CC BY
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Turczaninowia 24, 3: 65-76 (2021) DOI: 10.14258/turczaninowia.24.3.5 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 protected areas of Kon Tum Plateau (Central Highlands of Vietnam)
1 Joint Russian-Vietnamese Tropical Research and Technological Center, Nguyen Van Huyen Str., 63, Cau Giay, Hanoi, Vietnam 2 Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popova Str., 2, Saint Petersburg, 197376, Russia
Keywords, biodiversity, boletoid basidiomycetes, Central Highlands of Vietnam, Kon Chu Rang, Kon Ka Kinh, Kon Plong, Tay Nguyen, tropical forests.
Summary. The article continues the series of publications devoted to the boletoid fungi of Vietnam. It summarizes the results of the observation of their diversity in three nature protected areas of the Central Highlands (Tay Nguyen) -Kon Ka Kinh National Park, Kon Chu Rang (Kon Chu Rang) Nature Reserve and Kon Plong Protected Forest, where middle-mountain evergreen broad-leaved and coniferous-deciduous forests prevail at altitudes of 900-1500 m. An annotated list of 22 revealed species is presented. Of these, two species (Fistulinella aurantioflava and Tylopilus subotsuensis) have been recently described from this territory as new to science. Hortiboletus rupicapreus is reported "ad interim". Information on eight species (Aureoboletus sinobadius, Hourangia nigropunctata, Ionosporus longipes, Neoboletus multipunctatus, Phylloporus luxiensis, Pulveroboletus subrufus, Tylopilus atripurpureus, T. atroviolaceo-brunneus) is published for the first time for Vietnam. Strobilomyces aff. echinocephalus, S. aff. glabriceps, and Tylopilus aff. balloui (two different lines) are close to known species but genetically not identical. Six more specimens are identified only to the genus, and they probably represent species new to science. The color photographs of new to Vietnam and noteworthy species are presented. The nucleotide sequences obtained during the study were deposited in NCBI GenBank.
Болетовые грибы (Boletaceae, Basidiomycota) особо охраняемых природных территорий плато Контум (Центральное нагорье Вьетнама)
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
3Московский государственный университет им. М. В. Ломоносова, Биологический факультет, 119234, Ленинские горы, д. 1, стр. 12, г. Москва, Россия
Ключевые слова: биоразнообразие, болетоидные базидиомицеты, Конкакинь, Конплонг, Контьыранг, особо охраняемые природные территории, тропические леса, Тэйнгуэн, Центральное нагорье Вьетнама.
Поступило в редакцию 26.04.2021 Submitted 26.04.2021
Принято к публикации 03.08.2021 Accepted 03.08.2021
T. H. G. Pham1, 4, O. V. Morozova2, 5*, A. V. Alexandrova1, 3, 6
3Lomonosov Moscow State University, Leninskie Gory Str., 1, 12, Moscow, 119234, Russia
4ORCID iD: https://orcid.org/0000-0002-4137-7213 5E-mail: [email protected]; ORCID iD: https://orcid.org/0000-0002-7329-528X 6ORCID iD: https://orcid.org/0000-0003-2359-4780
Т. Х. Ж. Фам1, О. В. Морозова2, А. В. Александрова1, 3
Аннотация. Статья продолжает серию работ, посвященных болетовым грибам Вьетнама. Здесь обобщены результаты изучения разнообразия этой группы грибов трех охраняемых природных территорий Центрального нагорья (Тэйнгуэн) - национального парка Конкакинь, природного заповедника Контьыранг и охраняемого лесного массива Конплонг, где на высотах 900-1500 м преобладают среднегорные широколиственные и хвой-но-широколиственные леса. Приводится аннотированный список 22 идентифицированных видов. Из них два недавно описаны с этой территории как новые для науки виды (Fistulinella aurantioflava и Tylopilus subotsuen-sis). Hortiboletus rupicapreus приводится "ad interim". Информация о восьми видах (Aureoboletus sinobadius, Hourangia nigropunctata, lonosporus longipes, Neoboletus multipunctatus, Phylloporus luxiensis, Pulveroboletus subrufus, Tylopilus atripurpureus, T. atroviolaceobrunneus) публикуется впервые для Вьетнама. Strobilomyces aff. echinocephalus, S. aff. glabriceps и Tylopilus aff. balloui незначительно отличаются генетически от соответствующих известных видов. Еще шесть образцов определены только до рода и, вероятно, представляют новые для науки виды. Приводятся цветные фотографии новых для Вьетнама и редких видов. Нуклеотидные последовательности, полученные в ходе исследования, депонированы в GenBank NCBI.
