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Phytotaxa 181 (3): 163–170
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Copyright © 2014 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Genevieve Gates: 20 Jun. 2014; published: 3 Oct. 2014
http://dx.doi.org/10.11646/phytotaxa.181.3.4
163
A new angiocarpous Lactarius species from Thailand
ANNEMIEKE VERBEKEN1*, FELIX HAMPE1, KOMSIT WISSITRASSAMEEWONG1,2,3, KEVIN HYDE2,3,
URSULA EBERHARDT1,4 & JORINDE NUYTINCK5
1 Ghent University, Dpt. Biology, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium
2Institute of Excellence in Fungal Research, Mae Fah Luang University, 333 Moo 1, Thasud sub-district, Muang district, Chiang Rai
57100, Thailand
3School of Science, Mae Fah Luang University, 333 Moo 1, Thasud sub-district, Muang district, Chiang Rai 57100, Thailand
4 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany
5 Naturalis Biodiversity Center, Section National Herbarium of the Netherlands, P.O. Box 9517, 2300RA Leiden, The Netherlands
* corresponding author: mieke.verbeken@ugent.be
Abstract
Lactarius bisporus sp. nov. is described from primary tropical forest in Thailand. Morphological characters and DNA se-
quence data are given. Comparisons with the closely related angiocarpous taxon Lactarius pomiolens are provided.
Key Words: basidiomycetes, ectomycorrhiza, taxonomy, truffle-like Russulales
Introduction
As in numerous other agaricomycete groups, it is now accepted that also in the Russulales angiocarpous species have
evolved many times from gymnocarpous species, and that the shape of the basidiocarps has long been overestimated
as a phylogenetic character (Miller et al. 2001, Desjardin 2003, Eberhardt & Verbeken 2004, Nuytinck et al. 2004,
Verbeken et al. 2014). After the recent division of the milkcaps in three genera: Multifurca Buyck & V. Hofst. (Buyck
et al. 2008, 2010), Lactarius Pers., and Lactifluus (Pers.) Roussel (Buyck et al. 2008, 2010, Norvell 2011, Barrie
2011), all truffle-like milkcap species known so far seem to belong to the genus Lactarius. Before the inclusion of
angiocarpous Russulales in agaricoid genera was accepted, a number of genera were erected to include sequestrate
species. Milk-exuding species were often, but not exclusively, described in Arcangeliella Cavara or Zelleromyces
Singer & A.H. Sm. (Miller et al. 2001, Eberhardt & Verbeken 2004, Nuytinck et al. 2004).
The angiocarpous habit evolved in the three main subgenera: L. subg. Plinthogalus (Burl.) Hesler & A.H. Sm., L.
subg. Russularia (Fr. ex Burl.) Kauffman and L. subg. Piperites (Fr. ex J. Kickx f.) Kauffman (Verbeken et al. 2014).
Angiocarpous Russulales are mainly known from North America and Australia, but also occur in the tropics where
their diversity is probably underestimated (Eberhardt & Verbeken 2004, Verbeken et al. 2014). A recent expedition in
Northern Thailand revealed another new truffle-like milkcap, which is proposed here as Lactarius bisporus sp. nov.
Material and Methods
Morphological study
Macroscopic characters are all based on fresh material. Microscopic features were studied from dried material
mainly in Congo Red in L4 (Clémençon 1973). Spore ornamentation is described and illustrated as observed in Melzers
reagent. For details on terminology we refer to Verbeken (1998) and Verbeken & Walleyn (2010). Line-drawings
were made by A. Verbeken, with the aid of a drawing tube at original magnifications: 6000 × for spores, 1000 × for
individual elements and sections. Basidia length excludes sterigmata length. Spores were measured in side view in
Melzer’s reagent, excluding the ornamentation, and measurements are given as {(MIN) [AV-2×SD]–AV–[AV+2×SD]
VERBEKEN ET AL.
164 Phytotaxa 181 (3) © 2014 Magnolia Press
(MAX)}length × {(MIN) [AV-2×SD]–AV–[AV+2×SD] (MAX)}width in which AV = mean value for the measured collection
and SD = standard deviation. Q corresponds to spore “length/width ratio” and is given as (MINQa) Qa–Qb (MAXQb),
where Qa and Qb are the lowest and the highest mean ratio for the measured specimens, respectively. Colour pictures
of this species will be available at http://www2.muse.it/russulales-news/.
