M., Lim, Y.K., Chase, M.W., Clarkson, J.J., Knapp, S. & Leitch, A.R. (2008) Evolution of rDNA in Nicotiana allopolyploids: a potential link between rDNA homogenization and epigenetics. Annals of Botany 101 (6): 815–823. https://doi.org/10.1093/aob/mcn019
Krak, K., Caklova, P., Chrtek, J. & Fehrer, J. (2013) Reconstruction of phylogenetic relationship in a highly reticulate group with deep coalescence and recent speciation. Heredity 110: 138–151. http://dx.doi.org/10.1038/hdy.2012.100 
Krasnov, A.N. (1888) Opyt istorii razvitiya flory yuzhnoi chasti Vostochnogo Tyan’Shanya (An Essay of the history of development of the Flora in the Southern Part of Eastern Tien Shan). Zapiski Russkogo Geograficheskogo obschestva po obschey geografii 19: 1–413. [in Russian]
Krylov, P.N. (1909) Flora Altaya i Tomskoy gubernii: Rukovodstvo k opredeleniyu rasteniy Zap. Sibiri / (Soch.) P. Krylova (Flora of Altai and Tomsk province: Guide to the identification of plants Zap. Siberia / (Works) P. Krylova), T. 5. Salsolaceae—Betulaceae. Tipo-litografiya Sibirskogo T-va Pechatnogo Dela (Typo-lithograph of the Siberian Printing House), Tomsk, pp. 1089–1252. [in Russian] 
Krylov, P.N. (1930) Flora Siberiae Occidentalis, Ed. sec. et comp. Flora Altaica et provinciae Tomskensis, vol. 4. Salicaceae—Amaranthaceae. Editio sectionis Tomskensis Societatis botanicae Rossicae, Tomsk, 979 pp. [In Russian]
Kulikov, P.V. (2010) Opredelitel sosudistykh rasteniy Chelyabinskoy oblasti (Key for vascular plant identification of the Chelyabinsk Region). Ural Branch of the Russian Academy of Sciences, Yekaterinburg, 969 pp. [In Russian] 
Ledebour, C.F., Meyer, C.A. & Bunge, A. von (1830) Flora Altaica, vol. 2. G. Reimer, Berlin, pp. 464.
Ledebour, C.F. (1834) Icones plantarum novarum vel imperfecte cognitarum floram Rossicam, imprimis Altaicam, illustrantes, Cent. 5, facs. 1. Riga, London, Paris, Strasbourg & Brussels, 28 pp. 
Ledebour, C.F. (1849) Flora Rossica, vol. 3 (2). E. Schwiezerbart, Stuttgart, 373 pp. 
Linnaeus, C. (1753) Species Plantarum, vol. 1. Impensis Laurentii Salvii, Holmiæ [Stockholm], 560 pp.
Lovelius, O.L. (1978) Compositio specierum, Distributio geographica et cohaerentia oecologica generis Atraphaxis L. (Polygonaceae). In: Klokov, M.V. (Ed.) Novosti sistematiki vysshikh i nizshikh rasteniy (Novitates systematicae plantarum vascularium et non vascularium 1977), Naukova dumka, Kiev, pp. 85–108.
Lovelius, O.L. (1979) Synopsis generis Atraphaxis L. (Polygonaceae)[A systematic review of the genus Atraphaxis L.] Novitates systematicae plantarum vascularium, vol. 15, 1978. Leningrad, Nauka, pp. 114–128. [in Russian]
Marschall von Bieberstein, L.B.Fr. (1819) Flora Taurico-Caucasica, vol. 3. Typis Academicis, Charkov, 654 pp.
Milyutina, I.A., Goryunov, D.V., Ignatov, M.S., Ignatova, E.A. & Troitsky, A.V. (2010) The phylogeny of Schistidium (Bryophyta, Grimmiaceae) based on the primary and secondary structure of nuclear rDNA internal transcribed spacers. Molecular Biology 44 (6): 883–897. https://doi.org/10.1134/S0026893310060051
Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., Von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular biology and evolution 37 (5): 1530–1534. https://doi.org/10.1093/molbev/msaa015
Müller, K. (2005) SeqState—primer design and sequence statistics for phylogenetic DNA data sets. Applied Bioinformatics 4: 65–69.