Introduction
This article continues the series of publications devoted to the diversity of the boletoid fungi in Vietnam (Pham et al., 2018; Pham, Morozova, 2020). The results of the work in three nature protected areas of the Central Highlands (Tây Nguyên) - Kon Chu Rang (Kon Chu Rang) Nature Reserve, Kon Ka Kinh National Park and Kon Plong Protected Forest - are summarized here. Before the beginning of our research, there was no information on the mycobiota of this territory. Then, the data on the agaricoid genera Entoloma, Volvariella, Pluteus were published (Morozova et al., 2018; Malysheva et al., 2019, 2020).
An annotated list of 22 species of Boletaceae is presented here. Of these, two species (Fistulinella aurantioflava and Tylopilus subotsuensis) have been recently described by us from this territory as new to science (Crous et al., 2020, 2021). Hortiboletus rupicapreus is reported here "ad interim", it will be published soon. Eight species (Aureoboletus sinobadius, Hourangia nigropunctata, Ionosporus longipes, Neoboletus multipunctatus, Phylloporus luxiensis, Pulveroboletus subrufus, Tylopilus atripurpureus, and T. atroviolaceobrunneus) are published for the first time for Vietnam. But most of them (by the exclusion of Aureoboletus sinobadius and Tylopilus atripurpureus) were reported previously in the manuscript of the dissertation work of the first author (Pham, 2020).
Materials and methods
Collections were made by route method in different types of tropical forests on Kon Tum plateau in 2015-2017 in course of the investigation of mycobiota of the Central Highlands (Tây Nguyên).
The information on the nature conditions of the territories studied is provided below.
Kon Ka Kinh National Park
Kon Ka Kinh National Park (Vuon quoc gia Kon Ka Kinh) is located in the northern part of Gia Lai Province (Mang Yang County, A Yun Commune) between 14.15°N-14.5°N and 108.25°E-108.45°E and covers an area of 417.8 km2. About 80 % (331.46 km2) of the park's territory is covered with forests. The territory of the park lies between Pleiku and Kon Ha Nung plateaus and has mountainous terrain with an average altitude of 1200-1500 m a. s. l. (the highest Kon Ka Kinh - 1748 m). The rivers flowing through the park belong to the basins of the Ba, Pne, and A Yun rivers. The park's climate is tropical monsoon, with distinct rainy (May - November) and dry (December - April) seasons. The average annual temperature ranges from 21 to 25 °C. The average annual precipitation is from 2000 to 2500 mm, the peak of precipitation falls in July-August (400-450 mm). There is practically no precipitation in January - February (Sourcebook of Existing ..., 2004; Ha et al., 2011, 2014).
The forest vegetation of Kon Ka Kinh National Park is represented by three main types (Thai, 1978; Le et al., 2000). The most widespread are mountain evergreen broad-leaved forests, which occupy heights of 900-1000 m a. s. l. and higher. They are characterized by the dominance of species of Fagaceae (Castanopsis, Lithocarpus, Quercus), Lauraceae, Euphorbiaceae, Meliaceae, Sterculiaceae, Theaceae, Rubiaceae. In mountain evergreen mixed forests found south of the Kon Ka Kinh Mt, at altitudes from 1300 m, in addition to species from the families Fagaceae, Theaceae, Euphorbiaceae, Hamamelidaceae, Elaeocarpaceae, Myrtaceae, and Meliaceae, species of conifers present (Fokienia hodginsii, Dacrycarpus imbricatus, Dacrydium
elatum, Nageia fleuryi, Podocarpus neriifolius). Below 900 m a. s. l., low-mountain evergreen broad-leaved forests are developed, dominated by Dipterocarpaceae (Shorea siamensis, S. roxburghii), Fabaceae, Irvingiaceae, Euphorbiaceae, Moraceae, Burseraceae, Lauraceae, Apocynaceae, Meliaceae, Fagaceae, and Magnoliaceae. Rather large area is occupied by plantings of Pinus latteri.
