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Turczaninowia 26, 3: 194-198 (2023) DOI: 10.14258/turczaninowia.26.3.18 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
УДК 582.998.2:581.95(571+571.52) Saussurea arctecapitulata Lipsch. - new species for the flora of Russia
E. A. Pyak1 2, A. V. Rudyev1 3, D. V. Yusupovsky1 4, A. I. Pyak1 5*
1 Tomsk State University, Lenina Pr., 36, Tomsk, 634050, Russian Federation 2E-mail: epyak.tom@gmail.com; ORCID iD: https://orcid.org/0000-0001-8694-7278 3E-mail: tyomarydyev@gmail.com; ORCID iD: https://orcid.org/0000-0001-6022-1321 4E-mail: unitycoredown@gmail.com; ORCID iD: https://orcid.org/0000-0002-2447-7001 5E-mail: a_pyak@rambler.ru; ORCID iD: https://orcid.org/0000-0002-6055-7973 * Corresponding author
Keywords: biodiversity, floristic findings, Republic of Tuva, Siberia, vascular plants.
Summary. Saussurea arctecapitulata, a herbaceous perennial species, belonging to the type section of the genus Saussurea, previously known only from a few localities in the northern Mongolia, has been found in the Naryn River Valley during field trips to the territory of Republic of Tuva. This article presents an updated description accompanied by photos of the species in nature, updated data on the distribution of the species, as well as phylogenetic relationships with morphologically similar species of the section based on the variability of the site ITS1-5.8S-ITS2.
Saussurea arctecapitulata Lipsch. - новый вид для флоры России
Е. А. Пяк, А. В. Рудьев, Д. В. Юсуповский, А. И. Пяк
Томский государственный университет, пр. Ленина, д. 36, г. Томск, 634050, Россия
Ключевые слова: биоразнообразие, Республика Тыва, Сибирь, сосудистые растения, флористические находки.
Аннотация. Saussurea arctecapitulata - травянистый многолетник, относящийся к типовой секции рода Saussurea, ранее известный лишь из нескольких местонахождений на территории Северной Монголии. В ходе полевых экспедиций по территории Республики Тыва нами были обнаружены популяции данного вида в долине реки Нарын. В статье представлены дополненное описание, сопровождаемое фотографиями вида в природе, обновленные данные о распространении вида, а также результаты филогенетического анализа с включением в анализ морфологически близких видов секции на основе изменчивости участка ITS1-5.8S-ITS2.
Introduction
The genus Saussurea DC. comprises about 450-500 species of perennial, biennial herbs and subshrubs widely distributed in temperate and subarctic regions of Asia (Lipschitz, 1979; Smirnov et al., 2018). It is one of the largest genera in the tribe Cardueae Cass. with the center of species richness and greatest morphological diversity in the Sino-Hi-
malaya and adjacent regions (Xu et al., 2019), however, the number of species may be still significantly underestimated due to lack of floristic investigations in different parts of the genus distribution range, especially in remote mountain areas. In June 2019, during a floristic investigation carried out in the Sayan Mountains, a population of Saussurea arctecapitulata Lipsch. was found in the Sangilen Plateau, near the mouth of the Bayan-Gol River. This species was
Поступило в редакцию 22.08.2023 Принято к публикации 05.11.2023
Submitted 22.08.2023 Accepted 05.11.2023
described by S. J. Lipschitz in 1972 based on the herbarium materials collected by V. L. Komarov from northern Mongolia in 1902 (Lipschitz, 1972), and it was not previously reported for the flora of Russia. In August 2023, expeditionary research was carried out in this area, and several more populations of this species, apparently common in the valley of the Naryn River in the Sangilen Plateau, were found.