DNA extraction, PCR amplification and sequencing
DNA was extracted from fresh material stored in 2×CTAB buffer, using the methods described by Nuytinck &
Verbeken (2003) with slight modifications (Van de Putte et al. 2010). The internal transcribed spacer region of the
nuclear ribosomal DNA (ITS) was amplified and sequenced using primers ITS1-F and ITS4 (White et al. 1990, Gardes
& Bruns 1993). Sequencing (using the same primers) was conducted by MACROGEN (Amsterdam, The Netherlands).
Sequences were assembled and edited with the software SequencherTM v5.0 (GeneCodes Corporation, Ann Arbor,
Michigan, U.S.A.).
Alignment and phylogenetic analyses
Table 1 shows an overview of all specimens and sequences used in the phylogenetic analyses, including GenBank
accession numbers. Alignments were constructed with PRANK (Löytynoja & Goldman 2008). The alignment was
manually refined in BioEdit v7.0.9.0 (Hall 1999) and made available in TreeBASE (http://purl.org/phylo/treebase/
phylows/study/TB2:S15099). Maximum Likelihood (ML) analyses (Fig. 1) were performed in RAxML v7.0.3 (Stamatakis
2006), combining a ML search with the Rapid Bootstrapping algorithm for 1000 replicates and using the GTRGAMMA
model.
TABLE 1. Specimens and GenBank accession numbers of DNA sequences used in the molecular analyses.
Species Voucher collection Origin ITS accession no.
Arcangeliella camphorata USA EU644701
A. crassa USA AY558740
A. sp. Australia, Tasmania JF960610
Lactarius acris EU014 (UPS) Germany DQ421988
L. akahatsu AV2004-141 (GENT) Thailand KF133269
L. albocarneus AV98-080 (GENT) France KF241545
L. alboscrobiculatus LTH175 (CMU, SFSU, GENT) Thailand EF141538
L. angiocarpus DA00-448 (GENT) Zambia AY606942
L. atroviridis AV05-306 (GENT) USA KF133270
L. auriolla RW1601 (GENT) Sweden KF133257
L. azonites AV00-124 (GENT) Belgium KF241540
L. baliophaeus AV05-155 (GENT) Malawi GU258277
L. bisporus spec. nov. FH 12-160 (type, GENT) Thailand KF856293
L. borzianus Switzerland AF373599
L. camphoratus UE04.09.2004 (UPS) Sweden DQ422009
L. chichuensis Wang1236 (HKAS) China KF241541
L. chromospermus AV99-174 (GENT) Zimbabwe KF133260
L. chrysorrheus UE04.10.2002-8 (UPS) Italy KF133261
L. citriolens UE20.09.2004-03 (UPS) Sweden DQ422003
L. controversus AV00-117 (GENT) Italy KF241544
L. crassiusculus LTH369 (GENT) Thailand EF560684
L. cyanescens DS06-058 (GENT) Malaysia GU258278
L. cyathuliformis UE04.09.2004-2 (UPS) Sweden KF133266
L. deliciosus JN2001-046 (GENT) Slovakia KF133272
L. echinellus AV07-169 (GENT) Sri Lanka KF133287
L. echinus AV07-168 (type, GENT) Sri Lanka KF133273
L. falcatus KVP08-038 (type, GENT) Thailand KF133274
......continued on the next page
LACTARIUS FROM THAILAND Phytotaxa 181 (3) © 2014 Magnolia Press 165
TABLE 1. (Continued)
Species Voucher collection Origin ITS accession no.
