CC BY
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Turczaninowia 23, 4: 88-98 (2020) DOI: 10.14258/turczaninowia.23.4.9 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
УДК 582.284.52:581.95(579.154-751.2)
Boletoid fungi (Boletaceae, Basidiomycota) of the Bidoup - Nui Ba
National Park (Vietnam)
T. H. G. Pham1*, O. V. Morozova2
1 Joint Russian-Vietnamese Tropical Research and Technological Center, 63 Nguyen Van Huyen Str., Cau Giay, Hanoi, Vietnam.
*E-mail: giangvietnga@gmail.com
2 Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Str., RUS-197376, Saint Petersburg, Russia.
E-mail: ovm.leptonia@gmail.com
Keywords, biodiversity, Boletaceae, boletoid basidiomycetes, Central Highlands of Vietnam, protected areas, Tay Nguyen, tropical forests.
Summary. Data on the diversity of boletoid fungi of the Bidoup - Nui Ba National Park are presented here and analyzed. An annotated list of 13 species are published for the first time for the National Park and 8 for the first time for Vietnam (Porphyrellus nigropurpureus, Phylloporuspachycystidiatus, Ph. rubiginosus, Pulveroboletus brun-neopunctatus, Strobilomyces brunneolepidotus, S. calidus, Veloporphyrellus pseudovelatus, Xerocomus subparvus). Among plant communities of the National Park the mountain evergreen coniferous-broad-leaved forests dominated by Fagaceae, Lauraceae and Magnoliaceae and with the participation of representatives of Podocarpaceae and Pinaceae are characterized by the highest diversity of boletoid fungi. The species are illustrated with color photographs. The nucleotide sequences obtained during the study were deposited in NCBI GenBank.
1 Совместный Российско-Вьетнамский Тропический научно-исследовательский и технологический центр, ул. Нгуен Ван Хуен, д. 63, Кау Зай, г. Ханой, Вьетнам
2Ботанический институт им. В. Л. Комарова РАН, ул. Проф. Попова, д. 2, г. Санкт-Петербург, 197376, Россия
Ключевые слова: биоразнообразие, болетоидные базидиомицеты, особо охраняемые природные территории, тропические леса, Тэйнгуэн, Центральное нагорье Вьетнама, Boletaceae.
Аннотация. В настоящей работе представлены сведения о разнообразии болетовых грибов национального парка Бидуп-Нуйба и дан их краткий анализ. Приведен аннотированный список из 13 видов, впервые публикуемых для национального парка, из которых 8 - впервые для Вьетнама (Porphyrellus nigropurpureus, Phylloporus pachycystidiatus, Ph. rubiginosus, Pulveroboletus brunneopunctatus, Strobilomyces brunneolepidotus, S. calidus, Veloporphyrellus pseudovelatus, Xerocomus subparvus). Среди растительных сообществ национального парка наибольшим разнообразием болетовых грибов характеризуются горные вечнозеленые хвойно-широ-колиственные леса с преобладанием Fagaceae, Lauraceae, Magnoliaceae и участием представителей Podocarpaceae и Pinaceae. Виды проиллюстрированы цветными фотографиями. Нуклеотидные последовательности, полученные в ходе исследования, депонированы в NCBI Genbank.
* Corresponding author
Болетовые грибы (Boletaceae, Basidiomycota) национального парка Бидуп - Нуйба (Вьетнам)
Т. Х. Ж. Фам1, О. В. Морозова2
2
Поступило в редакцию 03.08.2020 Принято к публикации 20.11.2020
Submitted 03.08.2020 Accepted 20.11.2020
Introduction
Boletoid fungi due to well-developed mycelium and more or less large basidiomata play a significant role in tropical ecosystems, forming mycorrhizal associations with forest-forming tree species of more than 10 families, including Dipterocarpaceae, Fabaceae, Fagaceae, and Pinaceae, improving their nutrition and growth, increasing resistance to adverse environmental influences, as well as participating in the decomposition of organic matter (Thoen, Ba, 1989; Sato et al., 2007; Becerra, Zak 2011). Many of them are important as a source of nutrition for humans and as potential producers of biologically active substances for medical purposes, what is very actual in recent years. The Southeast Asia region is the center of diversity for this group.
