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J. Fungi 2023, 9, 621. https://doi.org/10.3390/jof9060621 www.mdpi.com/journal/jof
Article
New Species of Entoloma Subgenera Cubospora and Leptonia
(Agaricales, Basidiomycota) from Central Vietnam
Olga Morozova
1,
*
and Thi Ha Giang Pham
2,
*
1
Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Str.,
197022 Saint Petersburg, Russia
2
Joint Vietnam-Russia Tropical Science and Technology Research Centre, Nguyen Van Huyen,
Nghia Do, Cau Giay, Hanoi 122100, Vietnam
* Correspondence: omorozova@binran.ru (O.M.); giangvietnga@gmail.com (T.H.G.P.)
Abstract: Four new species of Entoloma from Kon Chu Rang Nature Reserve and Ta Dung National
Park were discovered during an investigation of the diversity of the mycobiota of Central Vietnam
and are described here on the base of the molecular and morphological data. Phylogenetic analysis
was based on nrITS1-5.8S-ITS2, nrLSU and tef1α regions. Illustrated descriptions of their macro- and
microscopic features and discussion on similar taxa are given. Entoloma cycneum and E. peristerinum
belong to the subgenus Cubospora. They are morphologically similar species and are characterized
by white or whitish basidiomata with yellowish or beige tinges and with mainly smooth, glabrous,
and hygrophanous pileus, longitudinally fibrillose or fibrillose-scaly white stipe, cuboid spores, and
more or less cylindrical cheilocystidia, arising from hymenophoral trama. Entoloma peristerinum
posseses initially more coloured beige conical pileus, discolouring to white with age and drying.
The pileus of E. cycneum is initially white, hemisphaerical to convex, usually with thin pubescence
near the margin. The species can be recognized also by the cheilocystidia form: serrulatum-type in
E. cycneum vs. porphyrogriseum-type in E. peristerinum. Another two species belong to the subgenus
Leptonia. Entoloma tadungense is close to E. percoelestinum from which it differs by smaller spores with
pronounced angles, presence of the cheilocystidia, and the lilac discolouration of the stipe. E.
dichroides is named after its similarity to E. dichroum, a dark blue coloured species with
pronouncedly angled basidiospores. It is distinguished by the basidiospores form—irregularly 5(–
6) angled with elongated apiculus, as well as by absence of the cheilocystidia and darker
basidiomata with conical pileus. The article also describes the history of the study of the genus
Entoloma in Vietnam with a list of 29 species mentioned in the publications for this country.
Keywords: taxonomy; systematics; Entolomataceae; molecular phylogeny; new species; Kon Chu
Rang Nature Reserve; Ta Dung National Park
1. Introduction
According to estimates for 2018 [1], 21 species of the genus Entoloma (Fr.) P. Kumm.
were known from the Central Vietnam, including unpublished data of the authors. In
subsequent years, five more new species were described from this territory [2–4]. The full
history of study of the genus Entoloma in Vietnam with a list of 29 species mentioned in
the publications for this country is described in the discussion part. However, the real
diversity of the genus is much higher, and many species are still waiting to be described.
As proven by recent molecular genetic studies, the shape of the basidiospores is a
key feature in understanding evolution and relatedness in Entolomataceae [5–7].
The cuboid and cuboid-like form of spores is remarkable and interesting from a
phylogenetic point of view in terms of the multiplicity of its occurrence in the course of
evolution. The molecular genetic data [7] supports a clear segregation of clades with
cuboid spores from cuboid-like ones (pentagonal, prismatic, pseudocuboid,
Citation: Morozova, O.; Pham,
T.H.G. New Spec ies of Entoloma
Subgenera Cubospora and Leptonia
(Agaricales, Basidiomycota) from
Central Vietnam. J. Fungi 2023, 9,
621. hps://doi.org/10.3390/
j
of9060621
Academic Editor: Vladimir Antonin
Received: 25 April 2023
Revised: 24 May 2023
Accepted: 25 May 2023
Published: 27 May 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Swierland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Aribution (CC BY) license
(hps://creativecommons.org/license
s/by/4.0/).
J. Fungi 2023, 9, 621 2 of 24
squamiferum). The species with rhomboid spores also cluster separately in a well-
supported clade but they are mixed with species with five or six angled spores [8,9].
Species with “true” cuboid spores (with six, more or less equal, quadrangular faces)
traditionally were included in the subgenus/genus Inocephalus [Entoloma subgen.
Inocephala Noordel., Inocephalus (Noordel.) P. D. Orton.]. However, the type of Inocephalus,
Entoloma inocephalum (Romagnesi) Dennis, described from Madagascar, has 5–7 angled
basidiospores in profile view [10]. The material identified as Entoloma inocephalum from
Vietnam [11] has the same morphology and is used as a reference collection for the
phylogenetic studies while authentic or Madagascar-collected material is not available [7].
It turns out that the /Inocephalus clade occupies an isolated position from the clades with
cuboid spores. Therefore, two new subgenera were described to arrange species with such
spores—Entoloma subgenus Cubospora Karstedt, Capelari, Largent, T. J. Baroni &
Bergemann with E. luteolamellatum (Largent & Aime) Blanco-Dios as a type species (with
a conical pileus) and Entoloma subgenus Cuboeccilia Karstedt, Capelari & Largent with E.
omphalinoides (Largent) Blanco-Dios as a type species (with depressed pileus) [7]. The
number of species with cuboid and cuboid-like spores described world-wide so far is
estimated to be 165 [12] or 120 [7].
The first monographic contributions devoted to groups with cuboid spores were
made by Horak [13–17], and refer to SE-Asia and South America. The following
publications deal with the diversity of cuboid-spored species in different regions, e.g.,
Asia [11,12,18–21], Africa/Madagascar [10,22–24], Australia/Tasmania/New Zealand
[25,26], South and Central America [27–30], and North America [31,32]. Generalization of
accumulated data based on phylogenetic analysis was conducted by F. Karstedt et al. [7].
The analysis of records shows that the centre of diversity of the group with “true” cuboid
spores is situated in tropical and subtropical regions of the Old and New Worlds. Two
more species of the subgenus Cubospora are described from Vietnam here.
Another two new species belong to the subgenus Leptonia in the modern sense [8].
The multigene phylogeny based on nrITS, nrLSU, and mtSSU [5] showed that subgenus
Leptonia in the previously considered volume [32–36] is polyphyletic. Sect. Leptonia of the
subgenus belongs to the /Nolanea-Claudopus clade, and Cyanula and Griseorubida to the
/Inocephalus-Cyanula clade. Based on these data, Cyanula has been raised to the subgenus
level [26]. Recent studies [37,38] have shown that a clade of species around Entoloma
ameides (sect. Ameides (Largent) Reschke et al.) previously classified under the subgenus
Nolanea now is grouped with the subgenus Leptonia as a sister to the /Leptonia clade with
high support. For that reason, the /Ameides clade is treated as a new section of subgenus
Leptonia [8]. The subgenus Leptonia now combines species with more or less coloured
(predominantly with blue, brown, but also yellow or green tinges) basidiomata,
characterized also by the presence of clamp-connections, absence of brilliant granules, and
more or less fibrillose to squamulose stipe, and includes around 50 species world-wide
[8,26,32,39]; Gasteroid forms also occur [37]. The highest diversity of this group was found
in the temperate zone of the Holarctic. In the tropics, representatives of the subgenus are
rare, and in Vietnam they are recorded for the first time.