Material and Methods
Samples for phylogenetic analyses and herbarium specimens were collected by the authors during field trips to Republic of Tuva, Republic of Altai (Russia), Khovd Aimag (Mongolia), Friuli-Venezia Giulia (Italy) in 2016-2023. In total, 8 taxa (S. arcte-capitulata, 6 morphologically related species from the type section of the genus, Jurinea multiflora (L.) B. Fedtsch. as an outgroups) were included in the study. All newly generated sequences were deposited in GenBank, for S. pseudoalpina N. D. Simpson, S. parviflora (Poir.) DC., S. neoserrata Nakai and J. multiflora we refer to GenBank data (Table 1). The studied specimens collected by the authors were deposited in the Herbarium of P. N. Krylov of Tomsk State University (TK, Tomsk, Russia). DNA extraction and sequencing. - Genomic DNA isolation from plant tissues was performed following a modified 2xCTAB protocol (Doyle J. J., Doyle J. L., 1990). To quantify DNA extracted from silica-gel dried samples, NanoDrop2000 spectrophotometer (Thermo Fisher Scientific, USA) was used. ITS1-5.8S-ITS2
fragments were amplified by polymerase chain reaction using the following primer combinations: ITS-p5(ccttatcayttagaggaaggag) / ITS-u4 (rgtttcttttcctc-cgctta) (Cheng et al., 2016), according to program, starting with 94 °C for 4 minutes, followed by 35 cycles of 30 seconds at 94 °C, 30 seconds at 53 °C and a final extension of 45 seconds at 72 °C with a final step of 10 minutes at 72 °C. Automated DNA sequencing was carried out directly from the purified with the PCR Purification Kit (Roche, Germany) PCR products using BigDye Terminator v.2 reagents in the ABI310 sequencer (PE-Applied Biosystems, USA). Sequence alignment and phylogenetic analyses. -DNA sequences were aligned in the MAFFT vs. 7 (Katoh, Standley, 2013), after which the alignment results were checked manually and edited using BioEdit vs. 7.0.1 (Hall, 1999). The best-fitting models of nucleotide substitution selected in MrMo-deltest v.2.4 using the Akaike Information Criterion (AIC) was GTR+I. Phylogenetic analyses was performed with MrBayes 3.1.2 (Ronquist, Huelsenbeck, 2003) using the Bayesian method. The analysis was performed using four incrementally heated Markov chains, simultaneously started from random trees and run for one million cycles, recording one tree every 1000 generations until the average standard deviation of split frequencies < 0.01 was achieved. The first 25 % of the trees were discarded as burn-in, after which the remaining trees were used to calculate a majority-rule phylogenetic tree edited using the FigTree 1.4.4 program (Rambaut, 2018).
Table 1
GenBank accession numbers and information about voucher specimens for the samples used in the phylogenetic analysis. Newly generated sequences highlighted in bold
№ Species Voucher specimens ITS1-5.8S-ITS2
1 Saussurea arctecapitulata TK-005031 OR565243
2 S. subacaulis AP2016002 OR565245
3 S. klementzii AP2019013 OR565244
4 S. alpina EP2019003 OR565246
5 S. pseudoalpina WYJ201607020 OR119937
6 S. pseudoalpina WYJ201308135 OQ539813
7 S. pseudoalpina WYJ201308201 0Q539809
8 S. parviflora LJQ-2008-GN-092 0Q539754
9 S. parviflora Ikeda200714073 0Q539755
10 S. neoserrata unpublished, no data AB254668
11 Jurinea multiflora WYJ201308102 MH003704
Results and discussion
Saussurea arctecapitulata Lipsch.: "Russia. Republic of Tuva, Erzinsky district, Sangilen Plateau,
Naryn River Valley, mouth of the Bayan-Gol River, 1577 m. 50°09'17.40"N, 95°58'28.1"E. 06 IX 2019. A. I. Pyak, E. A Pyak, S. Karachurina / Россия. Республика Тыва, Эрзинский район. Нагорье Сан-
Pyak E. A. et al.