The aim of the present work was to study the diversity of boletoid fungi in the Bidoup - Nui Ba National Park, located in the central part of the Dalat Plateau. Large tracts of mountain and foothill forests have been preserved here. The study of the fungal diversity of typical tropical forests complements the information on the biota of macromycetes in Vietnam and may be useful in the development of measures for the conservation of these valuable nature areas and species inhabiting them.
Bidoup - Nui Ba National Park
Bidoup - Nui Ba National Park (VQG Bidoup Nui Ba) is located in the northwest of Lam Dong province, Lac Doung District, in the Central Highlands of Vietnam (Tay Nguyen), in the central part of the Dalat plateau between 12°00'-12°19'N and 108°21'-108°44'E, occupying an area of 648 km2. The park relief is dominated by mountains with heights of more than 1400 m a. s. l., the highest peaks exceed 2000 m - Bidoup (2287 m), Lang Bian (2167 m) and Hon Giao (2062 m). The hydrographic network of the National Park belongs to Krong-No (Krong No) River basin (flowing to Mekong (Me Kong)) and Da Nhim River (flowing to Dong Nai (Dong Nai)). Climate is tropical monsoon. The average annual temperature is 18 °C, in winter in the mountains there can be short-term frosts down to -0.1 °C. Average annual rainfall is about 1800 mm (Eames, 1995; Sourcebook..., 2004; Brodribb, Field, 2008).
The vegetation cover is dominated by mountain and alpine evergreen broadleaf and mixed forests. Secondary pyrogenic pine forests with Pinus kesiya are widespread, accounting for about a third of the park's area.
The species of Fagaceae (Castanopsis pseudoserrata, C. indica, Lithocarpus stenopus, L. truncatus, Quercus braianensis) and Lauraceae (Cinnamomum, Litsea) dominate in the stand of evergreen broadleaf forests. In mixed forests, conifers complement the upper tier: Pinus dalatensis, P. krempfii, Fokienia hodginsii, Dacrycarpus imbricatus, Keteleeria evelyniana. Juglandaceae (Engelhardtia roxburghiana), Theaceae (Schima wallichii, Anneslea fragrans), Magnoliaceae, Myr-taceae, Hamamelidaceae (Rhodoleia championii), Aceraceae (Acer campbellii), Simplocaceae (Sym-plocos lucida) are also common in the stand. Above 1700 m a. s. l., crooked cloud forests with dominance of Ericaceae (Vaccinium sprengelii, V. viscifolium), Fagaceae (Lithocarpus silvicolarum, L. dalatensis Castanopsis wilsonii, Quercus poilanei) Sapotaceae (Palaquium ridleyi) are widespread (Eames, 1995; Tordoff et al., 2004; Kuznetsov et al., 2006; Tran, 2011; Pham, Le, 2012; Novozhilov et al., 2020).
Materials and methods
Collections were made in different types of tropical forests in Bidoup - Nui Ba National Park during 2010-2014 in course of the investigation of mycobiota of the Central Highlands (Tay Nguyen). Boletoid fungi in the territory of the National Park were collected in the following main biotopes:
Mountain evergreen coniferous-broadleaved forests dominated by Fagaceae, Lauraceae and Magnoliaceae and with the participation of representatives of the Podocarpaceae and Pinaceae (1500-1700 m a. s. l.).
Middle mountain open pyrogenic coniferous forest dominated by Pinus kesiya (1500-1700 m a. s. l.).
High mountain cloud forests dominated by Ericaceae and Fagaceae (1700-2000 m a. s. l.).
The collected specimens were examined using light and electron microscopy, as well as by analyzing DNA sequences (ITS and tef1a sites). Macromorphological features have been studied in fresh collections and later through detailed photographs taken in the field. Microscopical characters were studied with a light Zeiss Axioscope A1 microscope with AxioCam 1Cc 3 camera and program tools AxioVisionRel.4.6. (Carl Zeiss, Germany). SEM photos were also used for the identification of material. They have been made by L. A. Kartseva on a scanning analytical electron microscope JEOL JSM-6390LA of Core Facility Centre 'Cell and Molecular Technologies in Plant
Science' of Komarov Botanical Institute and by A. V. Alexandrova (Lomonosov State University) on scanning analytical electron microscopes JSM-6380LA and Camscan-S2 (Cambridge Instruments, 1990). Basidiospores, basidia, and hymenial cystidia were observed in squash preparations of small parts of the tubes in 5 % KOH. The pileipellis was examined on a radial section of the pileus, the stipitipellis - on longitudinal slice of the stipe in 5 % KOH. Basidiospore dimensions are based on 20 measurements, whereas cystidia and basidia dimensions are based on observing at least 10 structures per collection.