2. Materials and Methods
2.1. Collecting and Site Description
The material for this study was collected during the expeditions of the Joint Vietnam-
Russia Tropical Science and Technology Research Centre (VRTC) to the Central Highlands
of Vietnam.
The Kon Chu Rang Nature Reserve is located in the north-eastern part of Gia Lai
Province of Vietnam (Son Lang Commune, K’Bang District), between 14.50° N–14.58° N
and 108.5° E–108.65° E. The area of the reserve is 15,446 hectares, of which 99% are primary
and intact forests, which is the highest forest coverage in the country. The relief is hilly to
mountainous in the northern part, with heights from 800 to 1452 m a. s. l. (Kon Chu Rang
J. Fungi 2023, 9, 621 3 of 24
Mt). The average annual temperature is about 21 °C (from 28 °C in May to 12 °C in
January). The average annual precipitation is about 1900–2000 mm with the peak in
September (340 mm). The dry season is from January to April [40,41]. The main forest type
in the reserve is middle-mountain evergreen broad-leaved and mixed forest dominated
by Fagaceae (Lithocarpus, Quercus, Castanopsis), Lauraceae, Fabaceae, Clusiaceae,
Myrtaceae, Ericaceae, Burseraceae, and Magnoliaceae, mixed with gymnosperms
(Dacrycarpus imbricatus, Dacrydium elatum), distributed at elevations between 900 and 1500
m in the north-west of the nature reserve. The first data on the mycobiota of Kon Chu
Rang Nature Reserve were published only recently, including those on several subgenera
of Entoloma, and on the Boletaceae [3,4,42,43].
Ta Dung National Park is located in the Central Highlands of Vietnam within the
administrative boundary of Dak Som commune, Dak G’long district, Dak Nong province,
50 km northeast of Gia Nghia commune’s exam centre. It extends between 11.79° N–11.99°
N and 107.89° E–108.11° E, occupying a total area of 20,973.7 hectares. The National Park
is located on the Dak Nong Plateau and part of the Di Linh Plateau. Dak Nong Plateau is
an arched elevation with an average height of 600–1200 m and has many mountain ranges
with an average height of 1200–1500 m, with Ta Dung peak being 1982 m high, the lowest
elevation being the land. Ta Dung National Park is situated in an area with a tropical
highland monsoon climate regime with two distinct seasons, the rainy season lasts from
April to October, the dry season—from November to March. The average annual
temperature is 22.0 °C. The total average annual rainfall is 2339 mm; precipitation falls
mainly from May to October. Ta Dung National Park is characterized by subtropical
humid evergreen closed forests and mixed broadleaf and coniferous closed forests [44].
The mycobiota of the National Park previously was not studied.
Specimens were photographed in the field, and their macromorphological characters,
such as size, colour, shape, and surface of all parts of the basidiomata as well as odour,
were documented before drying. Colour codes refer to Kornerup & Wanscher [45]. GPS
coordinates of collection site, habitat, and substrate type were also documented for each
collection. Specimens were then dried either in airtight plastic containers with silica gel,
or with an electric dryer at a temperature ca. 50 °C, placed on a piece of absorbent paper
and packed in plastic Ziploc bags with small amounts of silica gel.
2.2. Morphological Study
Microscopic measurements and drawings were made with an AxioScope A1 light
microscope equipped with Zeiss AxioCam 1Cc3 digital camera with AxioVisionRel.4.6
software (CarlZeiss, Jena, Saxe-Weimar-Eisenach, Germany). Spores, basidia, and cystidia
were observed in squash preparations of small parts of the lamellae in 5% KOH or 1%
Congo Red in concentrated NH4OH. The pileipellis was examined from a radial section
of the pileus in 5% KOH. Basidiospore dimensions were based on 20 spores; cystidia and
basidia dimensions on at least 10 structures per collection. Basidia were measured without
sterigmata, and the spores without apiculus. Spore length to width ratios were reported
as Q. When studying the specimens and compiling morphological descriptions, we
followed the recommendations and terminology of [8]. The dried specimens were
deposited in the Mycological Herbarium of the Komarov Botanical Institute RAS (LE) and
in the Herbarium of the Joint Vietnam–Russia Tropical Science and Technology Research
Centre, Hanoi (VRTC).
2.3. DNA Extraction, Amplification, and Sequencing
PCR products were obtained without DNA purification step using the Thermo
Scientific Phire Tissue Direct PCR Master Mix (Thermo Fisher Scientific, MA, USA)
standard protocol. The ribosomal ITS1–5.8S–ITS2 region was amplified with the fungal
specific primers ITS1F and ITS4B [46];
hp://www.biology.duke.edu/fungi/mycolab/primers.htm, accessed on 1 March 2017.
Sequences of nrLSU-rDNA were generated using primers LR0R and LR5 [47]. Primers
J. Fungi 2023, 9, 621 4 of 24
EF1-983F and EF1-1567R were used to amplify approximately 500 bp of tef1 [48]. For ITS,
PCR was carried out under the following cycling parameters: initial denaturation: 98 °C
for 4 min; followed by 35 cycles: 98 °C for 1 min, 52 °C for 1 min, and 72 °C for 1 min, and
final extension at 72 °C for 3 min. For nrLSU: initial denaturing at 98 °C for 5 min; then 12
cycles of denaturing at 98 °C for 5 s, annealing at 67 °C for 1 min, extension at 72 °C for
1.5 min; then 35 cycles of denaturing at 98 °C for 5 s, annealing at 56 °C for 1 min, extension
at 72 °C for 1.5 min; and a final extension step of 72 °C for 10 min. For tef1-a: initial
denaturing at 98 °C for 5 min; then 8 cycles of denaturing at 98 °C for 5 s, annealing at 60
°C for 40 s, extension at 72 °C for 2 min; then 36 cycles of denaturing at 98 °C for 5 s,
annealing at 53 °C for 1.5 min, extension at 72 °C for 2 min; and a final extension step of
72 °C for 10 min.
PCR products were visualized using agarose gel electrophoresis and Gel Red
staining, and subsequently purified with the Fermentas Genomic DNA Purification Kit
(Thermo Fisher Scientific Inc., Waltham, MA, USA). Sequencing was performed with an
ABI model 3500 Genetic Analyzer (Applied Biosystems, CA, USA).
This work was carried out using equipment of the Core Facility Centre ‘Cell and
Molecular Technologies in Plant Science’ of the Komarov Botanical Institute. Raw data
were edited and assembled in MEGA X [49]. Newly generated sequences have been
deposited in the GenBank.
2.4. Alignment and Phylogenetic Analyses
For this study, 12 nrITS and 9 tef1α, and 10 nrLSU sequences were newly generated.
In addition, 38 nrITS and 20 tef1α, and 43 nrLSU sequences, including outgroups, were
retrieved from the GenBank database (www.ncbi.nlm.nih.gov/genbank, accessed 15
March 2023), using the BLASTn application (hps://blast.ncbi.nlm.nih.gov/Blast.cgi,
accessed 15 March 2023). The information on all these sequences is presented the Table 1.
Table 1. Specimens and GenBank accession numbers of DNA sequences used in the molecular
analyses (newly generated sequences are in bold).