Kon Chu Rang Nature Reserve
The KBTTN Kon Chu Rang is located in the northeastern part of Gia Lai Province (K'Bang District, Son Lang Commune), between 14.5°N-14.58°N and 108.5°E-108.65°E, and is bordered by the provinces of Binh Dinh, Quang Ngai, and Kon Tum. The area of the reserve is 159 km2, of which 156.1 km2 (99 %) are primary and intact forests. The relief ofthe reserve is mostly hilly, mountainous in the northern part, with heights from 800 to 1452 m a. s. l. (Kon Chu Rang). The average annual temperature is about 21 °C (maximum in May +28 °C, minimum in January +12 °C). The average annual precipitation is about 1900-2000 mm, the peak of precipitation falls in September (340 mm). The dry season is relatively short, with up to 60 mm of precipitation per month in January -April (New et al., 2002; Sourcebook of Existing ..., 2004). Numerous rivers flowing through the reserve belong to basin of the Kon River (Song Kon), which has a number of waterfalls, the most famous and tallest of which is 50 m high.
The main forest type is middle-mountain evergreen broad-leaved and mixed forest distributed at elevations between 900 and 1500 m in the northwest ofthe nature reserve. Canopy cover ofthis forest type is 70 to 80 %, and the tree flora is dominated by species from the Fagaceae (Lithocarpus, Quercus, Castanopsis), Lauraceae, Fabaceae, Clusiaceae, Myrtaceae, Ericaceae, Burseraceae, and Magnoliaceae, mixed with gymnosperms (Dacrycarpus imbricatus, Dacrydium elatum). Lowland evergreen forest occurs at elevations below 900 m. Only 2 % of the nature reserve is covered by secondary vegetation, mainly scrub with scattered trees.
Kon Plong Protected Forest
The Kon Plong Protected Forest is located in the northeast of Kon Tum Province (Kon Plong County), within the state forestry enterprises of Mang Canh II, Tan Lap, and the Ngok tem (Ngok tem) Water Protection Forest. The massif has an area of 650.8 km2 and is located on the highly dissected mountain
basalt plateau Kon Ha Nung (Kon Ha Nung), at an altitude of 1100 to 1757 m a. s. l. The studied territory is located in the basin of the Lo (Bäk Lo), Te (Nuöc Che) rivers and the lower reaches of the Khe (Bäk Khe) river within 14.7125°N-14.7589°N and 108.3°E-108.3233°E. The climate is characterized by moderate temperatures (average annual temperature is about +18 °C, and the coldest month +16 °C) and heavy rainfall (up to 2800 mm per year).
The vegetation cover above 1000 m a. s. l. is dominated by middle-mountain evergreen mixed forests, the upper tier of which is dominated by Fagaceae (Lithocarpus, Quercus, Castanopsis), Lauraceae, Hamamelidaceae, Theaceae, Podo-carpaceae (Dacrycarpus imbricatus, Dacrydium elatum, Podocarpus neriifolius), Pinaceae (Pinus dalatensis, Keteleeria evelyniana), Cupressaceae (Fokienia hodginsii), etc. There are areas of monodominant coniferous forests with Pinus latteri. At altitudes up to 900-1000 m a. s. l. low-mountain evergreen broadleaf forests are developed with the participation of Dipterocarpaceae (Shorea siamensis, Parashorea stellata, Dipterocarpus obtusifolius), Fagaceae, Myrtaceae, Sterculiaceae, Annonaceae, etc. On a large area, natural forests are disturbed by economic activity and are replaced by secondary forests, bamboo, shrub and herbaceous communities (Eames et al., 2001; Pham et al., 2001).