The molecular study is based on a phylogenetic species recognition (Taylor et al., 2000). To assess the taxonomic status of the identified phylogenetic lineages and put forward species hypotheses, we relied on the accepted boundaries of the possible variability of the ITS1-5.8S-ITS2 and tefla nDNA regions within the species (Petersen et al., 2008; Hughes et al., 2009).
DNA was extracted from herbarium material using NucleoSpin® Plant II kit (Macherey-Nagel, Düren, Germany). The ribosomal ITS1-5.8S-ITS2 region was amplified with primers ITS1F and ITS4B (Gardes, Bruns, 1993), and elongation translation factor (tefla) with Boletaceae-specific primers EF1-B-F1 and EF1-B-R (Wu et al., 2014). PCR products were purified with the Fermentas Genomic DNA Purification Kit (Thermo Fisher Scientific, Waltham, MA) and sequenced on an ABI model 3130 Genetic Analyzer (Applied Biosystems, Waltham, MA). Raw data were edited and assembled in MEGA X (Kumar et al., 2018). Newly generated sequences were deposited in NCBI GenBank (Table 2).
Collections studied are kept in the Herbarium of the Komarov Botanical Institute (LE).
History of the studies on of boletoid fungi in the Bidoup - Nui Ba National Park
Information on the history of the investigation of macromycetes in the Central Highlands of Vietnam, including the Bidoup - Nui Ba National Park, is presented in the works of Morozova et al. (2012) and Pham et al. (2018).
The table below (Table 1) summarizes the published data on Boletaceae in Bidoup - Nui Ba National Park, including species described as new to science from this area at different times.
A series ofworks on the diversity ofmacromycetes of the Central Highlands, containing information on 24 species of boletoid fungi, belongs to Lê Bâ Düng (Le, 2001, 2003). However, these works do
not indicate the exact locations of the fungi, so we cannot consider these findings as collected in the territory of the Bidoup - Nui Ba National Park.
It should be noted that previously species
were identified on the basis of morphological and ecological characters only, and often for identification of paleotropic species European and North American keys were used. Species described from Madagascar also present in the list. In light of recent data on the geographical distribution of fungi, such determinations are questionable and require confirmation by the molecular methods. In this regard, we take into account only the species, which collections were confirmed by molecular data (including those determined only to the genus) and / or described directly from the territory of the National Park. Attempts to isolate DNA from old type specimens, unfortunately failed.
Results and discussion
Thirteen species not previously recorded for the Bidoup - Nui Ba National Park are listed below, including eight species published for the first time for Vietnam (marked with "!").
Annotated list of species reported for the first
time for the Bidoup - Nui Ba National Park
Subfamily Austroboletoideae G. Wu et Zhu L.
Yang
Austroboletus cf. mucosus (Corner) Wolfe: "Vicinities of Giang Ly Ranger Station, 12°11'5.6"N, 108°41'12.3"E, 1540 m a. s. l., on soil in a mediummountain broad-leaved forest with the participation of conifers. 29 V 2014" (LE 315558, fig. 1a). Our collections differ from the protologue (Corner, 1972) in a very light color. Young basidiomata are almost white, turn yellowish with age, while in the original description they are completely yellow.
! Veloporphyrellus pseudovelatus Yan C. Li et Zhu L. Yang: "Vicinities of Klong Lanh Village, the left bank of the Da Nhim River, path to Bidoup Mt, 12°07'44"N, 108°39'22"E, 1570 m a. s. l., at the base of the Pinus kesiya in the secondary pine forest (Pinus kesiya). 25 VI 2010" (LE 315608, fig. 1b).
Subfamily Boletoideae Burnett
! Porphyrellus nigropurpureus (Hongo) Y. C. Li et Zhu L. Yang: "Vicinities of Giang Ly Ranger Station, the left bank of the Da Nhim River, on soil in a medium-mountain broad-leaved forest with the participation of Lauraceae, Myrtaceae, Fagaceae, Magnoliaceae. 24 VI 2010" (LE 254399, fig. 1c); ibid. 29 VI 2010 (LE 254400).
* The names of taxa are given in accordance with the data of the Index Fungorum resource (as of July 2020) with indication of the synonyms under which they were indicated in the corresponding publications.
