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Research Article
Rostrupomyces, a new genus to accommodate Xerocomus sisongkhramensis, and a new Hemileccinum species (Xerocomoideae, Boletaceae) from Thailand
expand article infoSanthiti Vadthanarat§, Bhavesh Raghoonundon§, Saisamorn Lumyong|, Olivier Raspé§#¤
‡ Ubon Ratchathani University, Ubon Ratchathani, Thailand
§ Mae Fah Luang University, Chiang Rai, Thailand
| Chiang Mai University, Chiang Mai, Thailand
¶ Academy of Science, The Royal Society of Thailand, Bangkok, Thailand
# Meise Botanic Garden, Meise, Belgium
¤ Service Général de l’Enseignement Supérieur et de la Recherche Scientifique, Fédération Wallonie-Bruxelles, Brussels, Belgium
Open Access

Abstract

A new genus, Rostrupomyces is established to accommodate Xerocomus sisongkhramensis based on multiple protein-coding genes (atp6, cox3, tef1, and rpb2) analyses of a wide taxon sampling of Boletaceae. In our phylogeny, the new genus was sister to Rubinosporus in subfamily Xerocomoideae, phylogenetically distant from Xerocomus, which was highly supported as sister to Phylloporus in the same subfamily Xerocomoideae. Rostrupomyces is different from other genera in Boletaceae by the following combination of characters: rugulose to subrugulose pileus surface, white pores when young becoming pale yellow in age, subscabrous stipe surface scattered with granulose squamules, white basal mycelium, unchanging color in any parts, yellowish brown spore print, and broadly ellipsoid to ellipsoid, smooth basidiospores. In addition, Hemileccinum inferius, also from subfamily Xerocomoideae, is newly described. Detailed descriptions and illustrations of the new genus and new species are presented.

Key words

atp6, Boletales, cox3, fungal diversity, multigene phylogeny, one new species, taxonomy, Tropical Asia

Introduction

Xerocomoideae Singer, which is one of the six subfamilies in Boletaceae Chevall, was established in 1945 with Xerocomus Quél. as the typus. At present, the subfamily consists of 12 genera, namely Alessioporus Gelardi, Vizzini & Simonini, Amylotrama Bloomfield, Davoodian, Trappe & T. Lebel, Aureoboletus Pouzar, Boletellus Murrill, Heimioporus E. Horak, Hemileccinum Šutara, Hourangia Xue T. Zhu & Zhu L. Yang, Phylloporus Quél., Pulchroboletus Gelardi, Vizzini & Simonini, Rubinosporus Vadthanarat, Raspé & Lumyong, Veloboletus Fechner & Halling, and Xerocomus (Šutara 2008; Gelardi et al. 2014; Wu et al. 2014; Zhu et al. 2015; Wu et al. 2016; Crous et al. 2020; Lebel et al. 2022; Vadthanarat et al. 2022). The typical characters of species in this subfamily are boletoid or phylloporoid, rarely sequestrate basidiomata; dry or viscid pileus with smooth or subtomentose to tomentose pellis; absence or rarely presence of a veil; off-white, yellowish white, yellowish to yellow context; at least some basidiome parts often bluing, sometimes reddening or unchanging; smooth or ornamented stipe surface; hymenophore yellowish to yellow to bright yellow or cream to dull yellow to yellow to gray in sequestrate forms; basidiospores with bacillate, reticulate, tiny warts, pinholes, longitudinally striate, pitted ornamentations, or occasionally smooth; spore deposit with more or less olive-brown tint, rarely dark ruby (e. g. Gelardi et al. 2014; Wu et al. 2014; Zhu et al. 2015; Wu et al. 2016; Crous et al. 2020; Lebel et al. 2022; Vadthanarat et al. 2022).

Hemileccinum, one of the genera belonging to the Xerocomoideae, was established in 2008 to accommodate two Boletus species, namely B. depilatus Redeuilh and B. impolitus Fr. In 2012, a new genus named Corneroboletus N.K. Zeng & Zhu L. Yang was established to accommodate Boletus indecorus Massee (Zeng et al. 2012). However, Corneroboletus was later synonymized with Hemileccinum (Wu et al. 2016). Hemileccinum currently comprises 13 species worldwide, namely H. albidum Mei Xiang Li, Zhu L. Yang & G. Wu, H. brevisporum Mei Xiang Li, Zhu L. Yang & G. Wu, H. brunneotomentosum (B. Ortiz) Nitson & J.L. Frank, H. depilatum (Redeuilh) Šutara, H. ferrugineipes Mei Xiang Li, Zhu L. Yang & G. Wu, H. floridanum J.A. Bolin, A.E. Bessette, A.R. Bessette, L.V. Kudzma, A. Farid & J.L. Frank, H. hortonii (A.H. Sm. & Thiers) M. Kuo & B. Ortiz, H. impolitum (Fr.) Šutara (typus), H. indecorum (Massee) G. Wu & Zhu L. Yang, H. parvum Mei Xiang Li, Zhu L. Yang & G. Wu, H. rubropunctum (Peck) Halling & B. Ortiz, H. rugosum G. Wu & Zhu L. Yang, H. subglabripes (Peck) Halling (Index Fungorum, accessed on 23 March 2023). Hemileccinum species share the following combination of characters: boletoid basidiomata, glabrous to subtomentose, smooth to rugose pileus surface, which turns violet with NH3 vapours; tubes depressed around the stipe apex, pores at first light yellow to deep yellow becoming olive-yellow in age, concolorous with tubes, unchanging; olive spore deposit; central stipe, whose surface is always ornamented with scales concolorous with stipe, unchanging; pale yellow to light yellow context, unchanging; pileipellis a trichodermium with broad hyphae or an epithelium, sometime with filamentous terminal elements; pleurocystidia present, fusoid to lageniform; spores boletoid, subfusoid or ellipsoid in face view, smooth under light microscope, irregularly tiny warted and pinholed or rarely smooth under SEM; clamp connections absent (Šutara 2008; Halling et al. 2015; Wu et al. 2016; Index Fungorum 443:1, 2020; Kuo and Ortiz-Santana 2020; Farid et al. 2021; Li et al. 2021).

The first study of poroid mushrooms from Thailand was published in 1902, with descriptions of five new species, namely Boletus lacunosus Rostr. [current name: Austroboletus rostrupii (Syd. & P. Syd.) E. Horak], Boletus costatus Rostr., Suillus changensis Rostr. [current name: Boletus changensis (Rostr.) Sacc. & D. Sacc.], Suillus hygrophanus Rostr. [current name: Boletus hygrophanus (Rostr.) Sacc. & D. Sacc.], and Suillus velatus Rostr. [current name: Veloporphyrellus velatus (Rostr.) Y.C. Li & Zhu L. Yang] (Rostrup 1902; Saccardo and Saccardo 1905; Horak 1980; Li et al. 2014). At that time, they were classified to belong to the Polyporaceae; however, later they were all moved to family Boletaceae. No new taxa in Boletaceae were described from Thailand during the following one hundred years. It is only in 2006 that again a new species, Rhodactina incarnata Zhu L. Yang, Trappe & Lumyong, was described from Chiang Mai Province, northern Thailand (Yang et al. 2006). In 2009, Spongiforma thailandica Desjardin, Manfr. Binder, Roekring & Flegel was described as a new genus and species from Nakorn Nayok Province, central Thailand (Desjardin et al. 2009). After that, molecular phylogenetic analyses have been widely used in Boletaceae taxonomy. Two more new Boletaceae genera including Cacaoporus Raspé & Vadthanarat and Rubinosporus Vadthanarat, Raspé & Lumyong were described from Chiang Mai Province, northern Thailand (Vadthanarat et al. 2019b, 2022). During that period, twenty-seven new species were also described from the country, among which nine belong in subfamily Xerocomoideae, namely Heimioporus subcostatus Vadthanarat, Raspé & Lumyong, Phylloporus castanopsidis M.A. Neves & Halling, P. dimorphus M.A. Neves & Halling, P. infuscatus M.A. Neves & Halling, Phylloporus pusillus Raspé, K.D. Hyde & Chuankid, P. rubiginosus M.A. Neves & Halling, P. subrubeolus Chuankid, K.D. Hyde & Raspé, Rubinosporus auriporus Vadthanarat, Raspé & Lumyong, Xerocomus sisongkhramensis Khamsuntorn, Pinruan & Luangsa-ard (Neves et al. 2012; Halling et al. 2014; Raspé et al. 2016; Vadthanarat et al. 2018; Chuankid et al. 2019; Vadthanarat et al. 2019a, 2019b, 2020; Chuankid et al. 2021; Raghoonundon et al. 2021; Vadthanarat et al. 2021; Tan et al. 2022; Vadthanarat et al. 2022).

In this study, several collections of boletes belonging to the subfamily Xerocomoideae were obtained from northern and northeastern Thailand. They were carefully studied based on morphology as well as family-wide and subfamily-wide phylogenetic analyses. Some of them were identified as a new Hemileccinum species. Some collections were identified as X. sisongkhramensis based on morphological characters and the megablast result of the ITS region. However, following multiple gene phylogenetic analyses based on four protein-coding gene (atp6, cox3, tef1, and rpb2), X. sisongkhramensis appeared phylogenetically distant from other Xerocomus species and distinct from existing genera in Boletaceae. Moreover, the detailed morphology did not fit any known Xerocomoideae genus. Therefore, Rostrupomyces is introduced to accommodate X. sisongkhramensis. Finally, a new Hemileccinum species is introduced with full descriptions and illustrations.

Materials and methods

Specimens collecting

Fresh basidiomata of boletes in subfamily Xerocomoideae were collected in Chiang Mai and Chiang Rai provinces in northern Thailand, and Ubon Ratchathani and Sisaket provinces in northeastern Thailand between 2015 and 2021. They were photographed in the field and then wrapped in aluminum foil for later description in the laboratory on the same day. The specimens were then dried in an electric drier at 45–50 °C. Examined specimens were deposited at MFU, BKF or CMUB herbaria.