Morphological study
Macromorphological features were studied on the basis of fresh and dried material, as well as through the analysis of photographs and descriptions taken in the field.
Micromorphological structures were studied using herbarium material. The collected specimens were examined using light and electron microscopy, as well as by analyzing DNA sequences (ITS and tefla sites). Microscopical characters were studied with a light Zeiss Axioscope A1 microscope with AxioCam1Cc 3 camera and program tools AxioVisionRel.4.6. (Carl Zeiss, Germany). 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.
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).
Molecular-genetic study
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).
Collections studied are kept in the Mycological Herbarium of the Komarov Botanical Institute (LE).
Results and discussion
Twenty-two species revealed in the Kon Tum plateau are listed below including ten species published for the first time for Vietnam (marked with "!").
Annotated list of species of Boletoid fungi of the Kon Chu Rang Nature Reserve, Kon Ka Kinh National Park, and Kon Plong Protected Forest
Subfamily Austroboletoideae G. Wu et Zhu L. Yang
Fistulinella aurantioflava T. H. G. Pham, A. V. Alexandrova et O. V. Morozova: "Kon Ka
Kinh National Park, 14.217129°N, 108.310132°E, 1220 m a. s. l., on slopes of mountain, on soil in the tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. A. V. Alexandrova" (LE 315616) (Pham, 2020, as "Pulveroboletus" curtisii; Crous et al., 2021).
Subfamily Boletoideae Burnett
Hortiboletus rupicapreus Svetash., A. V. Alexandrova, O. V. Morozova et T. H. G. Pham, ad interim: "Kon Ka Kinh National Park, 14.219917°N, 108.325667°E, 1200 m a. s. l., ridges on slopes, on soil in tropical mountain forests dominated by Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 18 V 2016. A. V. Alexandrova" (LE 312677); ibid., "14.205190°N, 108.316312°E, 1000 m a. s. l., on soil in plantations of Pinus kesiya with some Fagaceae. 15 V 2016. A. V. Alexandrova" (LE 312678).
Porphyrellus nigropurpureus (Hongo) Y. C. Li et Zhu L. Yang: "Kon Plong Protected Forest, Mang Canh Commune, 5 km north of Kondu Village, 14.73295°N, 108.31203°E, 1100 m a. s. l., on soil in polydominant rainforest with participation of Podocarpaceae, Magnoliaceae, Myrtaceae, Calophyllaceae, Elaeocarpaceae. 10 VI 2016. O. V. Morozova" (LE 315621).
! Strobilomyces aff. echinocephalus Gelardi et Vizzini: "Kon Ka Kinh National Park, 14.219917°N, 108.325667°E, 1200 m a. s. l., ridges on slopes, on soil in tropical mountain forests dominated by Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 18 V 2016. O. V. Morozova" (LE 312681, fig. 1a, b).
! Strobilomyces aff. glabriceps W. F. Chiu: "Kon Ka Kinh National Park, on slopes of the mountain, 14.217129°N, 108.310132°E, 1220 m a. s. l., on soil in tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. O. V. Morozova" (LE 312682, fig. 1c).
Strobilomyces seminudus Hongo: "Kon Ka Kinh National Park, 14.20414°N, 108.32143°E, 970 m a. s. l., on soil in a middle-mountain evergreen broadleaf forest with the participation of families Magnoliaceae, Myrtaceae, Theaceae, Lauraceae, Fagaceae. 20 V 2016. O. V. Morozova" (LE 315628, fig. 1d, e, f).
! Tylopilus atripurpureus (Corner) E. Horak: "Kon Ka Kinh National Park, 14.20414°N, 108.32143°E, 970 m a. s. l., on soil in a middle-mountain evergreen broadleaf forest with the
participation of families Magnoliaceae, Myrtaceae, Theaceae, Lauraceae, Fagaceae. 20 V 2016. O. V. Morozova" (LE 312683, fig. 1g).