L. flexuosus UE06.09.2002-1 (UPS) Sweden DQ421992
L. formosus LTH382 (type, CMU, SFSU, GENT) Thailand EF141549
L. fuliginosus MTB97-24 (GENT) Sweden JQ446111
L. helvus UE08.09.2004-1 (UPS) Sweden KF133263
L. hispidulus AB152 (GENT) Guinea KF133258
L. kabansus AV99-205 (GENT) Zimbabwe KF133259
L. lignyotus UE06.09.2003-5 (UPS) Sweden DQ421993
L. luridus OB11-011 (GENT) Belgium KF241547
L. mairei AV00-118 (GENT) Italy AY336950
L. mammosus UE09.09.2004-5 (UPS) Sweden KF133265
L. montoyae KD1065 (type, BSHC) India EF560673
L. necator AV04-231 (GENT) France KF133276
L. peckii JN2004-020 (GENT) USA KF133277
L. pomiolens AV07-159 (type, GENT) Sri Lanka KF133282
L. pubescens AV96-931 (GENT) Norway AY336958
L. quieticolor UE10.09.2004-1 (UPS) Sweden DQ422002
L. quietus UE16.09.2004 (UPS) Sweden KF133264
L. romagnesii UE29.09.2002-6 (UPS) France DQ421989
L. rubriviridis DED7312 (SFSU) USA EF685088
L. rufus JN2002-008 (GENT) Norway KF241543
L. saturnisporus DS07-490 (GENT) Sri Lanka KF133285
L. shoreae AV07-164 (type, GENT) Sri Lanka KF133278
L. sphagneti PL2805 (pers. herb. P. Leonard) UK KF133268
L. spinosulus AT2003068 (UPS) Sweden KF133262
L. stephensii RW2930 (GENT) Belgium AY331012
L. subdulcis JV2006-024 (GENT) Belgium KF133279
L. subplinthogalus AV04-219 (GENT) USA KF241539
L. subsericatus UE11.10.2004-8 (UPS) Sweden DQ422011
L. tenellus DKA3598 (BR) Benin KF133280
L. thyinos A.Voitk23-08-2004 (GENT) Canada KF133271
L. torminosus RW3183 (GENT) Czech Republic KF133281
L. trivialis UE27.08.2002-17a (UPS) Sweden DQ421991
L. uvidus KVP10-027 (GENT) Russia KF241546
L. vietus UE11.19.2004-1 (UPS) Sweden KF133267
L. vinaceorufescens JN2007-018 (GENT) Canada KF241542
Multifurca furcata RH7804 (NY) Costa Rica DQ421994
M. ochricompacta BB02.107 (PC) USA DQ421984
M. zonaria DED7442 (PC) Thailand DQ421990
Zelleromyces gardneri USA DQ453696
Z. giennensis Spain AF230900
Z. hispanicus Spain AF231913
Z. sp. Australia, Tasmania JF960852
Z. sp. Australia, Tasmania JF960853
Z. sp. Australia, Tasmania JF960854
VERBEKEN ET AL.
166 Phytotaxa 181 (3) © 2014 Magnolia Press
Results
The results of the ML analyses (Fig. 1) show that angiocarpous species are included in all major clades of the genus
Lactarius, corresponding to L. subgen. Russularia, L. subgen. Piperites and L. subgen. Plinthogalus. The newly
described species L. bisporus is not included in either of these subgenera, but belongs to Lactarius, as the placement in
a joined and well-supported clade with L. pomiolens Verbeken & Stubbe shows. The majority of angiocarpous species
are included in mixed clades together with agaricoid taxa. The clade of L. bisporus with two angiocarpous taxa is an
exception.
FIGURE 1. The obtained ML topology based on ITS sequences. Bootstrap values are indicated if they exceed 50%. Names in orange are
angiocarpous milk cap species for which ITS sequences are available on GenBank. (KF856293). Lactarius bisporus sp. nov. holotype is
indicated in red. The scale bar represents the expected number of nucleotide changes per site.
Taxonomy
Lactarius bisporus Verbeken & F. Hampe sp. nov. (Figs. 2–3)
Mycobank number: MB 807217
Diagnosis: Basidiomata 10–35 mm diam., 10–15 mm alta, subglobosa, subregularia. Peridium leviter velutinosum, ochraceum ad
luteobrunneum. Stipes absens. Gleba labyrinthuloidea, albida, cum locis parvis, cum paucis venis gelatinosis. Columella absens.
Latex excarsus, albidus, immutabilis. Sporae globosae ad subglobosae, 9.8–11.3–13.0 × 9.8–11.2–12.5 µm, amyloideae, spinis et
verrucis usque ad 2(3) µm altis ornatae, macula suprahilaris non amyloidea. Basidia 45–55 × 12–18 µm, bispora, subcylindrata ad
clavata. Cystidia 25–55 × 12–20 µm, cylindrata, clavata vel utriformia. Peridiopellis ex hyphis periclinis.
Typus:—THAILAND. Chiang Mai Province, Mae On District, about 3 km from Tharnthong Lodges, 725 m elev. N18°51’55’’, E99°17’23’’,
17 July 2012, leg. Felix Hampe, FH 12–160 (holotype, GENT!), MFLU 12–0650 (isotype, MFLU!).
LACTARIUS FROM THAILAND Phytotaxa 181 (3) © 2014 Magnolia Press 167
FIGURE 2. Basidiocarps of Lactarius bisporus (FH 12–160, holotype, photo by Felix Hampe).