Morphological study

Macroscopic descriptions were made based on the detailed field notes and photos of fresh basidiomata. Color codes were given based on Kornerup and Wanscher (1978). Macrochemical reactions (color reactions) were observed using aqueous solutions of 10% potassium hydroxide (KOH), and 28–30% NH4OH. Microscopic structures were observed from dried specimens rehydrated in 5% KOH or 1% ammoniacal Congo red. For the measurements of microscopic features, a minimum of 50 basidiospores or 20 for other structures, were randomly chosen and measured under a Nikon Eclipse Ni compound microscope using NIS-Elements D version 5.10 software. The notation ‘[x/y/z]’ represents the number of basidiospores ‘x’ measured from the number of basidiomata ‘y’ of the number of collections ‘z’. The measurements of microscopic structures are presented in the following format (a–) b–c–d (–e), in which ‘c’ represents the average, ‘b’ is the 5th percentile, ‘d’ is the 95th percentile, and ‘a’ and ‘e’ the extreme values, shown in parentheses. Q is the length/width ratio. Sections of the pileipellis were cut radially, perpendicularly to the surface halfway between the centre and margin of pileus. Sections of stipitipellis were taken halfway along the stipe length (Li et al. 2011; Hosen et al. 2013; Li et al. 2014; Zhu et al. 2015). All line drawings of microscopic features were drawn by free hand using an Olympus compound microscope model CX41 with Olympus Camera Lucida model U−DA. For scanning electron microscopy, small fragments of dried hymenophore were mounted directly onto a SEM stub with double-sided carbon tape. The samples were coated with gold, examined and photographed using a TESCAN MIRA’s 4th generation SEM.

DNA extraction, PCR amplification and DNA sequencing

Genomic DNA was extracted from tissue of dried specimen or fresh tissue preserved in CTAB, using a CTAB isolation procedure adapted from Doyle and Doyle (1990). Portions of the genes atp6, cox3, rpb2, and tef1 were amplified by polymerase chain reaction (PCR). The primer pairs ATP6-1M40F/ATP6-2M (Raspé et al. 2016), COX3M1-F/ COX3M1-R (Vadthanarat et al. 2019b), bRPB2-6F/bRPB2-7.1R (Matheny 2005), and EF1-983F/EF1-2218R (Rehner and Buckley 2005) were used to amplify atp6, cox3, rpb2, and tef1, respectively. PCR products were purified by adding 1 U of exonuclease I and 0.5 U FastAP alkaline phosphatase (Thermo Scientific, St. Leon-Rot, Germany) and incubated at 37 °C for 1 h, followed by inactivation at 80 °C for 15 min. Standard Sanger sequencing was performed in both directions by Macrogen with PCR primers, except for atp6, for which universal primers M13F-pUC(-40) and M13F(-20) were used. For tef1, additional sequencing was performed with two internal primers, EF1-1577F and EF1-1567R (Rehner and Buckley 2005).

Alignment and phylogeny inference

The two reads of newly generated sequences were assembled in GENEIOUS Pro v. 6.0.6 (Biomatters) and blasted against GenBank database to check that they were not from unrelated contamination. For the Boletaceae-wide tree, the introns in rpb2 and tef1 were removed based on the amino acid sequence of previously published sequences. The sequence datasets including the newly generated sequences and selected sequences representative of the whole family downloaded from GenBank, were separately aligned for each gene using MAFFT on the server accessed at http://mafft.cbrc.jp/alignment/server/ (Katoh and Standley 2013). Before combining the four gene partitions (atp6, cox3, rpb2 exons + tef1 exons), topological incongruence between the datasets was assessed using maximum likelihood (ML) on each of mitochondrial genes (atp6 + cox3) dataset and nuclear genes (rpb2 exons + tef1 exons) dataset. Paired trees were examined for conflicts involving only nodes with ML bootstrap (BS) ≥ 70%. After that, the Maximum likelihood phylogenetic inference was performed using RAxML (Stamatakis 2006) on the CIPRES web portal (RAxML-HPC2 on XSEDE; Miller et al. 2009). The phylogenetic tree was inferred by a single partitioned analysis with four character sets (one for each gene), using the GTRCAT model with 25 categories. The outgroup consisted of two Buchwaldoboletus and seven Chalciporus species from subfamily Chalciporoideae, based on previously published phylogenies. Statistical support of clades was obtained with 1,000 rapid bootstrap replicates. For Bayesian Inference (BI), the best-fit model of substitution among those implementable in MrBayes was estimated separately for each region using jModel-test (Darriba et al. 2012) on the CIPRES portal, based on the Bayesian Information Criterion (BIC). The selected models were HKY+I+G for atp6, GTR+I+G for cox3, K80+I+G for rpb2 exons, and SYM+I+G for tef1 exons. Partitioned Bayesian analysis was performed on the CIPRES web portal (MrBayes on XSEDE; Ronquist et al. 2012). Two runs of five chains were run for 15,000,000 generations and sampled every 1,000 generations. At the end of the run, the average deviation of split frequencies was 0.008563. The PSRF values were equal or greater than 1, and ESS values were greater than 200 for all parameters. A total of 11,252 trees were used to construct a 50% majority rule consensus tree and calculate the Bayesian posterior probabilities (BPPs).

A second, Xerocomoideae-wide tree, was also inferred from sequences of selected taxa in Xerocomoideae. Sequences were also separately aligned for each of the genes using the MAFFT online software, with introns included. Then, the topological incongruence between the datasets was also assessed using ML on each gene of five character sets, atp6, cox3, rpb2 exons, tef1 exons, and the three introns of tef1 + an intron of rpb2. Since there was no supported conflict, the ML phylogenetic tree was inferred by a single partitioned analysis with the five character sets (atp6, cox3, rpb2 exons, tef1 exons, and rpb2 intron + tef1 introns), using the same software and model that was used for family Boletaceae-wide phylogeny. Based on the latter, three Hourangia, three Phylloporus, and three Xerocomus species in the same subfamily Xerocomoideae were used as the outgroup. For BI, partitioned Bayesian analysis was performed with MrBayes 3.2.6 software for Windows. The selected models were GTR+I+G for atp6 and cox3, K80+I+G for rpb2 exons, and SYM+I+G for tef1 exons, HKY+I+G intron of rpb2 + introns of tef1. Two runs of five chains were sampled every 200 generations and stopped after 700,000 generations. At the end of the run, the average deviation of split frequencies was 0.007178. The PSRF values were equal or greater than 1, and ESS values were greater than 200 for all parameters. A total of 2,495 trees were used to construct a 50% majority rule consensus tree and calculate the BPPs.

Results

Phylogenetic analyses

A total of 39 sequences were newly generated in this study and deposited in GenBank. The ML phylograms from the mitochondrial and nuclear datasets were similar in topology without any supported conflict. The Boletaceae-wide, two-genome alignment contained 743 sequences comprising four genes (146 for atp6, 110 for cox3, 231 for rpb2, 256 for tef1) from 262 voucher specimens (Table 1) corresponding to 254 species, and was 2946 characters long (DOI: 10.6084/m9.figshare.23301077). ML and BI trees of the concatenated four-character set showed similar topologies without any supported conflicts (Bootstrap Support values, BS ≥ 70% and posterior probabilities, PP ≥ 0.90; Fig. 1). In the four-gene ML phylogram, the six subfamily clades were retrieved, namely the Austroboletoideae G. Wu & Zhu L. Yang, Boletoideae Singer, Chalciporoideae G. Wu & Zhu L. Yang, Leccinoideae G. Wu & Zhu L. Yang, Xerocomoideae, and Zangioideae G. Wu, Yan C. Li & Zhu L. Yang. The Pulveroboletus group introduced by Wu et al. (2014, 2016) was not monophyletic; however, the monophyly of each genus in this group was highly supported. All the Xerocomus (Rostrupomyces) sisongkhramensis collections included formed a highly supported (BS = 100%, PP = 1) monophyletic group, sister to Rubinosporus (BS = 99%, PP = 1) clustered in subfamily Xerocomoideae with high support (BS = 99%, PP = 1). The other selected Xerocomus species, including the type species X. subtomentosus (voucher VDKO 0987), formed another, distinct monophyletic group (BS = 89%, PP = 1), sister to Phylloporus (BS = 79%, PP = 1). The two genera also clustered in a supported clade together with Hourangia (BS = 100%, PP = 1). Regarding Hemileccinum, all selected species formed a highly supported clade (BS = 100%, PP = 1) consisting of fourteen species-level clades, including twelve known species, one new species from Thailand (this study), and one undescribed species from China. The new species Hemileccinum inferius clustered in a supported clade (BS = 76%, PP = 0.98) together with the American H. hortonii, the Chinese H. rugosum, and an undescribed Hemileccinum species from China (voucher HKAS53421).

Figure 1. 

Boletaceae-wide Maximum Likelihood phylogenetic tree inferred from the four-gene dataset (atp6, cox3, rpb2, and tef1) (introns excluded), showing the position of the new genus Rostrupomyces in Xerocomoideae. Bootstrap support values (BS ≥ 70%) and the corresponding Bayesian posterior probabilities (PP ≥ 0.90) are shown above the supported branches. The two Buchwaldoboletus and seven Chalciporus species (subfamily Chalciporoideae) were used as outgroup. All taxa belonging to subfamilies Austroboletoideae, Boletoideae, Chalciporoideae, Leccinoideae, and Zangioideae were collapsed into subfamily clades. All generic clades in subfamily Xerocomoideae (excluding Hemileccinum and Rostrupomyces) and Pulveroboletus group with high supports, were also collapsed.

Table 1.

List of collections used for DNA analyses, with origin, GenBank accession numbers, and reference(s).