! Tylopilus atroviolaceobrunneus Yan C. Li et Zhu L. Yang: "Kon Ka Kinh National Park, on soil
in a tropical middle-mountain polydominant forest with a predominance of tree families Myrtaceae, Meliaceae, Fagaceae, Dipterocarpaceae. 15 V 2016. O. V. Morozova" (LE 315627, fig. 1h).
Table
Boletaceae species in the nature protected areas studied, with voucher and Genbank numbers
Protected GenBank numbers
areas
Species Kon Ka Kinh Kon Chu Rang Kon Plong Alt. (m) LE numbers ITS tef1a
Aureoboletus sinobadius + 1220 LE 312680 MZ451340 MZ424885
Boletellus areolatus + + 980 1030 LE 315577 LE 315576 MZ451341 MZ451342 -
Boletus sp. 1 + 1300 LE 312687 MZ451343 -
Boletus sp. 2 + 1050 LE 312698 MZ451344 -
Chalciporus sp. 1 + 1000 LE 312688 MZ451345 MZ424888
Chalciporus sp. 2 + 1000 LE 312689 MZ451346 MZ424889
Chiua viridula + + 1000 LE 315611 LE 315612 MZ451347 MZ424887
Crocinoboletus rufoaureus + 1280 LE 315607 LE 315561 - MZ424876 (from Bidoup - Nui Ba NP)
Fistulinella aurantioflava + 1220 LE 315616 MW784159 -
Hortiboletus rupicapreus + 1000 1200 LE 312677 LE 312678 MW784161 MW784162 MZ424893 MZ424894
Hourangia nigropunctata + 1270 LE 315626 MZ451348 -
Ionosporus longipes + 1000 LE 315589 MZ451349 MZ424890
Neoboletus multipunctatus + 830 1080 1280 LE 315625 LE 312474 LE 315637 MZ451350 MZ451351 -
Neoboletus obscureumbrinus + 1090 LE 315548 MZ451352 MZ424891
Phylloporus luxiensis + 1020 LE 315622 - MZ424877
Phylloporus rubiginosus + 1000 LE 315623 MZ451353 -
Phylloporus sp. 1 + 1000 LE 312684 - MZ424892
Phylloporus sp. 2 + 1020 LE 312685 MZ451354 -
Porphyrellus nigropurpureus + 1100 LE 315621 - MZ424878
Pulveroboletus subrufus + 1050 LE 312686 MZ451355 MZ424879
Strobilomyces aff. echinocephalus + 1200 LE 312681 - MZ424880
Strobilomyces aff. glabriceps + 1220 LE 312682 - MZ424881
Strobilomyces seminudus + 970 LE 315628 MZ451356 -
Tylopilus atripurpureus + 970 LE 312683 - MZ424884
Tylopilus atroviolaceobrunneus + 1000 LE 315627 - MZ424882
Tylopilus aff. balloui 1 + 1300 LE 312532 MZ451357 MZ424883
Tylopilus aff. balloui 2 + 1000 LE 312700 MZ451358 MZ424886
Tylopilus subotsuensis + 1000 LE 312527 - MW014272
Total 20 6 4
28
Fig. 1. Basidiomata in situ: a, b - Strobilomyces aff. echinocephalus (LE 312681); c - Strobilomyces aff. glabriceps (LE 312682); d, e, f - Strobilomyces seminudus (LE 315628); g - Tylopilus atripurpureus (LE 312683); h - Tylopilus atroviolaceobrunneus (LE 315627); scale bar = 1 cm.
! Tylopilus aff. balloui [1] (Peck) Singer: "Kon Ka Kinh National Park, 14.21988°N, 108.30936°E, 1300 m a. s. l., on the soil in a tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 20 V 2016. A. V. Alexandrova" (LE 312532).
! Tylopilus aff. balloui [2] (Peck) Singer: "Kon Chu Rang Nature Reserve, 14.50516°N, 108.58140°E, 1000 m a. s. l., on soil in middle-mountain evergreen mixed forest. 27 V 2016. A. V. Alexandrova" (LE 312700).