Nelson, G. & Platnick, N. (1981) Systematics and biogeography: Cladistics and vicariance. Columbia Univ. Press, New York, xi+567 pp.
Nikiforova, O.D. (2012) Atraphaxis L. In: Baikov, K.S. (Ed.) Conspectus Florae Rossiae asiaticae: Plantae vasculares. The Siberian branch of the Russian Acad. Sci., Novosibirsk, pp. 115 [in Russian] 
Osipov, K.I. (2001) Polygonaceae. In: Anenkhonov, O.A. (Ed.) Opredelitel rastenyi Buryatii (Key to Buryatia plants). The Siberian branch of the Russian Acad. Sci. Institute of Experimental and General Biology, Ulan-Ude, pp. 242–254. [in Russian]
Pallas, P.S. (1776) Reise durch verschiedene Provinzen des Russischen Reichs, Theil 3, Vom Jahr 1772 und 1773. Kayserliche Akad. der Wissenschaften, St. Petersburg, 454 pp.
Pavlov, N.V. (1936) Atraphaxis L. In: Komarov, V.L. (Ed.) Flora URSS, vol. 5. USSR Academy of Sciences, Moscow & Leningrad, pp. 501–527. [in Russian]
Peshkova, G.A. (1972) Stepnaya flora Baikalskoy Sibiri (Steppe flora of the Baikal Siberia). Nauka, Moscow, 207 pp. [in Russian] 
Peshkova, G.A. (2005) Atraphaxis L. In: Baikov, K.S. (ed.) Conspectus Florae Siberiae: Plantae vasculares. Nauka, Novosibirsk, pp. 63. [in Russian]
Punt, W., Hoen, P.P., Nilsson, S. & Thomas, L. (2007) Glossary of pollen and spore terminology. The Review of Palaeobotany and Palynology 143: 1–81.
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v.1.6. Available from: http://tree.bio.ed.ac.uk/software/tracer/ (accessed 7 August 2022).
Rechinger, K.H. & Schiman-Czeika, M. (1968) Polygonaceae. In: Rechinger, K.H. (Ed.) Flora iranica, vol. 56. Academishe Druck-u.-Verlagsanstalt, Graz, Austria, pp. 1–88. 
Regel, E.A. von (1880) Descriptiones Plantarum novarum et minus cognitarum. Fasc. VII. Acta Horti Petropolitani. T. 6, Fasc. II. (1879). pp. 287–538. 
Rieseberg, L.H. & Soltis, D.E. (1991) Phylogenetic consequences of cytoplasmic gene flow in plants. Evolutionary Trends in Plants 5 (1): 65–84. 
Rieseberg, L.H., Whitton, J. & Linder, C.R. (1996) Molecular marker incongruence in plant hybrid zones and phylogenetic trees. Acta Botanica Neerlandica 45 (3): 243–262.
Rodionov, A.V., Amosova, A.V., Krainova, L.M., Machs, E.M., Mikhailova, Y.V., Gnutikov, A.A., Muravenko, O.V. & Loskutov, I.G. (2020) Phenomenon of Multiple Mutations in the 35S rRNA Genes of the C Subgenome of Polyploid Avena L. Russian Journal of Genetics 56 (6): 674–683. https://doi.org/10.1134/S1022795420060095
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
Ronse De Craene, L.-P. & Akeroyd, J.R. (1988) Generic limits in Polygonum and related genera (Polygonaceae) on the basis of floral characters. Botanical Journal of the Linnean Society 98 (4): 321–371. https://doi.org/10.1111/j.1095-8339.1988.tb01706.x
Ronse De Craene, L.-P., Hong, S.-P. & Smets, E. (2000) Systematic significance of fruit morphology and anatomy in tribes Persicarieae and Polygoneae (Polygonaceae). Botanical Journal of the Linnean Society 134 (1–2): 301–337. https://doi.org/10.1111/j.1095-8339.2000.tb02356.x
Ryabinina, Z.N. & Knyazev, M.S. (2009) Opredelitel sosudistykh rasteniy Orenburgskoy oblasti. (Key for vascular plant identification of the Orenburg Region). Tovarishchestvo nauchnykh izdaniy KMK, Moscow, 758 pp. [In Russian] 
Sambrook, J., Fritsch, E.F. & Maniatis, T. (1989) Molecular cloning: a laboratory manual Ed. 2, vols. 1–3. Cold spring harbor laboratory press. New York, pp.xxxviii + 1546 pp.