Etymology:—referring to the strictly 2-spored basidia.
Basidiocarps angiocarpous, semihypogeous, 10–35 mm diam., 10–15 mm high, subglobose, rather regular.
Peridium with minutely velutinous surface, chamois-leather-like, ochraceous yellow to yellow-brown. Stipe absent.
Columella absent. Gleba whitish, strongly labyrinthuloid, with small loci, with some, but very few, gelatinous veins
between the loci. Latex scarce, whitish hyaline, unchanging on the gleba and unchanging on white paper, unchanging
with KOH. Taste bitter to astringent, disagreeable but not burning acrid. Smell not remarkable. Basidiospores globose
to subglobose, 9.8–11.3–13.0 × 9.8–11.2–12.5 µm, Q = 1.00–1.01–1.04, n = 20; ornamentation amyloid, consisting of
isolated, up to 2(3) µm high spines and isolated warts; spines conical, subacute, with rounded to tapering, rarely forked
apex; apiculus up to 2.5(3) µm long; plage not distinct, inamyloid. Basidia 2-spored, subcylindrical to clavate, sometimes
irregular and somewhat tortuous, 45–55 × 12–18 µm, with 5–12 µm long sterigmata. Macrocystidia very variable in
shape, clavate, subcapitate, ventricose or lageniform, 25–55 × 12–20 µm, sometimes slightly but distinctly thick-walled.
Pseudocystidia present. Peridiopellis consisting of 2–3 µm broad hyphae which are rather periclinally arranged in the
subpellis, but form a densely intricate and interwoven layer on top and are embedded in a gelatinous matrix.
Habitat:—primary tropical forest with Dipterocarpus tuberculatus Roxb. Known only from the type collection.
Discussion
The ITS analysis shows that this new species of truffle-like Russulales belongs to the genus Lactarius, as all other
known angiocarpous milkcaps known so far. While it is obvious from the tree that the angiocarpous habit evolved
several times in the three major clades of the genus, representing L. subg. Piperites, L. subg. Russularia and L. subg.
Plinthogalus, we see that this new Thai species is most closely related with L. pomiolens, a species from Sri Lanka
(Verbeken et al. 2014). Lactarius pomiolens, also an angiocarpous taxon, forms one of the earlier diverging lineages
in the genus and has not been assigned to any of the existing subgenera. Both angiocarpous species L. bisporus and
L. pomiolens form a separate isolated small clade as does the group around L. kabansus Pegler & Piearce, a tropical
African species showing some superficial resemblances with L. subg. Plinthogalus and the morphologically very
isolated dark-spored L. chromospermus Pegler.
Morphologically, this new species clearly differs from L. pomiolens by the 2-spored basidia and the isolated
spines and warts as spore ornamentation. The spores in L. pomiolens are extremely large for spores formed by 4-
spored basidia (10.5–12.0–13.3 × 10.2–11.5–12.8 µm) and have very high wings. As already stated before (Verbeken
et al. 2014) the spore ornamentation within truffle-like milkcaps shows large differences: isolated spines or warts
on one hand, high ridges on the other hand, but this character does not seem to contain much phylogenetic signal.
VERBEKEN ET AL.
168 Phytotaxa 181 (3) © 2014 Magnolia Press
Furthermore, L. pomiolens has latex that turns immediately sulphur yellow while the latex in L. bisporus is white and
unchanging. As the name indicates, L. pomiolens is characterized by a strong and sweet apple-like smell.
Many species with normally 4-spored basidia show a low percentage of 2-spored ones, but exclusively 2-spored
basidia are rare in the genus. It is remarkable that though exclusively 2-spored basidia are very exceptional in agaricoid
milkcaps (such as e.g. L. acerrimus Britzelm.), they seem to be more common in angiocarpous representatives. Other
Asian 2-spored angiocarpous milkcap species are L. echinus Stubbe & Verbeken, L. falcatus Verbeken & Van de
Putte, Arcangeliella lactifera (B.C. Zhang & Y.N. Yu) J.M. Vidal and A. nanjingensis (B. Liu & K. Tao) J.M. Vidal.
Both L. echinus and A. lactifera differ from the newly proposed species by the longer (up to 4 µm) and more regular
spines. Lactarius falcatus has lower spines, at most 2 µm long, which are typically curved at the apex and also differs
macroscopically by latex that is turning immediately bright pale yellow. Martellia nanjingensis has smaller spores (8–
11.5 × 7.5–10 µm) with spines up to 1.5 µm long and differs macroscopically by the brown to dark brown basidiocarps
(Tao et al. 1993).