Species Voucher Origin atp6 cox3 rpb2 tef1 Reference(s)
Afroboletus aff. multijugus JD671 Burundi MH614651 MH614794 MH614747 MH614700 Vadthanarat et al. (2019b)
Afroboletus costatisporus ADK4644 Togo KT823958 MH614795* KT823991 KT824024 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Afroboletus luteolus ADK4844 Togo MH614652 MH614796 MH614748 MH614701 Vadthanarat et al. (2019b)
Amoenoboletus granulopunctatus HKAS 86007 China MW560079 MZ741478 Wu et al. (2021)
Amoenoboletus granulopunctatus HKAS 80250 China MW560080 MW566746 Wu et al. (2021)
Amylotrama banrockensis AD-C58672 Australia MN413637 Lebel et al. (2022)
Amylotrama clelandii MEL2432546 Australia MN413630 Lebel et al. (2022)
Anthracoporus cystidiatus HKAS55375 China MT110410 KT990816* Li and Yang (2021); Wu et al. (2016)*
Anthracoporus holophaeus HKAS59407 China KT990506 KT990888 Wu et al. (2016)
Anthracoporus nigropurpureus HKAS52685 China KT990459 KT990821 Wu et al. (2016)
Aureoboletus auriflammeus CFMR:BOS-699 USA MK766269 MK721060 Kuo and Ortiz-Santana (2020)
Aureoboletus catenarius HKAS54467 China KT990349 KT990711 Wu et al. (2016)
Aureoboletus duplicatoporus HKAS50498 China KF112754 KF112230 Wu et al. (2014)
Aureoboletus formosus GDGM44441 China KT291751 KT291744 Zhang et al. (2015)
Aureoboletus gentilis ADK4865 Belgium KT823961 MH614797* KT823994 KT824027 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Aureoboletus glutinosus GDGM44477 China MH700229 MH700205 Zhang et al. (2019)
Aureoboletus innixus CFMR:BOS-544 USA MK766270 MK721061 Kuo and Ortiz-Santana (2020)
Aureoboletus moravicus VDKO1120 Belgium MG212528 MH614798* MG212615 MG212573 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Aureoboletus nephrosporus HKAS74929 China KT990358 KT990721 Wu et al. (2016)
Aureoboletus pseudoauriporus JAB 80 USA MW737471 MW737490 Farid et al. (2021)
Aureoboletus raphanaceus GDGM 53127 China MN549706 MN549676 Zhang et al. (2019)
Aureoboletus singeri CFMR:BOS-468 Belize MK766274 MK721065 Kuo and Ortiz-Santana (2020)
Aureoboletus tenuis GDGM42601 China KT291754 KT291745 Zhang et al. (2015)
Aureoboletus thibetanus AFTOL-ID-450 China DQ534600* DQ366279 DQ029199 Binder and Hibbett (2006)*; Unpublished
Aureoboletus tomentosus HKAS90216 China KT990355 KT990717 Wu et al. (2016)
Aureoboletus viscidipes HKAS77103 China KT990360 KT990723 Wu et al. (2016)
Aureoboletus viscosus OR0361 Thailand MH614655 MH614801 MH614751 MH614704 Vadthanarat et al. (2019b)
Australopilus palumanus REH-9433 Australia MK766276 MK721067 Kuo and Ortiz-Santana (2020)
Austroboletus cf. dictyotus OR0045 Thailand KT823966 MH614802* KT823999 KT824032 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Austroboletus cf. subvirens OR0573 Thailand MH614656 MH614803 MH614752 MH614705 Vadthanarat et al. (2019b)
Austroboletus olivaceoglutinosus HKAS57756 China KF112764 KF112212 Wu et al. (2014)
Baorangia major OR0209 Thailand MG897421 MK372295* MG897441 MG897431 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Baorangia pseudocalopus HKAS63607 China KF112677 KF112167 Wu et al. (2014)
Baorangia rufomaculata BOTH4144 USA MG897415 MH614805* MG897435 MG897425 Phookamsak et al. 2019; Vadthanarat et al. (2019b)*
Binderoboletus segoi TWH8035 Guyana OP358290 OP358307 This study
Boletellus aff. ananas NY815459 Costa Rica KF112760 KF112308 Wu et al. (2014)
Boletellus aff. emodensis OR0061 Thailand KT823970 MH614806* KT824003 KT824036 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Boletellus ananas K(M)123769 Belize MH614658 MH614807 MH614754 MH614707 Vadthanarat et al. (2019b)
Boletellus areolatus TNS-F-61444 or BLT-7 Japan AB989025 AB999754 Sata and Hattori (2015)
Boletellus aurocontextus TNS-F-61501 or BLT-65 Japan AB989037 AB999770 Sata and Hattori (2015)
Boletellus emodensis TNS-F-61564 or BLT-128 Japan AB989053 AB999782 Sata and Hattori (2015)
Boletus aereus VDKO1055 Belgium MG212530 MH614809* MG212617 MG212575 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Boletus albobrunnescens OR0131 Thailand KT823973 MH614810* KT824006 KT824039 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Boletus botryoides HKAS53403 China KT990375 KT990738 Wu et al. (2016)
Boletus edulis VDKO0869 Belgium MG212531 MH614811* MG212618 MG212576 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Boletus rubriceps MICH:KUO-08150719 USA MK766284 MK721076 Kuo and Ortiz-Santana (2020)
Borofutus dhakanus OR0345 Thailand MH614660 MH614814 MH614755 MH614709 Vadthanarat et al. (2019b)
Buchwaldoboletus lignicola HKAS76674 China KF112819 KF112277 Wu et al. (2014)
Buchwaldoboletus lignicola VDKO1140 Belgium MH614661 MH614815 MH614756 MH614710 Vadthanarat et al. (2019b)
Butyriboletus appendiculatus VDKO0193b Belgium MG212537 MH614816* MG212624 MG212582 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Butyriboletus cf. roseoflavus OR0230 China KT823974 MH614819* KT824007 KT824040 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Butyriboletus pseudoregius VDKO0925 Belgium MG212538 MH614817* MG212625 MG212583 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Butyriboletus roseopurpureus BOTH4497 USA MG897418 MH614818* MG897438 MG897428 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Butyriboletus subsplendidus HKAS50444 China KT990379 KT990742 Wu et al. (2016)
Butyriboletus yicibus HKAS55413 China KF112674 KF112157 Wu et al. (2014)
Cacaoporus pallidicarneus SV0221 Thailand MK372262 MK372299 MK372286 MK372273 Vadthanarat et al. (2019b)
Cacaoporus tenebrosus SV0223 Thailand MK372266 MK372303 MK372290 MK372277 Vadthanarat et al. (2019b)
Caloboletus calopus ADK4087 Belgium MG212539 MH614820 KP055030 KJ184566 Vadthanarat et al. (2018); Zhao et al. (2014a); Zhao et al. (2014b); Vadthanarat et al. (2019b)
Caloboletus firmus BOS-372 Belize MK766288 MK721080 Kuo and Ortiz-Santana (2020)
Caloboletus inedulis BOTH3963 USA MG897414 MH614821* MG897434 MG897424 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Caloboletus radicans VDKO1187 Belgium MG212540 MH614822* MG212626 MG212584 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Caloboletus yunnanensis HKAS69214 China KT990396 KJ184568 Zhao et al. (2014a); Wu et al. (2016)
Chalciporus aff. piperatus OR0586 Thailand KT823976 MH614824* KT824009 KT824042 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Chalciporus aff. rubinus OR0139 China MH614663 MH614758 MH614712 Vadthanarat et al. (2019b)
Chalciporus africanus JD517 Cameroon KT823963 MH614825* KT823996 KT824029 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Chalciporus piperatus VDKO1063 Belgium MH614664 MH614826 MH614759 MH614713 Vadthanarat et al. (2019b)
Chalciporus rubinus AF2835 Belgium KT823962 KT823995 KT824028 Raspé et al. (2016)
Chalciporus sp. OR0363 Thailand MH645586 MH645607 MH645602 MH645594 Vadthanarat et al. (2019b)
Chalciporus sp. OR0373 Thailand MH645587 MH645608 MH645603 MH645595 Vadthanarat et al. (2019b)
Chamonixia brevicolumna DBG_F28707 USA MK766291 MK721083 Kuo and Ortiz-Santana (2020)
Chamonixia caespitosa OSC117571 USA MK766293 MK721085 Kuo and Ortiz-Santana (2020)
Chiua virens OR0266 China MG212541 MH614828* MG212627 MG212585 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Chiua viridula HKAS74928 China KF112794 KF112273 Wu et al. (2014)
Crocinoboletus cf. laetissimus OR0576 Thailand KT823975 MH614833* KT824008 KT824041 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Crocinoboletus rufoaureus HKAS53424 China KF112710 KF112206 Wu et al. (2014)
Cupreoboletus poikilochromus GS10070 Italy KT157068 KT157072 Gelardi et al. (2015)
Cyanoboletus brunneoruber OR0233 China MG212542 MH614834* MG212628 MG212586 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Cyanoboletus pulverulentus RW109 Belgium KT823980 MH614835* KT824013 KT824046 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Cyanoboletus sinopulverulentus HKAS59609 China KF112700 KF112193 Wu et al. (2014)
Erythrophylloporus aurantiacus REH7271 Costa Rica MH614666 MH614829 MH614761 MH614715 Vadthanarat et al. (2019a)
Erythrophylloporus fagicola Garay215 Mexico MH614667 MH614830 MH614762 MH614716 Vadthanarat et al. (2019a)
Erythrophylloporus paucicarpus OR1151 Thailand MH614670 MH614831 MH614765 MH614719 Vadthanarat et al. (2019a)
Erythrophylloporus suthepensis SV0236 Thailand MH614672 MH614832 MH614767 MH614721 Vadthanarat et al. (2019a)
Fistulinella prunicolor REH9880 Australia MH614676 MH614840 MH614771 MH614725 Vadthanarat et al. (2019b)
Harrya chromapes HKAS50527 China KF112792 KF112270 Wu et al. (2014)
Harrya moniliformis HKAS49627 China KT990500 KT990881 Wu et al. (2016)
Heimioporus conicus HKAS53451 China KF112805 KF112226 Wu et al. (2016)
Heimioporus australis REH9288 Australia KP327703 Halling et al. (2015)
Heimioporus cooloolae REH9817 Australia KP327710 Halling et al. (2015)