Table 1
Species noted in publications, indicating the geographical origin of their types
Виды* Patouillard, 1909 Perreau, Joly, 1964 Dörfelt et al., 2004 Ton, 2009 Nguyen et al., 2015 Pham et al., 2015 Crous et al., 2018 Crous et al., 2019 Geographical origin of the type
Austroboletus subflavidus (МштШ) Wolfe + North America
Boletellus ananas (M. A. Curtis) Murrill + North America
Boletellus annamiticus (Pat.) E.-J. Gilbert [=Strobilomyces annamiticus Pat.; = Boletellus emodensis (Berk.) E.-J. Gilbert ss. auct.] + Vietnam: Bidoup -Nui Ba
Boletus auripes Peck + North America
Boletus bouriquetii R. Heim + Madagascar
Boletus edulis Bull. + Europe
Boletus queletii var. aurantiacus Dörfelt, Kiet et A. Berg [? =Crocinoboletuspinetorum N. R. Zeng, L.-L. Wu, Zhi Q. Liang, S. Jiang] + + Vietnam: Bidoup -Nui Ba
Boletus speciosus Frost + North America
Fistulinella olivaceoalba T. H. G. Pham, Yan C. Li et O. V. Morozova + Vietnam: Bidoup -Nui Ba
Neoboletus erythropus (Pers.) C. Hahn [=Boletus erythropus Pers.; =Tubiporus erythropus (Pers.) Maire] + Europe
Phylloporus rhodoxanthus (Schwein.) Bres. + Europe
Phylloporus sulcatus (Pat.) E.-J. Gilbert [=Paxillus sulcatus Pat.] + + Vietnam: Bidoup -Nui Ba
Pulveroboletus ravenelii (Berk. et M. A. Curtis) Murrill + North America
Rugiboletus extremiorientalis (Lj. N. Vassiljeva) G. Wu et Zhu L. Yang [=Leccinum extremiorientale (Lar. N. Vassiljeva) Singer] + Russian Far East
Strobilomyces strobilaceus (Scop.) Berk. [=Strobilomyces floccopus (Vahl) P. Karst.] + Europe
Veloporphyrellus vulpinus T. H. G. Pham, O. V. Morozova, A. V. Alexandrova et E. S. Popov + Vietnam: Bidoup -Nui Ba
Xerocomus langbianensis Dörfelt, Kiet et A. Berg bis + + Vietnam: Bidoup -Nui Ba
Total: 17 2 5 3 2 3 2 1 1
! Strobilomyces brunneolepidotus Har. Takah et Taneyama: "Vicinities of Hon Giao Ranger Station, path to Hon Giao Mt, 12°11'18.1"N, 108°42'50.6"E, 1700 m a. s. l., on soil in upper montane mossy evergreen broadleaf forest. 28 VI 2010" (LE 315609, fig. 1d).
! S. calidus Li H. Han, J. Xu & Zhu L. Yang: "Vicinities of Giang Ly Ranger Station, 12°10'42.1"N, 108°41'00.7"E, 1510 m a. s. l., on
soil in a medium-mountain broad-leaved forest with the participation of conifers. 27 V 2014 (LE 315562, fig 1e.).