FIGURE 3. a. Basidiospores (FH 12–160, holotype), b. basidia (FH 12–160, holotype), c. cystidia (FH 12–160, holotype), d. peridiopellis
(FH 12–160, holotype), (scale bar = 10 μm).
LACTARIUS FROM THAILAND Phytotaxa 181 (3) © 2014 Magnolia Press 169
Acknowledgements
The authors are grateful for the help and assistance from the colleagues at Mae Fah Luang University and Chiang Mai
University (Thailand) and Ghent University (Belgium). The third author is supported by the joint doctorate program of
the “Bijzonder Onderzoeksfonds Ghent University” (BOF).
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... Lactarius Pers. is a genus of milkcap mushrooms in the family Russulaceae and is one of the dominant ectomycorrhizal genera in many terrestrial ecosystems (Verbeken et al. 2014, Wang et al. 2015, Wang 2016, 2017a, 2017b, Shi et al. 2018, Wang et al. 2018a, Lee et al. 2019. Lactarius forms ectomycorrhizal associations with many ecologically and economically important tree species belonging to families Pinaceae, Salicaceae, Betulaceae, Fagaceae, Cistaceae, Dipterocarpaceae, Fabaceae, Phyllanthaceae and Myrtaceae in temperate, tropical and subtropical forests , Lee et al. 2019. ...
... Lactarius subg. Russularia is a monophyletic group in family Russulaceae (Wang et al. 2018a) and is a common group of milkcap occurring almost worldwide (Verbeken et al. 2014, Wisitrassameewong et al. 2014a, 2016. The subgenus is dominant in all temperate zones, poorly represented in South America, completely absent or negligible in tropical Africa but with an important distribution in South-East Asia (Wisitrassameewong et al. 2015). ...
... The subgenus is dominant in all temperate zones, poorly represented in South America, completely absent or negligible in tropical Africa but with an important distribution in South-East Asia (Wisitrassameewong et al. 2015). 40 species of this subgenus have been described from South-East Asia so far mainly from China, Papua New Guinea, India, Korea, Thailand and Japan (Das et al. 2004, Verbeken et al. 2014, Wisitrassameewong et al. 2014a, 2014b, 2016, Wang 2017a, 2017b, Wang et al. 2018b, Lee et al. 2019). ...
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Shiwalik Range, also called Outer Himalayas are geologically youngest and lowest mountain range of Himalaya which extends for more than 1,000 miles from the Teesta River in Sikkim state of northeastern India, through Nepal across northwestern India. Tropical moist deciduous forests of Shiwalik range are largely dominated by dipterocarp tree Shorea (sal), which is commonly known as sal forests. Ectomycorrhiza (ECM) play important role in sal ecosystem as it forms mutualistic association with a variety of basidiomycetous fungi. Occurrence of ECM fungi in tropical sal forests of Indian Shiwaliks was undertaken, and Russulaceae was found to be the dominant fungal family. In this paper the morphoanatomical details of mycorrhizal roots of dipterocarp tree Shorea associated with two russulaceous genera, Russula and Lactifluus were investigated for the first time. Ectomycorrhizae are distinguished by differences in the size and colour of mycorrhizal system, surface texture, extraradical mycelium, mantle, shape and size of hyphal cells, presence or absence of cystidia, cell shape of Hartig net, as well as the differing colour reaction in Melzer’s reagent. Ectomycorrhizal roots of sal associated with Russula species were light brown to greyish brown, and have plectenchymatous gelatinised outer mantle layers having abundant cystidial elements. Ectomycorrhizae of Lactifluus species were reddish brown to light brown, lack cystidial elements and have thin walled emanating hyphae. The presence of simple, septate agaricoid hyphae and laticifers in the inner mantle layer indicate the association of this species with sal. Both ECM share a pseudoparenchymatous inner mantle, paraepidermal Hartig net, as well as absence of rhizomorphs and clamp connections.
... Lactarius Pers. is a genus of milkcap mushrooms in the family Russulaceae and is one of the dominant ectomycorrhizal genera in many terrestrial ecosystems (Verbeken et al. 2014, Wang et al. 2015, Wang 2016, 2017a, 2017b, Shi et al. 2018, Wang et al. 2018a, Lee et al. 2019. Lactarius forms ectomycorrhizal associations with many ecologically and economically important tree species belonging to families Pinaceae, Salicaceae, Betulaceae, Fagaceae, Cistaceae, Dipterocarpaceae, Fabaceae, Phyllanthaceae and Myrtaceae in temperate, tropical and subtropical forests , Lee et al. 2019. ...