Species Location Voucher Numbe
r
Genbank Accession No. References
ITS LSU tef
Clitopilopsis hirneola (Fr.) Kühner (Clitopilus
hirneolus (Fr.) Kühner & Romagn.) Italy MEN 199956 — GQ289211 — [5]
Clitopilus prunulus (Scop.) P. Kumm. Belgium MEN 2003-09-14 — GQ289149 — [5]
Entoloma abortivum (Berk. & M.A. Curtis) Donk Russia: Far East LE 312419 MF476905 MF487792 — [42]
E. acutipallidum E. Horak & Cheype Brazil: Pará SP: FK1893 — MG018325 MH190147 [7]
E. albidoquadratum Manim. & Noordel. India: Kerala PM667 (L) — GQ289151 — [5]
E. allochroum Noordel. The Netherlands v. Waveren, 29-07-
1973, holotype (L) KC898372 — — [39]
E. altissimum (Massee) E. Horak Vietnam LE 262945 MF476912 — — [42]
E. argillaceum O.V. Morozova et al. Russia: Caucasus LE 311861, holotype — OL338531 OL405537 [38]
E. araneosum (Quél.) M.M. Moser Belgium MEN 200314 KC710056 GQ289153 — [5]
E. atricolor O.V. Morozova et al. Vietnam LE 295000, holotype KY777496 — — [42]
E. aurantiovirescens
Reschke, Lotz-Winter & Noordel. Panama KaiR623, holotype MZ611665 — — [50]
E. austriacum Courtec. [E. tjallingiorum var.
laricinum O.V. Morozova et al.] Russia: Far East LE 254343, holotype KC898413 KC898513 — [39]
E. bidupense O.V. Morozova & E.S. Popov Vietnam LE 262935 — NG_059265 — [42]
E. callichroum E. Horak & Noordel. Switzerland ZT 71/58, holotype KC898350 — — [39]
E. cervinum (Karstedt & Capelari) Blanco-Dios
[Inocephalus cervinus Karstedt & Capelari] Brazil SP: FK 0940 — — MH190138 [7]
E. cetratum (Fr.) M.M. Moser Russia: European
part LE 235480 KC898450 — — [39]
J. Fungi 2023, 9, 621 5 of 24
E. chalybaeum (Pers.) Noordel. Russia: European
part LE 254353 KC898445 KC898500 — [39]
E. chytrophilum Wölfel, Noordel. & Dähncke Spain: Canary
Islands L 855, holotype KC898434 KC898519 — [39]
E. coelestinum (Fr.) Hesler Russia: Ural LE 258103 KC898362 KC898524 — [39]
E. conferendum (Britzelm.) Noordel. MEN 200330 KC710055 KC710133 — [51]
E. cycneum O.V. Morozova et T.H.G. Pham Vietnam LE F-343654,
holotype OQ779461 OQ804518 OQ779183 this work
E. cycneum Vietnam LE F-343655 OQ779463 OQ804519 OQ779182 this work
E. cycneum Vietnam LE F-343656 OQ779462 OQ804521 OQ779181 this work
E. cycneum Vietnam LE F-343657 OQ779464 OQ804520 OQ779184 this work
E. cycneum Vietnam LE F-343658 OQ779465 — OQ779185 this work
E. cystidiophorum Dennis [Inocephalus
cystidiophorus (Dennis) Karstedt & Capelari] Brazil: São Paulo SP: FK1023 — MG018319 MH190140 [7]
E. dennisii E. Horak [Inocephalus dennisii (E.
Horak) Karstedt & Capelari] Puerto Rico CORT: 8263 TJB — — MH190164 [7]
E. dichroides O.V. Morozova et T.H.G. Pham Vietnam LE F-343682 OQ779472 OQ804527 — this work
E. dichroum (Pers.) P. Kumm. Russia: European
part LE 227472, neotype KC898440 — — [39]
E. dichroum Russia: European
part LE 234260 — KC898527 — [39]
E. euchroum (Pers.) Donk Russia: Caucasus LE 262995 KC898417 KC898516 — [39]
E. eugenei Noordel. & O.V. Morozova
Russia:
Primorsky
Territory
LE 253771, holotype KC898438 KC898529 — [39]
E. griseocyaneum (Fr.) P. Kumm. Russia: Caucasus LE 254351 KC898444 KC898498 — [39]
E. inocephalum (Romagn.) Dennis Vietnam LE 262922 KC898449 MH259311 MH190154 [7,39]
E. kovalenkoi O.V. Morozova, E.S. Popov & A.V.
Alexandrova Vietnam LE 312529 OK257210 OK257207 OK256169 [3]
E. kruticianum O.V. Morozova, M.Y. Dyakov,
E.S. Popov & A.V. Alexandrova
Russia: European
part LE 311767, holotype KU666558 KU710222 — [52]
E. lampropus (Fr.) Hesler Sweden UPS:BOT:F-176490,
neotype KC898377 KC898506 — [39]
E. lasium (Berk. & Broome) Noordel. & Co-
David [Pouzarella lasia (Berk. & Broome)
Largent & Abell-Davis]
Australia:
Queensland DLL9811 (BRI, CNS) — — MG702641 [7]
E. lepidissimum (Svrček) Noordel. Czech Republic PRM 755801,
holotype KC898364 KC898532 — [39]
E. luteolamellatum (Largent & Aime) Blanco-
Dios [Trichopilus luteolamellatus Largent &
Aime]
Guyana MCA 1480, holotype — MH190213 MG702644 [7]
E. luteum Peck [Inocephalus luteus (Peck) T.J.
Baroni] USA: New York CORT: 7771 TJB — MH190212 MH190161 [7]
E. mucronatum (Karstedt & Capelari) Blanco-
Dios [Inocephalus mucronatus Karstedt &
Capelari]
Brazil: São Paulo SP: FK1033, holotype — MH190174 MH190141 [7]
E. murrayi (Berk. & M.A. Curtis) Sacc. & P. Syd. China MHHNU 30602 MK250917 — — [53]
E. murrayi USA CORT: TJB 8210 — MH190193 — [7]
E. nitidum Quél. Slovakia MEN 200324 KC710122 GQ289175 — [5]
E. pallidoflavum (Henn. & E. Nyman) E.
Horak Vietnam LE 262934 OQ779469 MH190183 MH190155 [7], this
work
E. panniculus (Berk.) Sacc. Australia G1680
— MK278012 — [54]
E. percoelestinum O.V. Morozova, Noordel.,
Vila & Bulyonk. Spain LE 254390, holotype KF745927 KF745928 — [39]
J. Fungi 2023, 9, 621 6 of 24
E. peristerinum O.V. Morozova et T.H.G.
Pham Vietnam LE F-343653,
holotype OQ779466 OQ804522 OQ779188 this work
E. peristerinum Vietnam LE F-343649 OQ779468 — — this work
E. peristerinum Vietnam LE F-343650 OQ779467 OQ804524 OQ779186 this work
E. peristerinum Vietnam LE F-343652 — OQ804525 OQ779187 this work
E. peristerinum Vietnam LE F-343651 — OQ804523 OQ779189 this work
E. placidum (Fr.) Noordel. Sweden UPS:BOT:F-121714,
epitype, KC898394 KC898514 — [39]
E. plicatum (Largent) Blanco-Dios USA DAR: DLL10091 — — MG702627 [7]
E. prunuloides (Fr.) Quél. USA AFTOL-ID: 523 4765
TJB — — DQ457633 [55]
E. quadratum (Berk. & M.A. Curtis) E. Horak
[Inocephalus quadratus (Berk. & M.A. Curtis) T.J.