Schaal, B.A., Hayworth, D.A., Olsen, K.M., Rauscher, J.T. & Smith, W.A. (1998) Phylogeographic studies in plants: problems and prospects. Molecular Ecology 7 (4): 465–474. https://doi.org/10.1046/j.1365-294x.1998.00318.x
Shaw, J., Lickey, E.B., Schilling, E.E. & Small, R.L. (2007) Comparison of whole chloroplast genome sequences to choose noncoding regions for phylogenetic studies in angiosperms: the tortoise and the hare III. American Journal of Botany 94 (2): 275–288. https://doi.org/10.3732/ajb.94.3.275
Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: https://www.simplemappr.net/ (accessed 7 August 2022).
Simmons, M.P. & Ochoterena, H. (2000) Gaps as Characters in Sequence-Based Phylogenetic Analyses. Systematic Biology 49 (2): 369–381. https://doi.org/10.1093/sysbio/49.2.369
Sitnikov, A.P. (1986) Pseudogamy in Atraphaxis frutescens (Polygonaceae) in Tatar SSR. Gametogenesis, fertilization and embryogenesis of seed plants, ferns and mosses. In: Chebotar, A.A. (Ed.) IX Vsesoyuznoye Soveshanije po embriologii rastenyi. Shtiintsa, Kishinev, pp. 167–168. [in Russian]
Sitnikov, A.P. (1991) Variability of reproductive structures in the genus Polygonum L. and other members of Polygonaceae Juss. Dissertation cand. biol. nauk, Kasan, 179 pp. [in Russian]
Soltis, D.E. & Kuzoff, R.K. (1995) Discordance between nuclear and chloroplast phylogenies in the Heuchera group (Saxifragaceae). Evolution 49 (4): 727–742. https://doi.org/10.1111/j.1558-5646.1995.tb02309.x
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991) Universal primers for amplification of three non-coding regions of chloroplast DNA. Plant molecular biology 17 (5): 1105–1109. https://doi.org/10.1007/BF00037152
Thiers, B. (2022 [continuously updated]) Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s virtual herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 7 August 2022)
Tian, X.M., Liu, R.R., Tian, B. & Liu, J.Q. (2009) Karyological studies of Parapteropyrum and Atraphaxis (Polygonaceae). Caryologia 62 (4): 261–266. 
Turczaninow, N.S. (1856) Flora baicalensi-dahurica: seu, Descriptio plantarum in regionibus cis-et transbaicalensibus atque in Dahuria sponte nascentium, vol. 2 (1). Typis Universitatis Caesareae, Mosquae, 437 pp. 
Tzvelev, N.N. (1996) Flora of East Europe, vol. 9. Mir i Semja-95, SPb, 451 pp. [in Russian]
Volkova, P.A., Herden, T. & Friesen, N. (2017) Genetic variation in Goniolimon speciosum (Plumbaginaceae) reveals a complex history of steppe vegetation. Botanical Journal of the Linnean Society 184 (1): 113–121. https://doi.org/10.1093/botlinnean/box011
Voronkova, M.S., Banaev, E.V., Tomoshevich, M.A. & Kostikova, V.A. (2019) IAPT chromosome data 30/12. In: Marhold, K. et al. (ed.) IAPT chromosome data 30. Taxon 68 (5): 1130, E29.