Heimioporus fruticicola REH8962 Australia KP327696 Halling et al. (2015)
Heimioporus gaojiaocong HKAS80582 China KT990409 KT990770 Wu et al. (2016)
Heimioporus ivoryi REH8620 Costa Rica KP327683 Halling et al. (2015)
Heimioporus japonicus OR0114 Thailand KT823971 KT824004 KT824037 Raspé et al. (2016)
Heimioporus japonicus SV0016 Thailand MT136776 MT136766 MT136771 Vadthanarat et al. (2020)
Heimioporus mandarinus OR0218 Thailand MG212546 MG212632 MG212590 Vadthanarat et al. (2018)
Heimioporus subcostatus SV0235 Thailand MT136780 MT136770 MT136775 Vadthanarat et al. (2020)
Hemileccinum albidum KUN-HKAS81120 China MZ936320 MZ936352 Li et. al. (2021)
Hemileccinum inferius BR0260 Thailand OP358291 OP358312 OP358319 This study
Hemileccinum inferius SV0282 Thailand OP358292 This study
Hemileccinum brevisporum KUN-HKAS89150 China MZ936328 MZ936362 Li et. al. (2021)
Hemileccinum brevisporum HKAS59445 China KT990414 KT990775 Wu et al. (2016)
Hemileccinum depilatum AF2845 Belgium MG212547 MH614843* MG212633 MG212591 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Hemileccinum ferrugineipes KUN-HKAS115554 China MZ936330 MZ973011 Li et. al. (2021)
Hemileccinum floridanum AB16 USA MW737481 Farid et al. (2021)
Hemileccinum hortonii MICH:KUO-07050706 USA MK766377 MK721175 Kuo and Ortiz-Santana (2020)
Hemileccinum impolitum ADK4078 Belgium MG212548 MH614844* MG212634 MG212592 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Hemileccinum indecorum OR0863 Thailand MH614677 MH614845 MH614772 MH614726 Vadthanarat et al. (2019b)
Hemileccinum parvum KUN-HKAS115553 China MZ936333 MZ973010 Li et. al. (2021)
Hemileccinum rubropunctum REH-8501 USA MK766327 MK721122 Kuo and Ortiz-Santana (2020)
Hemileccinum rugosum HKAS84355 China KT990413 KT990774 Wu et al. (2016)
Hemileccinum sp. HKAS53421 China KF112751 KF112235 Wu et al. (2014)
Hemileccinum subglabripes MICH:KUO-07230802 USA MK766300 MK721092 Kuo and Ortiz-Santana (2020)
Hortiboletus amygdalinus HKAS54166 China KT990416 KT990777 Wu et al. (2016)
Hortiboletus campestris MICH:KUO-08240502 USA MK766302 MK721094 Kuo and Ortiz-Santana (2020)
Hortiboletus rubellus VDKO0403 Belgium MH614679 MH614847 MH614774 Vadthanarat et al. (2019b)
Hortiboletus subpaludosus HKAS59608 China KF112696 KF112185 Wu et al. (2014)
Hourangia cf. pumila OR0762 Thailand MH614680 MH614848 MH614775 MH614728 Vadthanarat et al. (2019b)
Hourangia cheoi HKAS52269 China KF112773 KF112286 Zhu et al. (2015)
Hourangia microcarpa HKAS53378 China KF112775 KF112300 Wu et al. (2014)
Hourangia nigropunctata HKAS 57427 China KP136978 KP136927 Zhu et al. (2015)
Hymenoboletus luteopurpureus HKAS46334 China KF112795 KF112271 Wu et al. (2014)
Imleria badia VDKO0709 Belgium KT823983 MH614849* KT824016 KT824049 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Imleria obscurebrunnea OR0263 China MH614681 MH614850 MH614776 MH614729 Vadthanarat et al. (2019b)
Imleria pallidus BOTH4356 USA MH614659 MH614812 MH614708 Vadthanarat et al. (2019b)
Indoporus squamulosus HKAS107153 China MT110409 MT110335 Li and Yang (2021)
Ionosporus longipes LEE1180 Malaysia MT085461 MH712031* MT085471 Chuankid et al. (2021); Khmelnitsky et al. (2019)
Kaziboletus rufescens HKAS74706 Bangladesh JQ928600 JQ928578 Hosen et al. (2021)
Lanmaoa angustispora HKAS74752 China KM605177 KM605154 Wu et al. (2015)
Lanmaoa asiatica OR0228 China MH614682 MH614851 MH614777 MH614730 Vadthanarat et al. (2019b)
Lanmaoa carminipes BOTH4591 USA MG897419 MH614852* MG897439 MG897429 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Lanmaoa pallidorosea BOTH4432 USA MG897417 MH614853* MG897437 MG897427 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Lanmaoa sublurida Farid 1023 USA MW737460 MW737485 Farid et al. (2021)
Leccinellum aff. crocipodium HKAS76658 China KF112728 KF112252 Wu et al. (2014)
Leccinellum aff. griseum KPM-NC-0017832 Japan KC552164 JN378450* Unpublished; Orihara et al. (2012)*
Leccinellum cremeum HKAS90639 China KT990420 KT990781 Wu et al. (2016)
Leccinum scabrum VDKO0938 Belgium MG212549 MH614858* MG212635 MG212593 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Leccinum schistophilum VDKO1128 Belgium KT823989 MH614859* KT824022 KT824055 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Leccinum variicolor VDKO0844 Belgium MG212550 MH614860* MG212636 MG212594 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Leccinum versipelle KPM-NC-0017833 Scotland KC552172 JN378454 Orihara et al. (2016); Orihara et al. (2012)
Leccinum vulpinum KPM-NC-0017834 Scotland KC552171 JN378456 Orihara et al. (2016); Orihara et al. (2012)
Mucilopilus castaneiceps HKAS75045 China KF112735 KF112211 Wu et al. (2014)
Mucilopilus paracastaneiceps HKAS50338 China KT990391 KT990755 Wu et al. (2016)
Mucilopilus ruber HKAS84555 China MT110436 MT110364 Li and Yang (2021)
Mycoamaranthus cambodgensis SV0197 Thailand MZ355900 MZ355909 Vadthanarat et al. (2022)
Neoboletus brunneissimus OR0249 China MG212551 MH614861* MG212637 MG212595 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Neoboletus ferrugineus HKAS77718 China KT990431 KT990789 Wu et al. (2016)
Neoboletus flavidus HKAS59443 China KU974144 KU974136 Wu et al. (2016)
Neoboletus hainanensis HKAS59469 China KF112669 KF112175 Wu et al. (2014)
Neoboletus junquilleus AF2922 France MG212552 MH614862* MG212638 MG212596 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Neoboletus magnificus HKAS74939 China KF112653 KF112148 Wu et al. (2014)
Neoboletus obscureumbrinus OR0553 Thailand MK372271 MK372294 MK372282 Vadthanarat et al. (2019b)
Neoboletus tomentulosus HKAS53369 China KF112659 KF112154 Wu et al. (2014)
Neoboletus erythropus VDKO0690 Belgium KT823982 MH614864* KT824015 KT824048 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Octaviania asterosperma AQUI3899 Italy KC552159 KC552093 Orihara et al. (2016)
Octaviania cyanescens PNW-FUNGI-5603 USA KC552160 JN378438 Orihara et al. (2016); Orihara et al. (2012)
Octaviania tasmanica MEL2128484 Australia KC552157 JN378437 Orihara et al. (2016); Orihara et al. (2012)
Octaviania zelleri MES270 USA KC552161 JN378440 Orihara et al. (2016); Orihara et al. (2012)
Parvixerocomus pseudoaokii OR0155 China MG212553 MH614865 MG212597 MG212597 Vadthanarat et al. (2019b)
Paxilloboletus latisporus ADK5072 Congo MZ707870 MZ707866 Badou et al. (2022)
Paxilloboletus africanus SAB0716 Guinea MZ707869 MZ707865 Badou et al. (2022)
Phylloporus bellus OR0473 China MH580778 MH614866* MH580818 MH580798 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus brunneiceps OR0050 Thailand KT823968 MH614867* KT824001 KT824034 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Phylloporus castanopsidis OR0052 Thailand KT823969 MH614868* KT824002 KT824035 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Phylloporus maculatus OR0285 China MH580780 MH580820 MH580800 Chuankid et al. (2019)
Phylloporus pachycystidiatus HKAS53422 China KF112777 KF112288 Wu et al. (2014)
Phylloporus pelletieri WU18746 Austria MH580781 MH614869* MH580821 MH580801 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus pusillus OR1158 Thailand MH580783 MH614870* MH580823 MH580803 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus rhodoxanthus WU17978 Austria MH580785 MH614871* MH580824 MH580805 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus rubeolus OR0251 China MH580786 MH614872* MH580825 MH580806 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus rubiginosus OR0169 China MH580788 MH614873* MH580827 MH580808 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus rubrosquamosus HKAS52552 China KF112780 KF112289 Wu et al. (2014)
Phylloporus scabripes CFMR:BOS-621 Belize MK766359 MK721156 Kuo and Ortiz-Santana (2020)
Phylloporus subbacillisporus OR0436 China MH580792 MH614875* MH580831 MH580812 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus subrubeolus BC022 Thailand MH580793 MH614876* MH580832 MH580813 Chuankid et al. (2019); Vadthanarat et al. (2019b)*
Phylloporus yunnanensis OR0448 China MG212554 MH614877* MG212640 MG212598 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Porphyrellus castaneus OR0241 China MG212555 MH614878* MG212641 MG212599 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Porphyrellus porphyrosporus MB97 023 Germany DQ534609 GU187800 GU187734 Binder and Hibbett (2006); Binder et al. (2010)
Pseudoaustroboletus valens HKAS82644 China MT110431 MT110359 Li and Yang (2021)
Pulchroboletus sclerotiorum FLAS F 60333 USA MF614169 MF614167 Crous et al. (2019)
Pulchroboletus sclerotiorum FLAS F 60334 USA MF614164 MF614165 Crous et al. (2019)
Pulveroboletus aff. ravenelii ADK4360 Togo KT823957 MH614882* KT823990 KT824023 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Pulveroboletus aff. ravenelii ADK4650 Togo KT823959 MH614883* KT823992 KT824025 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Pulveroboletus brunneopunctatus HKAS55369 China KT990455 KT990814 Wu et al. (2016)