Baroni]
Russia: Far East LE254355 KC898452 KC898504 — [39]
E. quadratum USA: New York CORT: TJB 8214 — — MH190162 [7]
E. sericellum (Fr.) P. Kumm. [Alboleptonia
sericella (Fr.) Largent & R.G. Benedict] Russia: Caucasus LE 254362 KC898453 — — [39]
E. sericellum USA: California HSU: DLL9524 — — MG702617 [7]
E. sericellum Belgium MEN 200315 — GQ289190 — [5]
E. serrulatum (Fr.) Hesler Russia: Caucasus LE 254361 KC898447 KC898501 — [39]
E. sublaevisporum Vila, Noordel. & O.V.
Morozova Spain LIP JVG 1070823T,
holotype KC898436 KC898518 — [39]
E. tadungense O.V. Morozova et T.H.G. Pham Vietnam LE F-343680,
holotype OQ779470 — — this work
E. tadungense Vietnam LE F-343681 OQ779471 OQ804526 — this work
E. tibiiforme (Largent & Aime) Blanco-Dios
[Trichopilus tibiiformis Largent & Aime] Guyana MCA2426, holotypus
(BRG, LSUM) — — MG702645 [7]
E. tjallingiorum Noordel. var. tjallingiorum Sweden UPS:BOT:F-016378,
holotype KC898412 KC898509 — [39]
E. turbidum (Fr.) Quél. Slovakia MEN 200351 KC710060 GQ289201 — [5]
E. umbrophilum Noordel. & Hauskn. [Leptonia
umbrophila (Noordel. & Hauskn.) Largent]
Australia:
Queensland DLL9766 (BRI, CNS) — — MG702638 [7]
E. undatum (Gillet) M.M. Moser Russia: European
part LE 312417 MF476910 MF487801 — [42]
E. venustum Wölfel & F. Hampe Germany L, Wö E17/10,
holotype KC898355 KC898523 — [39]
E. vernum S. Lundell Russia: European
part LE 312538 OL338282 OL338537 OL405539 [38]
E. violaceozonatum
Noordel. & Liiv Estonia L 275, holotype KC898448 KC898502 — [39]
E. virescens (Sacc.)
E. Horak ex Courtec.
[Inocephalus virescens (Berk. & M.A. Curtis)
Largent & Abell-Davis]
MEL:2379813 MF977981 — — Genbank
E. virescens Guyana MCA 2479 — GU384622 MG702629 [7]
Three datasets were analysed: nrITS, tef1α, and nrLSU. DNA sequences were aligned
with the MAFFT v.7.110 web tool [56] using the G-INS-i option, and then manually
modified where necessary in MEGA X [49]. To determine the phylogenetic positions of
the studied collections, both datasets were analysed using Bayesian Analysis (BA). BA was
performed using MrBayes 3.2.1 [57], under a GTR model. The analyses were run with two
parallel searches: four chains for 5 million generations for ITS and LSU and for 1 million
generations for tef1α, starting with a random tree. The trees were sampled every 100
generations. To check for convergence of MCMC analyses and to obtain estimates of the
posterior distribution of parameter values, Tracer v1.7.2 was used [58]. The phylogenetic
J. Fungi 2023, 9, 621 7 of 24
trees were edited in Adobe Illustrator CS4. Posterior probability (PP) values ≥ 0.95 are
considered significant.
3. Results
3.1. Phylogenetic Analysis
The full nrITS dataset contained 50 sequences with 1059 characters (gaps included).
The/Entocybe clade was selected as outgroup due to its basal position in the Entoloma
phylogeny [6]. Besides our specimens, the tree includes 7 more sequences of the subgenus
Cubospora retrieved from the GenBank NCBI data base, 16 representatives of the subgenus
Leptonia, and 1–3 representatives of the other main subdivisions of the Entoloma s.l.
Since in the GenBank the ITS data are absent for many species of the subgenus
Cubospora, a tree was constructed for this subgenus based on the tef1α as well. For the
subgenus Leptonia, such information is insufficient for analysis. The full tef1α dataset
contained 26 sequences with 535 characters (gaps included). It included Entoloma
prunuloides as an outgroup due to its basal position in the Entoloma phylogeny [53,54], and
E. luteolamellatum, the type-species of the subgenus Cubospora. In addition to our
specimens, 11 more representatives of this subgenus and 1–3 representatives of the other
main subgenera of the Entoloma s.l. were included in the analysis.
The full nrLSU dataset contained 52 sequences with 781 characters (gaps included).
Clitopilus prunulus and Clitopilopsis hirneola were chosen as an outgroup because of their
basal position in the Entolomataceae phylogeny [5]. In addition to new species specimens,
8 representatives of the subgenus Cubospora and 15 sequences of the subgenus Leptonia, as
well as 1–3 sequences of the other main subgenera of the Entoloma s.l., were added to the
analysis.
The results of the phylogenetic analysis are presented in the Figure 1 (nrITS), Figure
2 (tef1α), and Figure 3 (nrLSU). Entoloma cycneum and E. peristerinum form highly
supported branches within the/Cubospora clade in all trees. E. tadungense and E. dichroides
clearly nest within the/Leptonia clade in the ITS tree. They are not represented on the tef1α
tree. Generally, in the LSU tree, the topology mostly is not or hardly resolved due to small
differences between the sequences in the subgenera. At the same time, the subgenus
Cubospora forms a highly supported clade in which two new species (E. cycneum and E.
peristerinum) nest.
Subgenus Leptonia does not form a separate clade in the LSU tree, but it is represented
by several small clades and singletons. However, it is clearly seen here that Entoloma
tadungense clusters together with E. percoelestinum and E. coelestinum with high support.
Entoloma dichroides distinctly nests in the/dichroum clade, but it differs from the known
Leptonia species.
J. Fungi 2023, 9, 621 8 of 24
Figure 1. Phylogenetic tree derived from Bayesian analysis, based on nrITS1-5.8S-ITS2 region data.
Posterior probability (PP > 0.95) values from the Bayesian analysis are added at the nodes. The scale
bar represents the number of nucleotide changes per site. The new species are in bold. HT—
holotype.
J. Fungi 2023, 9, 621 9 of 24
Figure 2. Phylogenetic tree derived from Bayesian analysis, based on tef1α data. Posterior
probability (PP > 0.95) values from the Bayesian analysis are added at the nodes. The scale bar
represents the number of nucleotide changes per site. The new species are in bold. HT—holotype.
J. Fungi 2023, 9, 621 10 of 24
Figure 3. Phylogenetic tree derived from Bayesian analysis, based on nrLSU data. Posterior
probability (PP > 0.95) values from the Bayesian analysis are added at the nodes. The scale bar
represents the number of nucleotide changes per site. The new species are in bold. HT—holotype.
J. Fungi 2023, 9, 621 11 of 24
3.2. Taxonomy
Entoloma subgenus Cubospora Karstedt, Capelari, Largent, T. J. Baroni &
Bergemann, in Phytotaxa 391(1): 20 (2019).