Tylopilus subotsuensis T. H. G. Pham, A. V. Alexandrova et O. V. Morozova: "Kon Chu Rang Nature Reserve, 14.50042°N, 108.56338°E, 1000 m
a. s. l., on soil in middle-mountain evergreen mixed forest. 27 V 2016. A. V. Alexandrova" (LE 312527) (Crous et al., 2020).
Subfamily Leccinoideae G. Wu et Zhu L. Yang
! Ionosporus longipes (Masee) Khmeln., Davoodian, Raspé, S. M. L. Lee et Hailing: "Kon Chu Rang Nature Reserve, Son Lang Commune, vicinity of waterfalls, 14.51413°N, 108.54630°E, 1000 m a. s. l., on the soil in a middle-mountain evergreen mixed forest with a predominance of Podocarpaceae (Dacrydium elatum, Dacrycarpus imbricatus), Magnoliaceae, Burseraceae, Myrtaceae. 25 V 2016. O. V. Morozova" (LE 315589, fig. 2a).
Fig. 2. Basidiomata in situ: a - Ionosporus longipes (LE 315589); b - Aureoboletus sinobadius (LE 312680); c -Hourangia nigropunctata (LE 315626); d - Phylloporus luxiensis (LE 315622).
Subfamily Xerocomoideae Singer
! Aureoboletus sinobadius Ming Zhang et T.H. Li: "Kon Ka Kinh National Park, 14.21226°N, 108.31318°E, 1090 m a. s. l., on the soil in tropical mountain polydominant forests with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. A. V. Alexandrova" (LE 312680, fig. 2b).
Boletellus areolatus Hirot. Sato: "Kon Ka Kinh National Park, 14.20948°N, 108.31276°E, 980 m a. s. l., on soil in a middle-mountain evergreen broadleaf forest with the participation of families Magnoliaceae, Myrtaceae, Theaceae, Lauraceae, Fagaceae. 16 V 2016. O. V. Morozova" (LE 315577); "Kon Plong Protected Forest, Mang Canh Commune, 5 km north of Kondu Village, 14.72222°N, 108.316°E, 1030 m a. s. l., on soil in a middle-mountain polydominant rainforest, 23 IV 2015. A. V. Alexandrova" (LE 315576).
! Hourangia nigropunctata (W.F. Chiu) Xue T. Zhu et Zhu L. Yang: "Kon Ka Kinh National Park, 14.222330°N, 108.310347°E, 1270 m a. s. l., on the soil in a tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 20 V 2016. A. V. Alexandrova" (LE 315626, fig. 2c).
! Phylloporus luxiensis M. Zang: "Kon Ka Kinh National Park, 14.209184°N, 108.314681°E, 1020 m a. s. l., on the soil in a tropical mountain polydominant forest with a predominance of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. A. V. Alexandrova" (LE 315622, fig. 2d).
Phylloporus rubiginosus M. A. Neves et Halling: "Kon Ka Kinh National Park, 14.2187°N, 108.3168°E, 1000 m a. s. l., on the soil in tropical middle-mountain polydominant forest dominated by Lauraceae, Myrtaceae, Meliaceae, Fagaceae, Dipterocarpaceae. 15 V 2016. O. V. Morozova" (LE 315623).
Subfamily Zangoideae G. Wu, Y. C. Li et Zhu L. Yang
Chiua viridula Y.C. Li et Zhu L. Yang: "Kon Ka Kinh National Park, 14.20791°N, 108.31500°E, 995 m a. s. l., on the soil in a middle-mountain middle-mountain polydominant forest with a predominance of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. E. S. Popov" (LE 315611); "Kon Chu Rang Nature Reserve, vicinity of waterfalls, 14.51413°N, 108.54630°E, 1000 m a. s. l., on the soil in a middle-mountain evergreen mixed forest with a predominance of Podocarpaceae (Dacrydium
elatum, Dacrycarpus imbricatus), Magnoliaceae, Burseraceae (Canarium), Myrtaceae (Syzygium). 25
V 2016. O. V. Morozova" (LE 315614, fig. 3a, b); ibid. "01 VI 2016. O. V. Morozova" (LE 315612).