Pulveroboletus fragrans OR0673 Thailand KT823977 MH614884* KT824010 KT824043 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Pulveroboletus ravenelii REH2565 USA KU665635 MH614885* KU665637 KU665636 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Retiboletus aff. nigerrimus OR0049 Thailand KT823967 MH614886* KT824000 KT824033 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Retiboletus brevibasidiatus OR0570 Thailand MT085469 MT085479 MT085476 Chuankid et al. (2021)
Retiboletus brunneolus HKAS52680 China KF112690 KF112179 Wu et al. (2014)
Retiboletus fuscus OR0231 China MG212556 MH614887* MG212642 MG212600 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Retiboletus griseus MB03 079 USA KT823964 MH614888* KT823997 KT824030 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Retiboletus kauffmanii OR0278 China MG212557 MH614889* MG212643 MG212601 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Retiboletus nigerrimus HKAS53418 China KT990462 KT990824 Wu et al. (2016)
Rhodactina himalayensis CMU25117 Thailand MG212558 MG212602, MG212603 Vadthanarat et al. (2018)
Rhodactina rostratispora SV0170 Thailand MG212560 MG212645 MG212605 Vadthanarat et al. (2018)
Rossbeevera cryptocyanea KPM-NC17843 Japan KT581441 KC552072 Orihara et al. (2016)
Rossbeevera eucyanea TNS-F-36986 Japan KC552115 KC552068 Orihara et al. (2016)
Rossbeevera griseovelutina TNS-F-36989 Japan KC552124 KC552076 Orihara et al. (2016)
Rossbeevera pachydermis KPM-NC23336 New Zealand KJ001064 KP222912 Orihara et al. (2016)
Rossbeevera vittatispora TO-AUS-72 Australia KC552108 KC552065 Orihara et al. (2016)
Rostrupomyces sisongkhramensis BR0311 Thailand OP358293 OP358313 OP358320 This study
Rostrupomyces sisongkhramensis BR0313 Thailand OP358294 OP358314 OP358321 This study
Rostrupomyces sisongkhramensis BR0368 Thailand OP358295 This study
Rostrupomyces sisongkhramensis BR0371 Thailand OP358296 OP358322 This study
Rostrupomyces sisongkhramensis OR0915 Thailand OP358297 This study
Rostrupomyces sisongkhramensis OR0918 Thailand OP358298 This study
Rostrupomyces sisongkhramensis OR0919 Thailand OP358299 OP358308 OP358315 OP358323 This study
Rostrupomyces sisongkhramensis OR1004 Thailand OP358300 This study
Rostrupomyces sisongkhramensis OR1059 Thailand OP358301 This study
Rostrupomyces sisongkhramensis OR1392 Thailand OP358302 This study
Rostrupomyces sisongkhramensis OR1399 Thailand OP358303 This study
Rostrupomyces sisongkhramensis SV0155 Thailand OP358304 OP358309 OP358316 OP358324 This study
Rostrupomyces sisongkhramensis SV0219 Thailand OP358305 OP358310 OP358317 OP358325 This study
Rostrupomyces sisongkhramensis SV0225 Thailand OP358306 OP358311 OP358318 OP358326 This study
Royoungia rubina HKAS53379 China KF112796 KF112274 Wu et al. (2014)
Rubinosporus auriporus SV0101 Thailand MZ355897 MZ355906 MZ355904 MZ355902 Vadthanarat et al. (2022)
Rubinosporus auriporus SV0090 Thailand MZ355896 MZ355905 MZ355903 MZ355901 Vadthanarat et al. (2022)
Rubroboletus legaliae VDKO0936 Belgium KT823985 MH614890* KT824018 KT824051 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Rubroboletus rhodosanguineus BOTH4263 USA MG897416 MH614891* MG897436 MG897426 Phookamsak et al. (2019); Vadthanarat et al. (2019b)*
Rubroboletus rhodoxanthus HKAS84879 China KT990468 KT990831 Wu et al. (2016)
Rubroboletus satanas VDKO0968 Belgium KT823986 MH614892* KT824019 KT824052 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Rugiboletus andinus REH-7705 Costa Rica MK766316 MK721111 Kuo and Ortiz-Santana (2020)
Rugiboletus brunneiporus HKAS83209 China KM605168 KM605144 Wu et al. (2015)
Rugiboletus extremiorientalis OR0406 Thailand MG212562 MH614893* MG212647 MG212607 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Singerocomus inundabilis TWH9199 Guyana MH645588 MH645609 LC043089* MH645596 Henkel et al. (2016)*; Vadthanarat et al. (2019b)
Singerocomus rubriflavus TWH9585 Guyana MH645589 MH645610 MH645597 Vadthanarat et al. (2019b)
Spongiforma thailandica DED7873 Thailand MG212563 MH614894** MG212648 KF030436* Nuhn et al. (2013)*; Vadthanarat et al. (2018); Vadthanarat et al. (2019b)**
Spongispora temasekensis SING 0206334 Singapore MG674378 MG674377 Wu et al. (2018)
Spongispora temasekensis ACMF5 Singapore MZ803018 MZ824748 MZ803023 Raghoonundon et al. (2021)
Strobilomyces atrosquamosus HKAS55368 China KT990476 KT990839 Wu et al. (2016)
Strobilomyces echinocephalus OR0243 China MG212564 MG212649 MG212608 Vadthanarat et al. (2018)
Strobilomyces floccopus RW103 Belgium KT823978 MH614895* KT824011 KT824044 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Strobilomyces mirandus OR0115 Thailand KT823972 MH614896* KT824005 KT824038 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Strobilomyces verruculosus HKAS55389 China KF112813 KF112259 Wu et al. (2014)
Suillellus luridus VDKO0241b Belgium KT823981 MH614901* KT824014 KT824047 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Suillellus queletii VDKO1185 Belgium MH645590 MH645611 MH645604 MH645598 Vadthanarat et al. (2019b)
Suillellus subamygdalinus HKAS57262 China KF112660 KF112174 Wu et al. (2014)
Sutorius australiensis REH9441 Australia MG212567 MK386576** MG212652 JQ327032* Halling et al. (2012)*; Vadthanarat et al. (2018); Vadthanarat et al. (2019b)**
Sutorius eximius REH9400 USA MG212568 MH614902** MG212653 JQ327029* Halling et al. (2012)*; Vadthanarat et al. (2018); Vadthanarat et al. (2019b)**
Sutorius pachypus OR0411 Thailand MN067465 MN067500 MN067484 Vadthanarat et al. (2021)
Sutorius pseudotylopilus OR0378B Thailand MH614692 MH614903 MH614787 MH614740 Vadthanarat et al. (2019b)
Sutorius rubinus OR0379 Thailand MH614693 MH614904 MH614788 MH614741 Vadthanarat et al. (2019b)
Sutorius ubonensis SV0032 Thailand MN067472 MN067507 MN067491 Vadthanarat et al. (2021)
Tengioboletus glutinosus HKAS53425 China KF112800 KF112204 Wu et al. (2014)
Tengioboletus reticulatus HKAS53426 China KF112828 KF112313 Wu et al. (2014)
Turmalinea persicina KPM-NC18001 Japan KC552130 KC552082 Orihara et al. (2016)
Turmalinea yuwanensis KPM-NC18011 Japan KC552138 KC552089 Orihara et al. (2016)
Tylocinum griseolum HKAS50281 China KF112730 KF112284 Wu et al. (2014)
Tylopilus atripurpureus HKAS50208 China KF112799 KF112283 Wu et al. (2014)
Tylopilus felleus VDKO0992 Belgium KT823987 MH614906* KT824020 KT824053 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Tylopilus ferrugineus BOTH3639 USA MH614694 MH614907 MH614789 MH614742 Vadthanarat et al. (2019b)
Tylopilus otsuensis HKAS53401 China KF112797 KF112224 Wu et al. (2014)
Tylopilus vinaceipallidus OR0137 China MG212571 MH614912* MG212656 MG212613 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Tylopilus violaceobrunneus HKAS89443 China KT990504 KT990886 Wu et al. (2016)
Veloboletus limbatus REH9228 Australia MT747398 MT747397 MN413636 Crous et al. (2019)
Veloporphyrellus conicus REH8510 Belize MH614698 MH614913 MH614792 MH614745 Vadthanarat et al. (2019b)
Veloporphyrellus gracilioides HKAS53590 China KF112734 KF112210 Wu et al. (2014)
Veloporphyrellus pseudovelatus HKAS59444 China JX984519 JX984553 Li et al. (2014)
Veloporphyrellus velatus HKAS63668 China JX984523 JX984554 Li et al. (2014)
Xanthoconium affine NY00815399 USA KT990486 KT990850 Wu et al. (2016)
Xanthoconium purpureum MICH:KUO-07061405 USA MK766372 MK721170 Kuo and Ortiz-Santana (2020)
Xanthoconium sinense HKAS77651 China KT990488 KT990853 Wu et al. (2016)
Xerocomellus chrysenteron VDKO0821 Belgium KT823984 MH614914* KT824017 KT824050 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Xerocomellus cisalpinus ADK4864 Belgium KT823960 MH614915* KT823993 KT824026 Raspé et al. (2016); Vadthanarat et al. (2019b)*
Xerocomellus communis HKAS50467 China KT990494 KT990858 Wu et al. (2016)
Xerocomellus ripariellus VDKO0404 Belgium MH614699 MH614916 MH614793 MH614746 Vadthanarat et al. (2019b)
Xerocomus ferrugineus CFMR:BOS-545 USA MK766375 MK721173 Kuo and Ortiz-Santana (2020)
Xerocomus fulvipes HKAS76666 China KF112789 KF112292 Wu et al. (2014)
Xerocomus magniporus HKAS58000 China KF112781 KF112293 Wu et al. (2014)
Xerocomus rugosellus HKAS58865 China KF112784 KF112294 Wu et al. (2014)
Xerocomus spadiceus var. gracilis MICH:KUO-07080702 USA MK766378 MK721176 Kuo and Ortiz-Santana (2020)
Xerocomus subtomentosus VDKO0987 Belgium MG212572 MH614919* MG212657 MG212614 Vadthanarat et al. (2018); Vadthanarat et al. (2019b)*
Xerocomus tenax MICH:KUO-08241404 USA MK766379 MK721177 Kuo and Ortiz-Santana (2020)
Zangia citrina HKAS52684 China HQ326850 HQ326872 Li et al. (2011)
Zangia olivaceobrunnea HKAS52272 China HQ326857 HQ326876 Li et al. (2011)
Zangia roseola HKAS51137 China HQ326858 HQ326877 Li et al. (2011)