Entoloma cycneum O.V. Morozova & T.H.G. Pham, sp. nov. (Figure 4).
MycoBank: MB848527
Etymology. From “κύκνος” (Greek)—swan, due to colour similarity with white swan.
Holotype. Vietnam, Gia Lai Province, K’Bang District, Son Lang Commune, Kon Chu
Rang Nature Reserve, N 14.49436°, E 108.54428°, 1030 m a. s. l., on soil in middle-mountain
evergreen mixed forest with a predominance of Podocarpaceae (Dacrydium elatum,
Dacrycarpus imbricatus), Magnoliaceae, Burseraceae (Canarium), Myrtaceae (Syzygium), 30
May 2016, O.V. Morozova (LE F-343654). Isotype in VRTC (299VN16).
Diagnosis. Entoloma cycneum is characterized by initially white or whitish basidiomata
with mainly smooth glabrous with thin pubescence near the margin, hygrophanous
pileus, longitudinally fibrillose or fibrillose-scaly stipe, cuboid spores, and long, more or
less cylindrical cheilocystidia of serrulatum-type.
Basidiomata small to medium-sized. Pileus 10‒25 mm diam., firstly hemispherical,
then convex, plano-convex with small papilla, with a slightly involute then straight
crenulate margin, hygrophanous, smooth, mostly glabrous, but covered with thin fibrils
on the pileus margin, in wet conditions translucently striate almost up to the centre, white
to cream (3A1–2; 4A1–2), yellowish in places of damage, lighter towards margin, slightly
da rker in radial hygrop hanous stripe s, then dries u p to whi te. Lamellae moderately distant,
adnexed, adnate-emarginate, ventricose, initially white, then cream, pale pink, with white
denticulate edge. Stipe 30–60 × 1.5–3 mm, cylindrical, fistulous, white, longitudinally
fibrillose or fibrillose-scaly, at least at the top, white tomentose at base. Context white or
hyaline. Smell indistinct, taste not reported.
Basidiospores (7.2–)8.5–9.0(–9.7) × (7.2–)8.0–8.5(–8.7) µm, Q = 1.0–1.2, isodiametrical or
subisodiametrical, cuboid, rarely with 5 angles in side-view. Basidia 37–50 × 11–12.5 µm,
4-spored, clavate, constricted in the middle part, clamped. Cheilocystidia 95–160 × 7.5–9
µm, of serrulatum-type, flexuous-cylindrical, sometimes capitate or narrowly clavate,
septate, not pigmented, with granular content, forming sterile lamellae edge.
Hymenophoral trama regular, made up of cylindrical or inflated hyphae 4–10 µm wide.
Pileipellis a cutis of cylindrical hyphae up to 10 µm broad, some hyphae are rising.
Pileitrama consists of cylindrical hyphae 4–6 µm broad, intertwined by oleiferous hyphae
with brilliant content. Stipitipellis of cylindrical hyphae up to 7 µm broad. Caulocystidia in
bundles of rising hairs, cylindrical or slightly broadened towards apex, up to 250 µm
length and 12 µm broad. Clamp-connections present in all tissue. Brilliant granules
abundant.
Habitat and distribution—In small groups on soil in middle-mountain evergreen
mixed forest. Known from Vietnam.
Additional specimens examined. Vietnam, Gia Lai Province, K’Bang District, Son Lang
Commune, Kon Chu Rang Nature Reserve, N 14.49439°, E 108.54591°, 990 m a. s. l., on soil
in middle-mountain evergreen mixed forest with a predominance of Podocarpaceae
(Dacrydium elatum, Dacrycarpus imbricatus), Magnoliaceae, Burseraceae (Canarium),
Myrtaceae (Syzygium), 27 May 2016, O.V. Morozova (LE F-343656 (1687_243VN16); ibid.,
28 May 2016, O.V. Morozova (LE F-343655 (1689_255VN16); LE F-343657 (1690_256VN16);
Dak Nong Province, Dak Plao District, south-east macroslope of Mt. Ta Dung, trail along
the stream valley in the saddle between M’neun Tchirke and the eastern spur of Ta Dung,
N 11.87172°, E 108.08253°, 1310 m a. s. l., on soil in middle-mountain evergreen mixed
forest with a participation of Fagaceae and Lauraceae (Litsea glutinosa), 3 June 2022, T.H.G.
Pham (LE F-343658).
J. Fungi 2023, 9, 621 12 of 24
Figure 4. Entoloma cycneum: (a,b). basidiocarps; (c). basidiospores; (d). cheilocystidia; (e). pileipellis;
(f). caulocystidia (a,d–f, from LE F-343654, holotype; b—from LE F-343655). Scale bars (a,b) 1 cm,
(c–f) 10 µm.
Entoloma peristerinum O.V. Morozova & T.H.G. Pham, sp. nov. (Figure 5).
MycoBank: MB848528
Etymology. From “περιστερά” (Greek)—dove, due to colour similarity to white dove.
Holotype. Vietnam, Gia Lai Province, K’Bang District, Son Lang Commune, Kon Chu
Rang Nature Reserve, N 14.49667°, E 108.56106°, 980 m a. s. l., on soil in middle-mountain
evergreen mixed forest with a predominance of Podocarpaceae (Dacrydium elatum,
J. Fungi 2023, 9, 621 13 of 24
Dacrycarpus imbricatus), Magnoliaceae, Burseraceae (Canarium), Myrtaceae (Syzygium), 28
May 2016, O.V. Morozova (LE F-343653). Isotype in VRTC (276VN16).
Diagnosis. Entoloma peristerinum is characterized by initially cream, pale beige to beige
or greyish-beige becoming white basidiomata, with smooth, glabrous, hygrophanous
pileus, longitudinally fibrillose or fibrillose-scaly stipe, cuboid spores and cheilocystidia
mostly narrowly clavate or tapering towards the apex—of porphyrogriseum-type.
Basidiomata small to medium-sized. Pileus 10‒30 mm diam., firstly conical, then
broadly conical with distinct acute papilla, with deflexed then straight or undulating,
slightly crenulate margin, hygrophanous, smooth, glabrous, in wet condition
translucently striate almost up to the centre, cream, pale beige to beige or greyish-beige
(4A2; 4B2–3; 5B2–3), lighter towards margin, with dark radial hygrophanous stripes and
yellowish papilla, then dries up by white radial strokes, finally becomes completely white,
silky. Lamellae moderately distant, adnate-emarginate, ventricose, cream, pale pink, with
minutely serrate whitish edge. Stipe 30–70 × 1.5–3 mm, cylindrical, fistulous, white,
longitudinally fibrillose or fibrillose-scaly, at least at the top, white tomentose at base.
Context white. Smell indistinct, taste not reported.