Webb, D.A. (1964) Atraphaxis L. In: Tutin, T.G., Heywood, V.H., Burges, N.A., Valentine, D.H., Walters, S.M. & Webb, D.A. (Eds.) Flora Europaea, vol. 1. Cambridge University Press, Cambridge, pp. 89.
Wen, J. & Zimmer, E.A. (1996) Phylogeny and biogeography of Panax L.(the ginseng genus, Araliaceae): inferences from ITS sequences of nuclear ribosomal DNA. Molecular phylogenetics and evolution 6 (2): 167–177. https://doi.org/10.1006/mpev.1996.0069
Wendel, J.F. & Doyle, J.J. (1998) Phylogenetic incongruence: window into genome history and molecular evolution. In: Soltis, D.E., Soltis, P.S. & Doyle, J.J. (Eds.) Molecular systematics of plants II: DNA sequencing. Springer, Boston, MA, pp. 265–296.
White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR Protocols: a Guide to Methods and Applications 18 (1): 315–322.
Whitfield, J.B. & Lockhart, P.J. (2007) Deciphering ancient rapid radiations. Trends in ecology & evolution 22 (5): 258–265.
Yurtseva, O. & Mavrodiev, E.V. (2019) Caelestium genus novum (Polygonaceae, Polygoneae): evidence bаsed on the results of molecular phylogenetic analyses of tribe Polygoneae, established with consideration of the secondary structure of the ITS nrDNA regions Novitates systematicae plantarum vascularium 50: 47–79. https://doi.org/10.31111/novitates/2019.50.47
Yurtseva, O.V., Troitsky, A.V., Bobrova, V.K. & Voylokova, V.N. (2010) On taxonomical revision of Polygonum L. s.s. (Polygonaceae): phylogenetic and morphological data. Botanicheskii Zhurnal 95 (2): 226–247. [in Russian]
Yurtseva, O.V., Levina, M.S., Severova, E.E. & Troitsky, A.V. (2012) Morphology and taxonomy of Polygonum cognatum Meisn., P. alpestre C.A.Mey. and allied taxa from Central Asia and the Caucasus (Polygonaceae). Wulfenia 19: 141–180.
Yurtseva, O.V., Kuznetsova, O.I., Mavrodieva, M.E. & Mavrodiev, E.V. (2016a) What is Atraphaxis L. (Polygonaceae, Polygoneae): cryptic taxa and resolved taxonomic complexity instead of the formal lumping and the lack of morphological synapomorphies. PeerJ 4: e1977: 1–50. http://doi.org/10.7717/peerj.1977
Yurtseva, O.V., Kuznetsova, O.I. & Mavrodiev, E.V. (2016b) A broadly sampled 3-loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: I. Atraphaxis kamelinii spec. nov. from Mongolia. Phytotaxa 268 (1): 1–24. https://doi.org/10.11646/phytotaxa.268.1.1
Yurtseva, O.V., Severova, E.E. & Bovina, I.Yu. (2014) Pollen morphology and taxonomy of Atraphaxis (Polygoneae, Polygonaceae). Plant Systematics and Evolution 300 (4): 749–766. http://doi.org/10.1007/s00606-013-0917-4
Yurtseva, O.V., Severova, E.E. & Mavrodiev, E.V. (2017) Persepolium (Polygoneae): A new genus in Polygonaceae based on conventional Maximum Parsimony and Three-taxon statement analyses of a comprehensive morphological dataset. Phytotaxa 314 (2): 151–194. https://doi.org/10.11646/phytotaxa.314.2.1
Yurtseva, O.V., Badmaeva, N.K. & Mavrodiev, E.V. (2021) A broadly sampled 3-loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: II. Atraphaxis selengensis spec. nov., and A. davurica Jaub. & Spach from Russian Transbaikalia. Phytotaxa 484 (1): 44–74. https://doi.org/10.11646/phytotaxa.484.1.2 
Zuker, M. (2003) Mfold web server for nucleic acid folding and hybridization prediction. Nucleic Acids Research 31 (13): 3406–3415. https://doi.org/10.1093/nar/gkg595