For the subfamily Xerocomoideae-wide phylogeny, no supported topological incongruence between the character sets was detected. Then, the Xerocomoideae-wide phylogeny was inferred based on the alignment containing 155 sequences of four genes (22 for atp6, 20 for cox3, 53 for rpb2, 60 for tef1) from 60 voucher specimens corresponding to 55 taxa, and was 3,161 characters long (DOI: 10.6084/m9.figshare.23301077). The ML and BI tree topologies of the concatenated five-character-set alignment were similar without any supported conflict (Fig. 2). The Xerocomoideae-wide ML tree also showed a similar topology to the Boletaceae-wide tree. However, in this subfamily Xerocomoideae-wide tree, the support of the clade consisting of the new species Hemileccinum inferius, H. hortonii, H. rugosum, and an undescribed Hemileccinum species, was lower (BS = 53%, PP = 0.71) than in the Boletaceae-wide ML tree.

Figure 2. 

Xerocomoideae-wide phylogenetic tree inferred from the four-gene dataset (atp6, cox3, rpb2, and tef1) (introns included), including new genus Rostrupomyces and selected Xerocomoideae using Maximum Likelihood and Bayesian Inference methods (ML tree is presented). The three Hourangia, three Phylloporus, and three Xerocomus species in Xerocomoideae were used as outgroup. Bootstrap support values (BS ≥ 70%) and posterior probabilities (PP ≥ 0.90) are shown above the supported branches.

Taxonomy

Rostrupomyces Vadthanarat & Raspé, gen. nov.

MycoBank No: 849050

Etymology

Named in honor of Frederik Georg Emil Rostrup (1831–1907), Danish botanist, mycologist, and plant pathologist, celebrating the 120 years of his describing the first new species of Boletaceae from Thailand in 1902.

Diagnosis

Differs from other genera in Boletaceae by the following combination of characters: rugulose to subrugulose pileus surface, white pore when young becoming grayish yellow in age, subscabrous stipe surface with scattered granulose squamules, white basal mycelium, unchanging color in any parts, yellowish brown spore print, and broadly ellipsoid to ellipsoid, smooth basidiospores.

Description

Basidiomata stipitate-pileate. Pileus convex then plano-convex to plane; surface at first rugulose then subrugulose in age, finely tomentose to tomentose, dark brown to reddish brown, becoming light brown to brown to grayish orange, unchanging when bruised; context off-white then yellowish to dull pale orange in age, unchanging when cut. Stipe central, terete, cylindrical; surface subscabrous, yellowish white to pale yellow to orange white, with scattered brown to dark brown to reddish brown granulose squamules, unchanging when bruised; basal mycelium white; context solid, white becoming off-white to yellowish white in age, unchanging when cut. Hymenophore tubulate, slightly depressed to depressed around the stipe. Tubes pale yellow then grayish yellow, separable from the pileus context, unchanging when cut. Pores roundish then subangular to angular with age; when young white then yellowish white becoming grayish yellow, unchanging when touched. Spore print yellowish brown. Basidiospores ellipsoid to broadly ellipsoid, thin-walled, smooth under light microscope and SEM. Basidia 4-spored, clavate without basal clamp connection. Cheilo- and pleurocystidia narrowly fusiform to fusiform or narrowly utriform, thin-walled. Pileipellis an intricate trichoderm, made of moderately interwoven to loosely interwoven, thin-walled hyphae. Stipitipellis arranged parallel to the surface of the stipe, composed of moderately interwoven, thin-walled hyphae, with scattered groups of rising cells to clusters of narrowly clavate to clavate cells. Clamp connections not seen in any tissue.

Typus generis

Rostrupomyces sisongkhramensis (Khamsuntorn, Pinruan & Luangsa-ard) Vadthanarat, Raghoonundon & Raspé.

Distribution

Currently known only from northern and northeastern Thailand.

Notes

Rostrupomyces can be morphologically separated from Xerocomus by the different shape and surface of basidiospores, which are ellipsoid to broadly ellipsoid with smooth under light microscope and SEM in the new genus, whereas Xerocomus produce more or less oblong to fusiform basidiospores, usually with bacillate surface under SEM (Wu et. al. 2016). Rostrupomyces also produces yellowish brown spore print, whereas Xerocomus produces olive-brown spore print. Moreover, color change upon bruising does not occur in any part of Rostrupomyces basidiomes, whereas context and hymenophore of Xerocomus always turn more or less bluish to blue when bruised or cut (Wu et. al. 2016). The most resembling genus, Hemileccinum, shares some similar characters including rugulose to subrugulose pileus surface, yellow hymenophore which is depressed around the stipe apex, subscabrous stipe surface (less so in Hemileccinum), white basal mycelium, mostly unchanging color in any parts. However, Rostrupomyces can be morphologically distinguished from Hemileccinum by the differences in spore print color, and in the shape and surface of basidiospores. Rostrupomyces produces yellowish brown spore print, broadly ellipsoid to ellipsoid basidiospores with smooth surface under light microscope and SEM. Hemileccinum produces olive-brown spore prints, boletoid basidiospores that are smooth under light microscope, but ornamented with irregular warts and pinholes under SEM. Also, the pore surface of Rostrupomyces is white in young basidiomata and becomes pale yellow when mature whereas in Hemileccinum, the pore surface is yellow in all stages (Šutara 2008; Wu et al. 2016; Farid et al. 2021; Li et al. 2021).

Rostrupomyces sisongkhramensis (Khamsuntorn, Pinruan & Luangsa-ard) Vadthanarat, Raghoonundon & Raspé, comb. nov.

MycoBank No: 851393
Figs 3, 4, 5A–B

Xerocomus sisongkhramensis Khamsuntorn, Pinruan & Luangsa-ard. Basionym.

Diagnosis

Rostrupomyces sisongkhramensis is characterised by having dark to reddish brown, becoming brown to grayish orange pileus, with rugulose to subrugulose, finely tomentose to tomentose surface; yellowish to orange white, subscabrous, longitudinally fissurate stipe surface, with moderately scattered brown to dark brown to reddish brown granulose squamules; yellow hymenophore; unchanging color in any parts; yellowish brown spore print; and broadly ellipsoid to ellipsoid smooth basidiospores.

Description

Basidiomata medium-sized. Pileus 37–94(118) mm in diameter, convex at first then plano-convex to plane, sometimes with sub-depressed at the centre; margin inflexed at first then deflexed in age, exact or slightly exceeding (up to 1 mm); surface at first rugulose especially near the margin then subrugulose in age, dull, dry to moist, finely tomentose to tomentose covered with greenish yellow (3A3–4, 3B4) matted hyphae at places (especially when young), at first dark brown to reddish brown (6–8F4–8), becoming light brown to brown to grayish orange (6D/E5–6, 5B4–5) on light yellow to brownish orange (4A3–5, 5C4) background in age, gradually paler to the margin, unchanging when bruised; context (3)5–10(14) mm thick half-way to the margin, at first firm then soft in age, color distribution even, at first off-white, slightly brownish (7D/E4–5) near the pileipellis, then yellowish to orange white (4–5A2) or occasionally yellowish (3A3–4) above the hymenium especially in age, unchanging when cut. Stipe (33)41–97(108) × 6(7)–19(20) mm, central, terete, usually cylindrical for the most part but often with wider base, rarely club-shaped; surface subscabrous longitudinally fissurate, slightly shiny, yellowish white to pale yellow to orange white (3A3 to 4A2 to 5A2), occasionally pale yellow (3A3–4) near the cap, with moderately scattered brown to dark brown to reddish brown (7D/E/F4–7) granulose squamules, unchanging when bruised; basal mycelium little developed, white (1A1); context solid, firm, at first white (1A1) becoming off-white to yellowish white (4A2) occasionally pale yellow (3A3–4) especially in the above part near the stipe surface in age, yellowish to orange gray (4–5B2–3) virgate at places, unchanging when cut. Hymenophore tubulate, slightly depressed to depressed around the stipe, with slightly decurrent tooth, sometimes almost free, mostly segmentiform to subventricose. Tubes (3)4–13 mm long half-way to the margin, at first pale yellow (4A3) then grayish yellow (4B3) when mature, separable from the pileus context, unchanging when cut. Pores 0.2–0.8(1.3) mm wide half-way to the margin, irregularly arranged, roundish then subangular to angular in age; topography subregular, composite pores frequent; color distribution even, when young white (1A1) then yellowish white (4A2) becoming grayish yellow (4B3–5) infrequently with reddish brown spots (7–8E/F8) at places in age, unchanging when touched. Odour mild fungoid. Taste mild. Spore print yellowish brown (5F5) in mass.