... Lactarius subg. Russularia is a monophyletic group in family Russulaceae (Wang et al. 2018a) and is a common group of milkcap occurring almost worldwide (Verbeken et al. 2014, Wisitrassameewong et al. 2014a, 2016. The subgenus is dominant in all temperate zones, poorly represented in South America, completely absent or negligible in tropical Africa but with an important distribution in South-East Asia (Wisitrassameewong et al. 2015). ...
... The subgenus is dominant in all temperate zones, poorly represented in South America, completely absent or negligible in tropical Africa but with an important distribution in South-East Asia (Wisitrassameewong et al. 2015). 40 species of this subgenus have been described from South-East Asia so far mainly from China, Papua New Guinea, India, Korea, Thailand and Japan (Das et al. 2004, Verbeken et al. 2014, Wisitrassameewong et al. 2014a, 2014b, 2016, Wang 2017a, 2017b, Wang et al. 2018b, Lee et al. 2019). ...
... Sequestrate Russulaceae seem to be widespread throughout the globe, especially in arid and semi-arid regions of Australia and New Zealand (Bougher 1997, Lebel 1998, 2001, 2003a, b, Lebel & Trappe 2000, Bougher & Lebel 2001, Trappe & Claridge 2003, Lebel & Tonkin 2007 and North America (Zeller & Dodge 1919, 1937, Singer & Smith 1960, Smith 1963, Miller & Lebel 1999, Fogel & States 2001, Desjardin 2003, Smith et al. 2006). However, several sequestrate species have also been documented in tropical forests in Africa (Dring & Pegler 1978, Eberhardt & Verbeken 2004, Verbeken & Walleyn 2010, Beenken et al. 2016) and Asia (Corner & Hawker 1953, Heim 1959, Zhang & Yu 1990, Tao et al. 1993, Verbeken et al. 2014a, as well as in temperate Nothofagus forests in Patagonia (Trierveiler-Pereira et al. 2015). ...
... Piperites) and Lactarius subg. Russularia, all of them receiving significant or subsignificant support, in agreement with previous analyses of Lactarius (Eberhardt & Verbeken 2004, Buyck et al. 2008, Verbeken et al. 2014a, b, Barge et al. 2016 a result similar to that of Verbeken et al. (2014b), probably due to the insufficient phylogenetic signal in the species analyzed, gaps in the phylogenetic diversity because of an incomplete sampling, or too much phylogenetic noise in the dataset employed. However, a significantly supported clade formed by L. borzianus, L. josserandii (= Z. hispanicus) and some gymnocarpic species, such as L. fulvissimus and L. subsericatus, in accordance with results of Verbeken et al. (2014b), Liu et al. (2015), or Barge et al. (2016). ...
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... In this respect it is worth to mention that sister to both our new species collected in dry dipterocarp forests are sequestrate species (Figs. 1, 2) 30 and both species can be members of two independent seasonal drought tolerant lineages within Amoeninae. Despite our effort, we did not collect any sequestrate species of Russula from dry dipterocarp forests, but during our field excursions, members of our expedition collected sequestrate Lactarius and Entoloma 31,32 . ...
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Dry dipterocarp forests are among the most common habitat types in Thailand. Russulaceae are known as common ectomycorrhizal symbionts of Dipterocarpaceae trees in this type of habitat. The present study aims to identify collections of Russula subsection Amoeninae Buyck from dry dipterocarp forests in Thailand. A multi-locus phylogenetic analysis placed Thai Amoeninae collections in two novel lineages, and they are described here as R. bellissima sp. nov. and R. luteonana sp. nov. The closest identified relatives of both species were sequestrate species suggesting that they may belong to drought-adapted lineages. An analysis of publicly available ITS sequences in R. subsect. Amoeninae did not confirm evidence of any of the new species occurring in other Asian regions, indicating that dry dipterocarp forests might harbor a novel community of ectomycorrhizal fungi. Macromorphological characters are variable and are not totally reliable for distinguishing the new species from other previously described Asian Amoeninae species. Both new species are defined by a combination of differentiated micromorphological characteristics in spore ornamentation, hymenial cystidia and hyphal terminations in the pileipellis. The new Amoeninae species may correspond to some Russula species collected for consumption in Thailand, and the detailed description of the new species can be used for better identification of edible species and food safety in the region.
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