Saussurea arctecapitulata Lipsch. - new species for the flora of Russia
гилен, долина реки Нарын, устье реки Баян-Гол, 1577 m. 50°09'17.40"N, 95°58'28.15"E. 06 IX 2019. А. И. Пяк, Е. А. Пяк, С. Карачурина" (TK [TK-004155]); "Russia. Republic of Tuva, Erzinsky district. Sangilen Plateau, Naryn River Valley, 1840 m. 50°12'15.7"N, 96°12'59.7"E. 03 VIII 2023. E. A Pyak, A. V. Rudyev, A. I. Pyak / Россия. Республика Тыва, Эрзинский район. Нагорье Сангилен, долина р. Нарын, пойма, сырые луговины и заросли кустарников, 1840 m. 50°12'15.7"N, 96°12'59.7"E. 03 VIII 2023. Е. А. Пяк, А. В. Рудьев, А. И. Пяк" (TK [TK-005031]).
Phylogenetic analyses. - Phylogenetic tree constructed based on the results of the ITS1-5.8S-ITS2 site variability is shown in the Fig. 1. The ingroup belonging to the species of the section Saussurea was retrieved as the monophyletic group (1.00 PP) and including two main clades. S. parviflora and S. ne-oserrata which constitute the first clade are sisters to the rest of the ingroup which includes S. pseu-doalpina related by sister relationship to S. alpina (L.) DC. with S. subacaulis (Ledeb.) Serg. (grouped together with 0.97 PP), S. klementzii Lipsch. and S. arctecapitula. The relationships between these taxa are unresolved due to polytomies.
Morphological description. - Herbs perennial, (5-)10-40 cm tall (Fig. 2). Root collar covered with remains of petioles of previous years leaves. Stems solitary, erect, simple, wingless. Leaves firm, leathery,
abaxially pale green, adaxially green, both surfaces glabrous or sparsely pubescent only along midvein and margins, apex acuminate, margin sinuate-dentate. Basal leaves withered at anthesis, petiolate; petioles short, usually up to 5 cm. Lower and middle stem leaves shortly petiolate to sessile; leaf blade elliptic, 4-10 x 1.5-3 cm. Upper stem leaves sessile, narrowly elliptic to linear-lanceolate, 3-9 x 1-2 cm. Capitula (1-)4-20, shortly pedunculate, in a dense corymbiform synflorescence. Involucre narrowly campanulate, 12-15 x 5-7. Phyllaries in 4-6 rows, abaxially from light green to dark colored, pubescent; inner phyllaries narrowly elliptic, apex subobtuse, outer phyllaries equal or slightly shorter, apex acuminate with a straight or slightly recurved cusp. Receptacle densely covered with persistent whitish subulate-filiform bristles. Corolla purple. Anthers dark-colored, 4-4.5 mm long, basal appendages obtuse, with lacerate tails, up to 1 mm long. Pappus of two series, inner row of whitish yellow (darker at the base) plumose bristles connated at base, 10.5-11.5 mm long, outer series of scabrid unequal bristles. Achenes glabrous, cylindric, 4-5 mm, usually more or less purple due to purple dots. The main characteristics that make it possible to distinguish S. arctecapitulata from morphologically related species, which include S. klementzii, S. subacaulis, S. tilesii (Ledeb.) Ledeb., S. parviflora, S. pseudoalpina, are presented in Table 2.