Basidiospores (7.4–)8–9.5(–9.8) × (6.5–)7–8(–9.2) µm, Q = 1.0–1.2(–1.3), isodiametrical or
subisidiametrical, cuboid, rarely with 5 angles in side-view. Basidia 47–60 × 11–14.5 µm, 4-
spored, clavate, constricted in the middle part, clamped. Cheilocystidia 75–215 × 12–15 µm,
of porphyrogriseum-type [8], cylindrical, narrowly clavate or tapering towards the apex,
with granulose content, usually forming sterile lamellae edge; in young basidiomata
lamellae edge can be heterogeneous with rare cheilocystidia. Hymenophoral trama regular,
made up of cylindrical or inflated hyphae 3–8 µm wide. Pileipellis a cutis of cylindrical
hyphae up to 10 µm broad. Pileitrama consists of cylindrical hyphae 4–6 µm broad,
intertwined by oleiferous hyphae with brilliant content. Clamp-connections present in all
tissue. Brilliant granules abundant.
Habitat and distribution—In small groups on soil in middle-mountain evergreen
mixed forest. Known from Vietnam.
Additional specimens examined. Vietnam, Gia Lai Province, K’Bang District, Son Lang
Commune, Kon Chu Rang Nature Reserve, N 14.49439°, E 108.54591°, 990 m a. s. l., on soil
in middle-mountain evergreen mixed forest with a predominance of Podocarpaceae
(Dacrydium elatum, Dacrycarpus imbricatus), Magnoliaceae, Burseraceae (Canarium),
Myrtaceae (Syzygium), 28 May 2016, O.V. Morozova (LE F-343651 (1688_254VN16)); ibid.,
N 14.49667°, E 108.56106°, 980 m a. s. l., 28 May 2016, O.V. Morozova (LE F-343650
(1691_276bVN16)); ibid., N 14.49436°, E 108.54428°, 1030 m a. s. l., 30 May 2016, O.V.
Morozova, LE F-343652 (1692_300VN16), LE F-343649 (1696_312VN16)).
J. Fungi 2023, 9, 621 14 of 24
Figure 5. Entoloma peristerinum: (a,b). basidiocarps; (c). basidiospores; (d). cheilocystidia; (e).
basidium; (f). pileipellis; (g). caulocystidia (a,c–g, from LE F-343653, holotype; b—from LE F-
343652). Scale bars (a,b) 1 cm, (c–e) 10 µm, (f,g) 20 µm.
Notes—Entoloma cycneum and E. peristerinum are morphologically very similar
species characterized by white or whitish with yellowish or beige tinged basidiomata with
mainly smooth, glabrous, and hygrophanous pileus, longitudinally fibrillose or fibrillose-
scaly white stipe, cuboid spores and more or less cylindrical cheilocystidia, arising from
hymenophoral trama. Entoloma peristerinum posseses initially more coloured beige conical
pileus, discolouring to white with age and drying. The pileus of E. cycneum is initially
J. Fungi 2023, 9, 621 15 of 24
white, hemisphaerical to convex, usually with thin pubescence near the margin. E.
peristerinum differs from E. cycneum also by more differentiated porphyrogriseum-type [8]
cheilocystidia, vs. serrulatum-type in E. cycneum. Yellow tinge can present in old or
damaged basidiomata of both species.
Phylogenetically both new species are also close to each other but rather distant from
other known species in the subgenus Cubospora [7]. Entoloma cervinum (Karstedt &
Capelari) Blanco-Dios and E. acutipallidum E. Horak & Cheype from the South America
are the closest species according to the phylogenetical analysis. Morphologically they
differ from our species by darker, distinctly coloured pileus [28].
Several species with cuboid spores and predominantly white or whitish basidiomata
have been described at different times from different regions of the Earth.
The pileus of Entoloma alboumbonatum Hesler from North America is darker coloured,
and only umbo is white. This species is characterized by the clavate or capitate
cheilocystidia, presence of the pleurocystidia, and absence of the clamp connections [31].
E. albidoquadratum Manim. & Noordel., described from India, is characterized by non
hygrophaneous, non-translucently striate pileus, presence of pleurocystidia, and large
spores [20]. E. minutoalbum E. Horak is a species of Southern Hemisphere, a common
fungus of the subantarctic Nothofagus forests of Tierra del Fuego and New Zealand.
Morphologically it differs from new species by small basidiomata (with the pileus less
than 10 mm diam.), smaller spores, and absence of the cheilocystidia [13]. E. albogracile E.
Horak is also species from the Southern Hemisphere—Papua New Guinea. It is
characterized by the small size, pileus covered with minute scales and fibrils, and by the
absence of clamp-connections [13]. E. laccaroides T.H. Li, E. Horak & Xiao Lan He is
recognized by the umbilicate pileus, and numerous conspicuous broadly fusoid to
utriform pleurocystidia [12].
Due to the discolouration new species would be compared with cuboid-spored light-
yellow species. Entoloma dennisii from Trinidad is a rather robust species with pileus up
to 5 cm broad, deep to pale yellow colour, and small spores. E. pallidoflavum differs by the
predominance of light-yellow colour in the pileus, yellow content of the oleiferous
hyphae, and smaller spores [13].
Entoloma subgen. Leptonia (Fr.) Noordel., in Persoonia 11: 146 (1981), emend. O. V.
Morozova, Noordel. & Vila (2014); emend. Noordeloos et al. (2022).
Entoloma tadungense O.V. Morozova & T.H.G. Pham, sp. nov. (Figure 6).
MycoBank: MB848530
Etymology. Named after the type-locality—Ta Dung National Park.
Holotype. Dak Nong Province, Dak Glong District, Ta Dung National Park, south-
eastern macroslope of the ridge of the Ta Dung Mt, south-eastern slope of the Ta Dung Mt,
TK 1805, N 11.86780°, E 108.11692°, 1240 m a.s.l., on soil in evergreen broad-leaved forest
with Lithocarpus spp., Quercus sp., Schima sp., Acer flabellatum, Dilleniaceae, Myristicaceae,
11 Oct. 2022, T.H.G. Pham, O.V. Morozova (LE F-343680). Isotype in VRTC (87VN22).
Diagnosis. Entoloma tadungense is distinguished among the other Leptonia species by
the tiny dark blue basidiomata with discolouring to lilac of the stipe, and by the small
spores with rather pronounced angles.
Basidiomata small-sized, mycenoid. Pileus 3–10 mm broad, conical, or hemispherical
with umbo, becoming almost applanate, not hygrophanous, not translucently striate, with
deflexed then straight margin, radially fibrillose, squamulose at centre, firstly uniformly
dark blue, blackish blue (19F7–8; 20E6–8; 21F7–8), glabrescent, radially cracking showing
the white background, and pallescent (up to 20C5–6; D5–6) with edge. Lamellae
moderately distant, adnate emarginate, ventricose, white, becoming pinkish, with entire
concolourous edge. Stipe 1.5–30 × 0.5–2 mm, cylindrical or broadened towards to base,
fistulous, initially distinctly longitudinally fibrillose, glabrescent with age up to almost
smooth, especially in the apex, firstly concolourous with pileus—dark blue, blackish blue
(19F7–8; 20E6–8), then discoloured up to lilac, whitely tomentose at base. Context thin,
concolourous with the surface. Smell indistinct, taste not reported.