Pulveroboletus-group
Crocinoboletus rufoaureus (Massee) N. K. Zeng, Zhu L. Yang et G. Wu: "Kon Plong Protected Forest, 5 km north of Kondu Village, headwaters of the La and Khe rivers, 14.7455°N, 108.3019°E, 1280 m a. s. l., middle-mountain polydominant forest dominated by Podocarpaceae, Magnoliaceae, Myrtaceae, Calophyllaceae, Elaeocarpaceae, Betulaceae. 05 VI 2016. A. V. Alexandrova" (LE 315607).
! Neoboletus multipunctatus N. K. Zeng, H. Chai et S. Jiang: "Kon Ka Kinh National Park, 14.211846°N, 108.313694°E, 1080 m a. s. l., on soil in a tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. A. V. Alexandrova" (LE 315625, fig. 3c); ibid, "14.21946°N, 108.30940°E, 1280 m a. s. l. 20
V 2016. A. V. Alexandrova" (LE 312474); ibid., "Krong commune, K'bang district, 14.310920°N, 108.438320°E, 830 m a. s. l., on soil in tropical middle-mountain polydominant forest with the participation Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 13 V 2017. A. V. Alexandrova" (LE 315637).
Neoboletus obscureumbrinus (Hongo) N. K. Zeng, H. Chai et Zhi Q. Liang [= Sutorius obscureumbrinus (Hongo) G. Wu et Zhu L. Yang]: "Kon Ka Kinh National Park, 14.21226°N, 108.31318°E, 1090 m a. s. l., on the soil in tropical mountain polydominant forests with the participation of representatives of the families Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16
V 2016. A. V. Alexandrova" (LE 315548, fig. 3e, f).
! Pulveroboletus subrufus N.K. Zeng et Zhu L. Yang: "Kon Ka Kinh National Park, 14.21052°N, 108.31434°E, 1050 m a. s. l., on the soil in a tropical mountain polydominant forest with the participation of Myrtaceae, Meliaceae, Anacardiaceae, Fagaceae, Theaceae. 16 V 2016. A. V. Alexandrova" (LE 312686, fig. 3d).
Totally, the presence of 28 species of Boletaceae in the studied area were confirmed by the molecular data (Table). Strobilomyces aff. echinocephalus, S. aff. glabriceps, and Tylopilus aff. balloui (two different lines) morphologically are close to known species but genetically not identical with the
Fig. 3. Basidiomata in situ: a, b - Chiua viridula (LE 315614); c - Neoboletus multipunctatus (LE 315625); c - Pul-veroboletus subrufus (LE 312686); e, f - Neoboletus obscureumbrinus (LE 315548).
sequences kept in the NCBI GenBank. In addition, 6 more genotypes were revealed on the base of the molecular study. They could not be attributed to any known species either by molecular or morphological data; there are: Boletus sp. 1, Boletus sp. 2, Chalciporus sp. 1, Chalciporus sp. 2, Phylloporus sp. 1, Phylloporus sp. 2.
The species are distributed between 17 genera belonging to 5 subfamilies and the Pulveroboletus group. Among the territories studied, the highest diversity of boletoid species was found in the Kon Ka Kinh National Park due to the higher diversity of the communities represented there and more favorable weather conditions during the collection period. The mountain evergreen broad-leaved and coniferous-deciduous forests of this park are richest in the diversity of the Boletaceae species.
Acknowledgments
The authors are grateful to staff of the Joint Russian-Vietnamese Tropical Research and Tech-
nological Center for organizing the expeditions and to management of the Kon Ka Kinh National Park, Kon Chu Rang Nature Reserve, and Kon Plong Protected Forest for their help and permission to collect in the forests. We thank Dr. E. S. Popov (Ko-marov Botanical Institute) for his assistance with collecting and providing literature. The study of Olga Morozova was conducted in the framework of a research project of the Komarov Botanical Institute of the Russian Academy of Sciences (no. AAAA-A18-118022090078-2) 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). The study of Alina Alexandrova was conducted as part of the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University (no. 121032300081-7).
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