Table 2
Comparative features of morphologically similar species to Saussurea arctecapitulata
Species Stem Leaves Phyllaries
Saussurea arctecapitulata (5-)10-40 cm, wingless, simple both surfaces green (abaxially pale green, adaxially green), glabrous or sparsely pubescent only along mid-vein and margins pubescent; inner subobtuse; outer equal or slightly shorter, apex acuminate with a straight or slightly recurved cusp
Saussurea klementzii (5-)10-50 cm, wingless, simple or apically branched both surfaces green (abaxially pale green, adaxially green), gland-dotted; stem leaves are half-stem-embracing with auricles glabrous or sparsely pubescent; inner subobtuse; outer clearly shorter, apex acuminate with a straight or slightly recurved cusp
Saussurea subacaulis 5-15(-20) cm, wingless, simple abaxially grayish green and tomen-tose, adaxially green and glossy pubescent, apex acuminate; outer equal or slightly shorter than inner
Saussurea pseudoalpina 5-25(-40) cm; wingless or rarely with short and narrow wings; simple abaxially grayish green, gland-dotted and arachnoid, adaxially green, sparsely pubescent pubescent, gland-dotted; inner subobtuse; outer clearly shorter, apex acuminate
Saussurea parviflora 20-100(-150) cm, narrowly winged; simple or apically branched both surfaces green (abaxially pale green, adaxially green), gland-dotted; glabrous or abaxially sparsely pubescent with uniseriate trichomes glabrous or sparsely pubescent along margins, apex subacute to obtuse; outer clearly shorter
Saussurea tilesii 5-10(-15) cm, wingless, simple both surfaces usually gland-dotted, abaxially grayish green and tomen-tose, adaxially green and glossy pubescent; inner subobtuse; outer slightly shorter, apex acuminate with a straight or slightly recurved cusp
Fig. 1. Bayesian 50 % majority-rule consensus phylogenetic tree from nrDNA ITS1-5.8S-ITS2 site for Saussurea arctecapitulata and the related taxa with indication of Bayesian Posterior Probability (PP) values for the internal tree nodes.
Fig. 2. Saussurea arctecapitulata: А - synflorescence; B - achene; C - involucre; D - receptacle bristles; E - habit (photos by E. A. Pyak).
Pyak E. A. et al.
Saussurea arctecapitulata Lipsch. - new species for the flora of Russia
Distribution. - Saussurea arctecapitulata has been considered as an endemic of Mongolia, known from the Khangai Mountains and Western Prihubsugul region (Lipschitz, 1972). However, the specimens from the Khangai Mountains, preserved in LE and MW belong to S. parviflora (Poir.) DC., a common species widely distributed in the Altai and Sayan Mountains and adjacent regions. Two species can be distinguished by involucre (with phyllaries of equal length in S. arctecapitulata vs. imbricate in S. parviflora) and stems ((5-)10-40 cm, wingless vs. 20-100(-150) cm, narrowly winged). To our knowledge, there is no other specimen collected in the Khangai Mountains that can be assigned to S. arctecapitulata. In Wehrden et al. (2009) S. arctecapitulata was mentioned from Gobi Altai as a diagnostic species of Kobresietum myosuroidis communities restricted to the moistest north-facing slopes of the Gobi Gurvan Saykhan, but the specimen preserved in HAL belongs to S. pseudoalpina N. D. Simpson, widely distributed in Altai Mountains. Thus, to date, the species is known only from the crossborded territory, from Khövsgöl Aimag of Mongolia (Lake Khubsugul) and Republic of Tuva of Russia (Naryn River Valley).
Specimens examined: "Mongolia. Lake Kosogol. 18 VII 1902. V. Komarov / Монголия. Озеро Ко-согол. 18 VII 1902. В. Комаров" (LE [LE 01025564; LE 01025565]); "[Mongolia]. Dzuun Saikhan, Yoliin Am catchment, mountain meadow (grazed) replacing Betula - Salix scrub on valley ground, 23002350 m. 43°29'N, 104°06'E. 02 VII 1996. G. Meihe, S. Meihe. 96-010-10" (HAL [HAL 50155]); "Mongolia. Khangai, 30 km from Tevchruleg Village, subalpine lawns on top of Mount Khairkhan, 2500 m. 24 VII 1980. I. A. Gubanov. 990 / Монголия. Хангай, 30 км на ЮЮЗ от пос. Тэвшрулег Архангайско-го аймака, субальпийские лужайки на вершине г. Хайрхан, 2500 m. 24 VII 1980. И. А. Губанов. 990" (LE; MW); "Mongolia. Khangai, upper reaches of the Urd-Tamir River, near the Holsain-Daba Pass, 2600-2700 m. 4 VIII 1980. I. A. Gubanov. 1124 / Монголия. Хр. Хангай, верховья р. Урьд-Тамира у перевала Холсайн-Даба, 2600-2700 m. 4 VIII 1980. I. A. Gubanov. 1124" (LE; MW).
Acknowledgements
The research was supported by the grant of the Russian Science Foundation (project No. 23-2400400).
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