Macrochemical reactions : KOH, brownish orange on pileus, yellowish to pale dull orange on pileus context and stipe surface, none or yellowish on stipe context, yellowish brown to brownish orange on hymenium; NH4OH, yellowish to brownish orange (occasionally with purple aura) on pileus, yellowish to pale orange on stipe surface, yellowish to brownish on hymenium, none or yellowish on pileus context and stipe context.

Spores [591/10/10] (6.3–)6.9–7.9–9.1(–9.8) × (4.5–)4.8–5.5–6.2(–6.5) µm Q = (1.2–)1.29–1.44–1.63(–1.79). From the type (6.5–)6.9–7.7–8.8(–9.5) × (4.7–)5–5.5–6.2(–6.5) µm, Q = (1.2–)1.25–1.41–1.54(–1.63), N = 106, broadly ellipsoid to ellipsoid, thin-walled, smooth under light microscope and SEM, yellowish hyaline in water or KOH, inamyloid. Basidia 4-spored, (22–)22–26–31(–31) × (9–)9–11–13(–13) µm, clavate without basal clamp connection, hyaline to yellowish hyaline in KOH; sterigmata up to 4 µm long. Cheilocystidia (30–)30–43–58(–59) × (9–)9–11–15(–15) µm, frequent, narrowly fusiform to fusiform with obtuse apex or narrowly utriform, thin-walled, hyaline in KOH. Pleurocystidia (33–)33–43–63(–63) × (8–)8–11–13(–13) µm, infrequent, narrowly fusiform to fusiform with obtuse apex, thin-walled, hyaline in KOH. Hymenophoral trama subregular to slightly divergent, 38–82 µm wide, with subregular mediostratum 8–24 µm wide, composed of cylindrical, 4–12 µm wide hyphae, hyaline in KOH. Pileipellis an intricate trichoderm, 70–130 mm thick, made of moderately interwoven (when young) to loosely interwoven in age, thin-walled, smooth, hyaline hyphae 4–18 mm wide, branching and anastomosing at places; terminal cells 12–65 × 4–18 mm, narrowly fusiform to fusiform to broadly fusiform with slightly acuminate or obtuse apex, hyaline to yellowish pale brown in KOH. Pileus context made of strongly interwoven, thin-walled hyphae, up to 12 µm wide, hyaline in KOH. Stipitipellis arranged parallel to the surface of the stipe, composed of moderately interwoven, cylindrical, thin-walled, 3–10 µm wide hyphae, anastomosing and branching at places, sparsely scattered with groups of rising cells to clusters (up to 87 µm high) of narrowly clavate to clavate cells (21–36 × 4–9 µm), hyaline to yellowish hyaline in KOH. Caulocystidia not seen. Stipe context parallelly arranged, composed of moderately interwoven, cylindrical, thin-walled, 3–18 µm wide hyphae, hyaline to yellowish hyaline in KOH. Clamp connections not seen in any tissue.

Habitat and distribution

Solitary or in small groups (up to 4 basidiomata), or fasciculate by 2 to 3 basidiomata, on sandy loam to sandy clay loam soil in open dry dipterocarp forest and dipterocarp forest dominated by Dipterocarpaceae trees namely Anthoshorea roxburghii, Dipterocarpus obtusifolius, D. tuberculatus, D. intricatus, Pentacme siamensis, and Shorea obtusa with or without scattered Fagaceae trees. Currently known from the type locality (Nakhon Phanom province), Sisaket and Ubon Ratchathani provinces in northeastern Thailand, and also in Chiang Mai and Chiang Rai provinces in northern Thailand.

Specimens examined

Thailand, Chiang Mai Province, Muang District, Doi Suthep-Pui National Park, 18°47'39.4"N, 98°55'21.5"E, elev. 915 m, 20 July 2015, Olivier Raspé, OR1004 (CMUB, BKF, BR); ibid., 18°48'04.2"N, 98°55'44.3"E, elev. 775 m, 21 July 2015, Santhiti Vadthanarat, SV0155 (CMUB, BKF); Mae On District, 18°51'57.4"N, 99°17'22.9"E, elev. 660 m, 1 June 2015, Olivier Raspé, OR0915 (CMUB, BR); ibid., 18°51'57.0"N, 99°17'23.0"E, elev. 660 m, 1 June 2015, Olivier Raspé, OR0918 (CMUB, BR); ibid., 18°51'57.0"N, 99°17'23.0"E, elev. 660 m, 1 June 2015, Olivier Raspé, OR0919 (CMUB, BR); ibid., 18°52'13.0"N, 99°18'25.0"E, elev. 760 m, 15 August 2015, Santhiti Vadthanarat, SV0219 (CMUB, BR); ibid., 18°51'57.4"N, 99°17'22.0"E, elev. 700 m, 16 August 2015, Santhiti Vadthanarat, SV0225 (CMUB, BR); ibid., 18°51'57.7"N, 99°17'26.5"E, elev. 685 m, 1 June 2017, Santhiti Vadthanarat, SV0397 (CMUB, BR); ibid., 18°52'15.6"N, 99°18'11.5"E, elev. 800 m, 11 July 2017, Olivier Raspé, OR1392 (CMUB, BR); ibid., 18°52'15.6"N, 99°18'11.5"E, elev. 800 m, 11 July 2017, Olivier Raspé, OR1399 (CMUB, BR); ibid., 18°52'16.7"N, 99°18'13.0"E, elev. 800 m, 9 June 2021, Santhiti Vadthanarat, SV0512 (CMUB, BR); ibid. 18°52'7.9"N, 99°17'42.0"E, elev. 780 m, 10 June 2021, Santhiti Vadthanarat, SV0517 (CMUB, BR); ibid. 18°52'16.4"N, 99°17'40.5"E, elev. 820 m, 10 June 2021, Santhiti Vadthanarat, SV0518 (CMUB, BR); ibid. 18°52'12.0"N, 99°17'31.2"E, elev. 700 m, 10 June 2021, Bhavesh Raghoonundon, BR0311; ibid. 18°52'26.8"N, 99°18'15.5"E, elev. 845 m, 10 June 2021, Bhavesh Raghoonundon, BR0313; Chiang Rai Province, Phan District, 19°48'50.0"N, 99°51'57.0"E, elev. 730 m, 22 June 2021, Bhavesh Raghoonundon, BR0368; ibid. 19°48'50.0"N, 99°51'57.0"E, elev. 730 m, 22 June 2021, Bhavesh Raghoonundon, BR0371; Sisaket Province, Kanthararom District, Kok Yang Yai roadside market, 17 September 2016, Santhiti Vadthanarat, SV0345 (CMUB); Ubon Ratchathani Province, Trakan Phuet Phon District, Huay Fai, 15°32'44.3"N, 105°10'17.4"E, elev. 165 m, 28 July 2015, Olivier Raspé, OR1059 (CMUB, BR).

ITS sequence accession number (SV0155): PP354891.

Notes

The BLAST result based on ITS sequence obtained from one of the examined specimens (voucher SV0155, GenBank accession number PP354891) was 100% identical to the holotype of X. sisongkhramensis (voucher BBH 48255, accession number OP462477) which was reported by Tan et al. 2022. This suggested that our collections belonged to X. sisongkhramensis. Morphological characters of our collections mostly fit the original description of the species. However, some variations were observed between ours and the original description as follows: Tan et al. (2022) mentioned the absence of cheilocystidia in X. sisongkhramensis while we could observe them in our collections; they were narrowly fusiform to fusiform with obtuse apex or narrowly utriform, thin-walled. The protologue mentioned broadly clavate to subclavate (40–60 × 8–15 μm) caulocystidia. However, in our observation only groups of rising terminal cells of shape and size similar to the caulocystidia in Tan et al. (2022), were observed. What Tan et al. (2022) considered as caulocystidia were what we described as undifferentiated terminal cells of the stipitipellis. In the species protologue, the pileipellis and stipitipellis were described as composed of thick-walled hyphae (no measurement mentioned). However, only thin-walled hyphae were observed in our collections.

Figure 3. 

Fresh basidiomata of Rostrupomyces sisongkhramensis A OR0915 B OR0919 C OR1004 D SV0155, white pores surface in young basidioma (white arrow) E SV0219 F SV0225. Scale bars: 1 cm (A–F).

Rostrupomyces sisongkhramensis is morphologically similar to Hemileccinum duriusculum Mei-Xiang Li, Zhu L. Yang & G. Wu, which was recently described from China. The two species share some morphological characters including basidiome size and color, scattering of granular squamules on the stipe surface, pale yellow to grayish yellow hymenophore that is depressed around the stipe apex, and unchanging color in any parts. However, H. duriusculum differs by its strikingly venose pileus surface, finer granular squamules on the stipe surface, and subfusiform basidiospores ornamented with irregular warts under SEM (Liu et al. 2024). Rostrupomyces sisongkhramensis is also somewhat similar to a European Leccinum species originally described from Italy, Leccinum albostipitatum den Bakker & Noordel., which has a similar shade of pileus color (light orange), whitish stipe covered with whitish squamules when young to reddish brown in age. However, L. albostipitatum can be differentiated by having an inflexed margin which exceeds the hymenophore by up to 4 mm, yellowish white to very pale brown hymenophore that becomes brownish when bruised, a clear blue discoloration of the stipe base when touched, context staining vinaceous then grayish to blackish when cut, smooth fusiform basidiospores, distribution in Europe, and association with Populus L. trees (den Bakker and Noordeloos 2005).

Figure 4. 