J. Fungi 2023, 9, 621 16 of 24
Basidiospores (6.0–)6.8–6.9(–8.3) × (4.2–)4.8–4.9(–5.7) µm, Q = 1.3–1.4(–1.6),
heterodiametrical, with 6–7 relatively sharp angles in side-view. Basidia 19.5–24 × 6.6–8.3
µm, 4-spored, narrowly clavate to subcylindrical, clamped. Lamellae edge fertile or
heterogeneous. Cheilocystidia 24–67 × 9–17 µm, of poliopus-type [8], clavate or lageniform,
more frequent near the pileus margin. Hymenophoral trama regular, made up of narrow,
cylindrical hyphae up to 3 µm wide. Pileipellis a cutis with a transition to a
plagiotrichoderm and a trichoderm of cylindrical to slightly inflated hyphae 10–20 µm
wide with blue intracellular pigment. Stipitipellis a cutis with a transition to a
plagiotrichoderm with rising hyphae forming hairs up to 5 µm wide. Clamp-connections
present, but rare. Brilliant granules absent.
Habitat and distribution—In small groups on soil in middle-mountain evergreen
mixed forest. Known from Vietnam.
Additional specimens examined. Dak Nong Province, Dak Glong District, Ta Dung
National Park, south-eastern macroslope of the ridge of the Ta Dung Mt, south-eastern
slope of the Ta Dung Mt, TK 1805, N11.86780, E 108.11692, 1240 m a.s.l., on soil in
evergreen broad-leaved forest with Lithocarpus spp., Quercus sp., Schima sp., Dilleniaceae,
Myristicaceae, Acer flabellatum, 11 Oct. 2022, T.H.G. Pham, O.V. Morozova, LE F-343681,
duplicate in VRTC (88VN22); ibid., 15 October 2022, LE F-343683, duplicate in VRTC
(138VN22), LE F-343684, duplicate in VRTC (139VN22).
Notes—Entoloma tadungense is a species of the subgenus Leptonia due to presence of
clamp connections, absence of brilliant granules, longitudinally fibrillose stipe surface and
plagiotrichoderm to trichoderm pileipellis. It resembles Entoloma percoelestinum O.V.
Morozova, Noordel., Vila & Bulyonk. by the small-sized dark blue mycenoid basidiomata
[39]. Microscopically the absence of cheilocystidia and small spores also make them
similar. However, E. tadungense can be recognized by smaller spores with pronounced
angles, as well as the lilac discolouration of the stipe. Molecular data support their
differences (p-distance from the closest species E. percoelestinum (ITS1-5.8S-ITS2 region)—
4.9%). The similar non tropical species with dark blue colour and small size of the
basidiomata distinguish from the new species in the following: E. coelestinum (Fr.)
Hesler—by the smooth stipe and slightly larger spores, E. lepidissimum (Svrček) Noordel.
and E. venustum—by the coloured lamellae, distinctly larger spores, and presence of the
cheilocystidia, E. chytrophilum possesses large nodulose spores [39]. The American species
E. subcoelestinum (Largent) Blanco-Dios is characterized by the initially coloured lamellae,
moniliform cells in the pileipellis, and larger indistinctly angular spores [32].
J. Fungi 2023, 9, 621 17 of 24
Figure 6. Entoloma tadungense: (a–d). basidiocarps; (e). basidiospores; (f). cheilocystidia and
basidium; (g). pileipellis; (h). caulocystidia (a–f,h from LE F-343680, holotype; g—from LE F-
343681). Scale bars (a–d)—1 cm; (e,f,h)—10 µm; (g)—20 µm.
Entoloma dichroides O.V. Morozova & T.H.G. Pham, sp. nov. (Figure 7).
MycoBank MB 848531
Etymology. Named after its similarity to Entoloma dichroum.
Holotype. Dak Nong Province, Dak Glong District, Ta Dung National Park, northwest
of the Ta Dung Mt, TK 1781, N11.923056°, E 108.00194°, 1000 m a.s.l., on soil in evergreen
J. Fungi 2023, 9, 621 18 of 24
broad-leaved forest with Parashorea chinensis, Rhodoleia championii, Fagaceae, Lauraceae,
Hypericaceae, 1 June 2022, Pham T.H.G., LE F-343682. Isotype in VRTC (HG09).
Diagnosis. Entoloma dichroides is a species of subgenus Leptonia, characterized by the
dark blue basidiomata with squamulose surface of the pileus and stipe, initially white
lamellae, absence of the cheilocystidia, and spores with 5(–6) pronounced angles and
elongated apiculus.
Basidiomata medium-sized, mycenoid. Pileus 40 mm diam, conical to conico-convex,
with deflexed then straight margin, not hygrophanous, not translucently striate, dark blue
(20F6–8), entirely fibrillose-squamulose with prominent dense squamules on whitish
background. Lamellae narrowly adnexed to almost free, ventricose, moderately distant,
initially white becoming yellowish-pink, brownish-pink when, with an entire
concolourous edge. Stipe 60 × 3–4 mm, cylindrical, slightly broadened at base, hollow, dark
blue, concolourous with the pileus, entirely fibrillose-squamulose with dark blue fibrils
(20F6–8) on whitish background. Context white. Smell weak, taste not reported.
Basidiospores 9.5–12.5 × 6.5–8.5 µm, on average 10.8 × 7.2 µm, Q = 1.4–1.6(1.7), Qav = 1.5,
5(–6)-angled, heterodiametric with pronounced angles and elongated apiculus. Basidia 31–
41 × 10–13 µm, 4-spored, clamped. Lamella edge fertile. Cheilocystidia absent. Hymenophoral
trama regular, made up of narrow cylindrical hyphae 2–6 µm wide. Pileipellis a trichoderm
of cylindrical entangled 6–10 m wide hyphae with fusiform or irregularly shaped terminal
elements, 90–200 × 13–19 µm, with dark blue intracellular pigment. Stipitipellis an
entangled trichoderm of cylindrical hyphae, 6–12 m wide with cylindrical to lageniform,
terminal elements, forming squamules, 4–12 µm wide and up to the 200 long, with dark
blue intracellular pigment. Clamp-connections abundant. Brilliant granules absent.
Habitat and distribution—Solitary in middle-mountain evergreen mixed forest.
Known from Vietnam.
Notes—Entoloma dichroides is similar to European E. dichroum and Australian E.
panniculus due to dark blue basidiomata with squamulose surface of the pileus and stipe,
initially white lamellae, and spores with pronounced angles [26]. New species differs from
them by darker basidiomata with conical pileus, basidiospores with elongated apiculus,
and absence of the cheilocystidia. The Eastern species E. eugenei Noordel. & O.V.
Morozova is also close. It is recognized by the more robust basidiomata and presence of
the cheilocystidia [21]. The p-distance (ITS1-5.8S-ITS2 region) of the new species from E.
dichroum—6.9%, from E. eugenei—10.4%.
J. Fungi 2023, 9, 621 19 of 24
Figure 7. Entoloma dichroides: (a–d). basidiocarps; (e,f). basidiospores; (g). basidiola and basidium;
(h). stipitipellis; (i). pileipellis (all from LE F-343682). Scale bars (a–d)—1 cm; (e–i)—10 µm.
4. Discussion
The genus Entoloma is the second largest genus in the order Agaricales with over 1000
species worldwide [59]. However, studies devoted to it in Vietnam have not yet been
carried out enough.