Subfamily Leccinoideae G. Wu et Zhu L. Yang Pseudoaustroboletus valens (Corner) Y. C. Li et Zhu L. Yang, in Li, Li, Zeng, Cui et Yang: "Vicinities of Giang Ly Ranger Station, 12°11'05.6"N, 108°41'12.3"E, 1540 m a. s. l., on soil in a medium-
mountain broad-leaved forest with the participation (LE 312475). "Ibid. 29 V 2014" (LE 315553, fig. of conifers. 28 VI 2010" (LE 315593). "Ibid., leg. 1f). A. E. Kovalenko, Ngoc Nguyen Bao. 04 XI 2012"
Table 2
Distribution of Boletaceae species of the Bidoup - Nui Ba National Park by biotope types with Genbank numbers
Species Types of biotops Altitude, m a. s. l. Voucher numbers Genbank numbers
1 2 3 ITS tefla
Aureoboletus longicollis + 1500 LE 315562 LE 312514 MT893598 MW033312
Austroboletus cf. mucosus + 1540 LE 315558 MT893602 -
Austroboletus sp. + 1550 LE 312524 MT893601 -
Boletus queletii var. aurantiacus [? =Crocinoboletus pinetorum] + 1600 V 136, HAL 1768, holotype - -
Boletellus annamiticus + 1500 FH 3720 (LBA 161), lectotype - -
Boletellus sp. + 1600 LE 315630 LE 254394 - -
Crocinoboletus rufoaureus + + 1700 LE 315594 LE 315561 LE 312520 - -
Fistulinella olivaceoalba + 1850 LE 312004, holotype MH718344 MH733592
Leccinellum sp. + 1550 LE 312515 - MW033313
Phylloporus pachycystidiatus + 1550 LE 312522 MT893596 -
Phylloporus rubiginosus + 1480 LE 312523 MT893597 -
Phylloporus sulcatus (Pat.) E.-J. Gilbert [=Paxillus sulcatus Pat.] + 1500 FH 3725 (LBA 210), holotype - -
Phylloporus sp. + 1550 LE 312519 - MW033321
Porphyrellus nigropurpureus + 1500 LE 254399 LE 254400 MT893603 MT893604 -
Pseudoaustroboletus valens + 1540 LE 312475 LE 315593 LE 315553 MT893599 MW033322
Pulveroboletus brunneopunctatus + 1520 LE 315598 - -
Strobilomyces brunneolepidotus + 1700 LE 315609 - MW033320
Strobilomyces calidus + 1510 LE 312518 - MW033317
Strobilomyces sp. 1 + 1500 LE 312516 - MW033316
Strobilomyces sp. 2 + 1400 LE 312517 - MW033319
Veloporphyrellus pseudovelatus + 1570 LE 315608 - MW033318
Veloporphyrellus vulpinus + 1500 LE 315544, holotype MN511177 MN597966
Xerocomus langbianensis + 1600 Dl 9903, HAL 769, isotype - -
Xerocomus subparvus + 1490 LE 315595 MT893600 MW033315
Zangia roseola + 1500 LE 315610 - MW033314
Total: 25 20 3 3
Subfamily Xerocomoideae Singer
Aureoboletus longicollis (Ces.) N. K. Zeng et Ming Zhang, in Zeng, Zhang et Liang: "Vicinities of Giang Ly Ranger Station, 12°10'46.8"N, 108°41'08.5"E, 1500 m a. s. l., on soil in a mediummountain broad-leaved forest with the participation of conifers. 25 V 2014" (LE 312514, fig. 2a). "Ibid. 12°10'43.3"N, 108°41'05.3"E. 27 V 2014" (LE 315562).
! Phylloporuspachycystidiatus N. K. Zeng, Zhu L. Yang & L. P. Tang: "Vicinities of Klong Lanh Village, the left bank of the Da Nhim River, path to Bidoup Mt, 1550 m a. s. l., on soil in a mediummountain broad-leaved forest with the participation of Lauraceae, Myrtaceae, Fagaceae, Magnoliaceae. 25 VI 2010" (LE 312522, fig. 2b).
! Ph. rubiginosus M. A. Neves & Halling: "Vicinities of Giang Ly Ranger Station, 12°10'54.1"N, 108°41'00.5"E, 1480 m a. s. l., on soil in a medium-mountain broad-leaved forest with the participation of conifers. 25 V 2014" (LE 312523, fig. 2c).
! Xerocomus subparvus Xue T. Zhu et Zhu L. Yang, in Wu, Li, Zhu, Zhao, Han, Cui, Li, Xu et Yang: "Vicinities of Giang Ly Ranger Station, 12°10'46.8"N, 108°41'00"E, 1490 m a. s. l., on soil in a medium-mountain broad-leaved forest with the participation of conifers. 25 V 2014" (LE 315595, fig. 2d).
Subfamily Zangioideae G. Wu, Y. C. Li et Zhu
L. Yang
Zangia roseola (W. F. Chiu) Y. C. Li et Zhu L. Yang, in Li, Feng et Yang: "Vicinities of Klong Lanh Village, path to Bidoup Mt, 12°07'57.3"N, 108°39'02.8"E, 1500 m a. s. l., on soil in a mediummountain broad-leaved forest with the participation of conifers. 24 V 2014" (LE 315610, fig. 2e).