The first most important contribution to the study of Vietnamese mycobiota was
made by N. Patouillard, who described many new species from Northern and Central
J. Fungi 2023, 9, 621 20 of 24
Vietnam based on collections by V. Demange, L. Duport, P. A. Eberhardt, E. Poilane, and
others. First records of the genus Entoloma (Rhodophyllus clypeatus, Rh. sericeus) were also
published by him [60,61], including originally described from Vietnam Rh. submurinus
[61]. Heim and Malençon [62] published information on E. madidum. The first checklist
summarizing all the data on the species composition of fungi and slime-molds in Vietnam
was published in 1998 and included 829 species, of which only 6 belonged to the genus
Entoloma [63]. The same species are presented in the list of plant species of Vietnam [64].
In 2003, Le Ba Dung listed 300 species of macromycetes for the Central Highlands, with
only two Entoloma species: E. lividum and E. madidum [65]. Later, Ngô Anh and Nguyễn
Thị Kim Cúc recorded two more species of the genus in Thua Thien Hue Province—E.
prunuloides in Bach Ma National Park [66] and E. abortivum in Phong Dien Nature Reserve
[67]. The data on Vietnamese mycobiota have been summarized in a monography
published in 2011–2012, which provides information on the ecology and distribution of
more than 900 species of macromycetes [68,69] with only 3 Entoloma species.
More intensive studies of the genus Entoloma in Vietnam began in the frame of the
work of the Joint Vietnam–Russia Tropical Science and Technology Research Centre in
collaboration with the Komarov Botanical Institute RAS. In 2012, Morozova et al. [11]
reported 12 species of the genus Entoloma (Agaricales, Basidiomycota) for the Bidoup—
Nui Ba (Lam Dong Province) and Cat Tien (Dong Nai Province) National Parks, of which
11 species were recorded for Vietnam for the first time, and a new species for science, E.
myriadophyllum O.V. Morozova, was described.
The above information was based only on morphological data. In connection with
the revision of the system of the genus Entoloma [8], these data are not entirely credible
and require confirmation. In cases where herbarium material is absent or old and
destroyed, this is not possible. However, sequences were later obtained for some of these
records. Some of the identifications were revised, and some of these findings were
described as species new to science—E. daphnis and E. bidupense [4,42]. Our specimens
with cuboid spores were used in the work of Karstedt and colleagues [7], where additional
data on tef1α, mtSSU, rpb2, and LSU markers were obtained for them.
Further descriptions of new species have already been confirmed by molecular data.
Thus, Entoloma flavovelutinum and E. porphyroleucum from the Bu Gia Map National Park
(Binh Phuoc Province) [2,70], E. nigrovelutinum from Chu Yang Sin National Park (Dak Lak
Province) [71], and E. atricolor [72], E. arion, E. argus, E. daphnis, E. icarus [4], E. kovalenkoi
[3], from the Kon Сhu Rang Nature Reserve were described.
As a result, the data on 29 species of Entoloma have been published for Vietnam so
far. Only for 14 of them the molecular data have been published and submied into the
GenBank. The resulting list is presented here.
List of the species of the genus Entoloma mentioned in the publications for Vietnam,
1910–2022 (species, supported for the molecular data are marked by the asterisk (*)):
Entoloma abortivum (Berk. & M.A. Curtis) Donk [67]
*E. altissimum (Massee) E. Horak [11]
*E. argus O.V. Morozova, E.S. Popov, A.V. Alexandrova & Noordel. [4]
*E. arion O.V. Morozova, E.S. Popov, T.H.G. Pham & Noordel. [4]
*E. atricolor O.V. Morozova, Noordel., E.S. Popov & A.V. Alexandrova [42,72]
*E. bidupense O.V. Morozova & E.S. Popov [11] (as E. violaceoserrulatum Noordel.), [42]
*E. carneum Z.S. Bi [7,11]
E. chalybeum (Fr.) Noordel. var. lazulinum (Fr.) Noordel. [11], (non est)
E. clypeatum (L.) P. Kumm. [60] (as Rh. clypeatus Fr.), [63,64,68]
*E. daphnis O.V. Morozova, E.S. Popov, T.H.G. Pham & Noordel. [4,11] (as E. nubilum
Manim., Leelav. et Noordel.)
E. depluens (Batsch) Hesler [63,64]
*E. flavovelutinum O.V. Morozova, E.S. Popov, A.V. Alexandrova & Xiao L. He [70]
*E. icarus O.V. Morozova, E.S. Popov & Noordel. [4]
*E. inocephalum (Romagn.) Dennis [7,11]
J. Fungi 2023, 9, 621 21 of 24
*E. kovalenkoi O.V. Morozova, E.S. Popov & A.V. Alexandrova [3]
E. sinuatum (Bull.) P. Kumm. [65,68] (both as E. lividum (Bull.) Quél.)
E. longistriatum (Peck) Noordel. [11]
E. madidum Gillet [62] (as Rhodophyllus madidus (Fr.) Quél.), [63–65]
*E. myriadophyllum O. V. Morozova [11]
*E. nigrovelutinum O.V. Morozova & A.V. Alexandrova [71]
*E. pallidoflavum (Henn. & E. Nyman) E. Horak [7,11]
E. cf. platyphylloides (Romagn.) Largent [73]
E. poliopus (Romagn.) Noordel. var. alpigenum (J. Favre) Bon [11], (non est)
*E. porphyroleucum O.V. Morozova, Noordel. & Dima [2]
E. prunuloides (Fr.) Quél. [63,64,66]
*E. quadratum (Berk. et M. A. Curtis) E. Horak [11]
E. sericeum (Bull.) Quél. [61] (as Rh. sericeus (Fr.) Quél.), [63,64]
E. submurinum (Pat.) E. Horak [61] (as Rh. submurinus Pat.), [63,64]
E. sulcatum (T.J. Baroni et Lodge) Noordel. et Co-David [11]
Four more species are described in the presented article. For E. pallidoflavum, the
information on ITS sequence is published here.
In conclusion, it is important to note that the area of tropical forests has been
drastically reduced in recent years, including in Vietnam. Along with them, the species
confined to them disappear, often without even being assigned a name. Nature reserves
and national parks serve to save biologically valuable forest areas. However, even here
the species are vulnerable. It sometimes happens that type localities are destroyed during
road reconstructions (e.g., E. atricolor, E. arion). It remains to be hoped that they will
continue to be found in the adjacent forests. In central Vietnam, the greatest diversity of
fungi of the genus Entoloma was observed in middle-mountain evergreen broad-leaved
forests. They require special aention, study, and careful treatment.
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 the
species inhabiting them.
Author Contributions: Conceptualization, collecting, formal analysis, investigation, writing—
original draft preparation—O.M. and T.H.G.P. O.M. and T.H.G.P. contributed equally and should
be considered as co-first authors. All authors have read and agreed to the published version of the
manuscript.
Funding: The field work was funded by Joint Vietnam-Russia Tropical Science and Technology
Research Centre (project Е1.5 No3). The work with collections was provided in the framework of an
institutional research project of the Komarov Botanical Institute RAS (122011900032-7).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: The DNA sequence data obtained from this study have been deposited
in GenBank NCBI (hps://www.ncbi.nlm.nih.gov/genbank/, accessed 13 April 2023).
Acknowledgments: The authors are grateful to the management and staff of the Kon Chu Rang
Nature Reserve and Ta Dung National Park for their great help in the field work and for permission
to collect in the forests. Many thanks to E.S. Popov (Komarov Botanical Institute RAS) for the help
in the field work and with the literature.
Conflicts of Interest: The authors declare no conflict of interest.
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