Pulveroboletus-group
Crocinoboletus rufoaureus (Massee) N. K. Zeng, Zhu L. Yang et G. Wu, in Zeng, Wu, Li, Liang et Yang: "Vicinities of Hon Giao Ranger Station, path to Hon Giao Mt, 12°11'18.1"N, 108°42'50.6"E, 1700 m a. s. l., on soil in upper montane mossy evergreen broadleaf forest. 28 VI 2010" (LE 315594). "Ibid. 26 V 2014" (LE 315561, fig. 2f). "Vicinities of Klong Lanh Village, path to Bidoup Mt, 12°07'57.3"N, 108°39'02.8"E, 1500 m a. s. l., on soil in a medium-mountain broad-leaved forest with the participation of conifers. 24 V 2014" (LE 312520).
! Pulveroboletus brunneopunctatus G. Wu et Zhu L. Yang, in Wu, Li, Zhu, Zhao, Han, Cui, Li,
Xu et Yang: "Vicinities of Giang Ly Ranger Station, 12°11'06.7"N, 108°41'13.9"E, 1520 m a. s. l., on soil in a medium-mountain broad-leaved forest with the participation of conifers. 29 V 2014" (LE 315598, fig. 2g).
In addition, thanks to molecular genetic methods, 6 genotypes were revealed, which could not be attributed to any known species either by molecular or morphological data. There are: Austroboletus sp., Boletellus sp., Leccinellum sp., Phylloporus sp., and two species of Strobilomyces.
Thus, we can say about the presence of 25 species of Boletaceae family in the studied area, which were confirmed by the molecular data or were taken into account as species with a type locality in a National Park (Table 2). The species are distributed between 14 genera, belonging to five subfamilies and the Pulveroboletus group.
Among plant communities of the National Park the mountain evergreen coniferous-broad-leaved forests dominated by Fagaceae, Lauraceae and Magnoliaceae and with the participation of representatives of Podocarpaceae and Pinaceae are characterized by the highest diversity of boletoid fungi. The most favorable conditions for their growth have developed in these types of forest: the largest number of possible symbionts grows; the temperature and humidity indicators are optimal. In addition, the largest number of observations was carried out in these biotopes. In secondary pine and high mountain foggy and crooked forests the diversity of the boletoid fungi was found to be distinctly lower.
The available information on the nutritional properties of tropical boletes is very limited. We can only assume them, based on knowledge of the phylogenetic relationships in this group of fungi. Boletellus annamiticus and Crocinoboletus rufoaureus are most likely edible, as the very closely related species Boletellus emodensis and Crocinoboletus laetissimus are eaten (L^ssoe et al., 2018; Wu et al., 2019). Zangia roseola considered edible (Wu et al., 2019). Aureoboletus viscosus and some species of the genus Strobilomyces regarded as potentially edible (L^ss0e et al., 2018). Therefore, the related Aureoboletus longicollis and other species of the genus Strobilomyces can also be considered as potentially edible. It is known that Pulveroboletus brunneopunctatus, as well as other species of this genus, are poisonous (Chen et al., 2014, L^ss0e et al., 2018). However, this question requires further study.
Fig. 1. Basidiocarps in situ: a - Austroboletus cf. mucosus (LE 315558); b - Veloporphyrellus pseudovelatus (LE 315608); c - Porphyrellus nigropurpureus (LE 254399); d - Strobilomyces brunneolepidotus (LE 315609); e -Strobilomyces calidus (LE 315562); f - Pseudoaustroboletus valens (LE 315553).
Fig. 2. Basidiocarps in situ: a - Aureoboletus longicollis (LE 315562); b - Phylloporus pachycystidiatus (LE 312522); c - Phylloporus rubiginosus (LE 312523); d -Xerocomus subparvus (LE 315595); e - Zangia roseola (LE 315610); f - Crocinoboletus rufoaureus (LE 315561); g - Pulveroboletus brunneopunctatus (LE 315598).
Acknowledgments
The authors are grateful to staff of the Joint Russian-Vietnamese Tropical Research and Technological Center for organizing the expeditions and to management of the Bidoup - Nui Ba National Park for their help and for permission to collect in the forests. We thank Drs. E. S. Popov (Komarov Botanical Institute) for the help with collecting, A. V. Alexandrova (Moscow State University) and L. A. Kartseva (Komarov Botanical Institute) - for their
assistance with the SEM photos. We are grateful to Curators of Herbariums HAL and FH for the opportunity to study type collections. The work was conducted in the framework of a research project of the Komarov Botanical Institute of the Russian Academy of Sciences (no. AAAA-A19-119020890079-6) using equipment of its Core Facility Centre 'Cell and Molecular Technologies in Plant Science' with the financial support of Russian Foundation for Basic Research (project no. 20-04-00349).
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