Microscopic features of Rostrupomyces sisongkhramensis A basidiospores B basidia C cheilocystidia D pleurocystidia E pileipellis F stipitipellis showing a cluster of narrowly clavate to clavate cells which slightly scattered on the stipe surface. Scale bars: 10 µm (A–D); 25 µm (D–E); 50 µm (E–F). All line drawings were made from SV0155.

Phylogenetically, R. sisongkhramensis is closely related to Rubinosporus auriporus Vadthanarat, Raspé & Lumyong, the only known species in the genus, which was described from the same region as Rostrupomyces (northern Thailand). However, it can be differentiated from R. sisongkhramensis by having grayish red to pastel red to reddish brown pileus; even stipe surface with scattered bright yellow to yellowish white to orange to light brown minute squamules; shorter tubes especially when young; golden yellow hymenophore; and the striking dark ruby spore print (Vadthanarat et al. 2022).

Figure 5. 

Scanning electron micrographs of basidiospores A–B Rostrupomyces sisongkhramensis (SV0155) C–D Hemileccinum inferius (SV0282).

Hemileccinum inferius Vadthanarat, Raghoonundon & Raspé, sp. nov.

MycoBank No: 849063
Figs 6, 7

Etymology

inferius” refers to the only lower part of the stipe ornamented with reticulum

Holotype

Thailand, Chiang Mai Province, Muang District, Doi Suthep-Pui National Park, 18°47'52.8"N, 98°54'21.2"E, elev. 1,170 m, 1 July 2016, Santhiti Vadthanarat, SV0282 (holotype: CMUB, isotype: BKF, MFUB). ITS sequence accession number PP354892.

Diagnosis

Hemileccinum inferius can be differentiated from resembling Hemileccinum species by a grayish red to reddish brown to dark brown, plane to sub-depressed, subrugulose to pitted pileus; and yellow to yellowish white, cylindrical with subbulbous stipe, with surface even on the upper half and subscabrous to delicately reticulate on the lower half, as well as smooth basidiospores even when observed under SEM.

Description

Basidiomata medium-sized. Pileus 66–68 mm in diameter, plane to sub-depressed at the centre; margin deflexed in age, elastic, slightly exceeding (up to 1 mm); surface subrugulose to pitted especially near the margin, dull, moist to slippery when wet, tomentose, grayish red (8B/C3–4) to reddish brown (8D/E4–6) to dark brown to reddish brown (7–8F4–6), unchanging when bruised; context 8–10 mm thick haft-way to the margin, firm to soft, pale yellow (1A3), slightly brown (7E5) near the pileus surface, light yellow (1A4) above the hymenium in age, unchanging when cut. Stipe 65–76 × 14–18 mm, central, terete, cylindrical with subbulbous base; surface even on the upper half then subscabrous to delicately reticulate on the lower half, dull, dry to moist, light yellow (2A4–6) to yellowish white to pale yellow (2A2–3) at the base, occasionally with reddish brown to dark brown spots (8D5–8, 8F7) at places, minutely covered with pale yellow to light brown to dark brown (2A3–4 to 7D/E4, 7F8) squamules on the upper half, slightly fibrillose following a reticulate pattern at the middle of the stipe getting less so to the base, unchanging when bruised; basal mycelium white (1A1); context solid, firm, pale yellow (2A3–5) especially in the above half near the stipe surface becoming yellowish white (2A2) to off-white at the base, unchanging when cut. Hymenophore tubulate, slightly depressed around the stipe, with slightly decurrent tooth, subventricose. Tubes 7–8 mm long half-way to the margin, yellow to grayish yellow (2A7 to 2B7) near the pileus context then olive (2E5) near the pores, separable from the pileus context, unchanging when bruised. Pores 0.3–0.8(1.2) mm wide at mid-radius, subangular to angular, even, grayish yellow (2B5), unchanging when touched, irregularly arranged; topography subregular. Odour mild fungoid. Taste mild. Spore print olive brown (4E7).

Macrochemical reactions : KOH, brownish orange on pileus and hymenophore, pale orange on pileus context and stipe surface, and stipe context; NH4OH, brownish orange with purple aura on pileus, yellowish to brownish orange with purple aura on stipe surface, yellowish to greenish or slightly blue on pileus context and stipe context.

Figure 6. 

Fresh basidioma of Hemileccinum inferius A, B SV0282 (holotype). Scale bars: 1 cm (A, B).

Spores [118/2/2] (10.5–)11.4–12.9–14.6(–15.3) × (3.8–)4.2–4.8–5.6(–6.1) µm Q = (2.06–)2.4–2.68–3.05(–3.32). From the type (10.8–)11.5–12.7–14.2(–14.5) × (4.1–)4.3–4.8–5.5(–6.1) µm, Q = (2.06–)2.33–2.66–3.06(–3.1), N = 68, narrowly ellipsoid to subcylindrical with a slight suprahilar depression, thin-walled, smooth under light microscope and SEM (Fig. 5C–D), yellowish to brownish hyaline in water, yellowish hyaline in KOH, inamyloid. Basidia 4-spored, (23–)24–27–31(–32) × (11–)11–12–14(–14) µm, clavate without basal clamp connection, hyaline to yellowish hyaline in KOH; sterigmata up to 4 µm long. Cheilocystidia (30–)34–54–72(–72) × (7–)8–10–14(–14) µm, narrowly fusiform with elongated obtuse apex, frequent, thin-walled, hyaline to yellowish hyaline in KOH. Pleurocystidia (34–)34–51–69(–70) × (10–)10–11–13(–13) µm, frequent near the pores, narrowly fusiform with elongated obtuse apex, thin-walled, hyaline to yellowish hyaline in KOH. Hymenophoral trama slightly divergent, 62–150 µm wide composed of cylindrical, 4–12 µm wide hyphae, with subregular mediostratum 30–100 µm wide, hyaline in KOH. Pileipellis a hyphoepithelium, 80–112 μm thick, the pileipellis composed of ellipsoid to broadly ellipsoid or cylindrical, thin-walled, more or less vertically arranged, occasionally branching or anastomosing, with metablematic, elongated-cylindrical hyphae (2–4 µm wide hyphae), branching or anastomosing at places, hyaline to yellowish hyaline in KOH; terminal cells of 2 types: 1) ellipsoid to broadly ellipsoid, 8–15 × 12–20 µm; and 2) clavate to broadly clavate with obtuse apex, 10–20 × 4–7 µm. Pileus context made of moderately interwoven, ellipsoid to broadly ellipsoid, thin-walled hyphae, 10–23 µm wide, hyaline in KOH. Stipitipellis arranged parallel to the surface of the stipe (40–50 µm thick), composed of moderately interwoven, cylindrical, thin-walled, 2.5–4 µm wide hyphae, anastomosing at places, moderately scattered with groups of rising cells to clusters (50–60 µm high) of thin-walled clavate to broadly clavate cells (20–30 × 10–15 µm), hyaline to yellowish hyaline in KOH. Caulocystidia not seen. Stipe context composed of parallel, 8–22 µm wide hyphae, hyaline in KOH. Clamp connections not seen in any tissue.

Figure 7. 

Microscopic features of Hemileccinum inferius A basidiospores B basidia C cheilocystidia D pleurocystidia E pileipellis F stipitipellis showing a cluster of clavate to boardy clavate like cells which moderately scattered on the stipitipellis. Scale bars: 10 µm (A–D); 25 µm (D–E); 50 µm (E–F). All drawings were made from holotype type (SV0282).

Habitat and distribution

Solitary, on loamy soil in hill evergreen forest dominated by Fagaceae scattered with a few Dipterocarpus obtusifolius, at 985–1,170 m elevation. Currently known from Chiang Mai Province, northern Thailand.

Additional specimens examined

Thailand, Chiang Mai Province, Mae Taeng District, 19°06'59"N, 98°44'23"E, elev. 985 m, 6 June 2021, Bhavesh Raghoonundon, BR0260 (MFLU).

Notes

Hemileccinum inferius is described based on collections from Thailand. The comparison of the new species with the seven known Asian species follows. Hemileccinum albidum differs from H. inferius by gray-brown to chrome yellow to ochraceous or golden brown pileus; longer and slender stipe (up to 160 mm); shorter basidiospores (10–12.5 × 4.0–5.5 µm); occurrence at higher elevations (1,968–2,490 m; Li et al. 2021). Hemileccinum brevisporum is similar in pileus color but has shorter basidiospores (9–11 × 4–5 μm); and it occurs under Fagaceae and Pinaceae, at higher elevations (1,700–2,120 m); Li et al. 2021). Hemileccinum duriusculum is macromorphologically quite similar, but differs by a strongly venose pileus surface, even when young, the absence of reticulum on the lower half of the stipe, as well as shorter cheilo- and pleurocystidia (Liu et al. 2024). Hemileccinum ferrugineipes has similar pileus surface and color but can be differentiated by the apparent pale red-brown color on the lower part of the stipe; and shorter basidiospores (11.0–12.5 × 4–5 μm; Li et al. 2021). Hemileccinum indecorum is clearly different in having dark red to reddish brown basidiomata with mucilaginous surface densely covered with whitish to dirty white, small conical to subconical to irregularly shaped squamules; incurved margin; and yellowish hymenophore that slowly turns brownish to reddish brown when bruised (Horak 2011; Zeng et al. 2012). Hemileccinum parvum has smaller basidiomata (pileus 3.3–3.6 cm diam, stipe 60–97 × 4–9 mm); paler pileus (brownish to yellowish); pale yellow context that slowly turns pale blue when cut (Li et al. 2021). Hemileccinum rugosum has paler pileus (light orange to reddish orange); very distinctly rugose to wrinkled pileus surface; and shorter basidiospores (9–13 × 4–5 µm; Wu et al. 2016).

Hemileccinum inferius is also similar to an American species, H. floridanum, which has reddish brown to chestnut brown wrinkled and uneven pileus, whitish to pale yellow stipe, white basal mycelium, yellow hymenophore, and smooth basidiospore under both light microscope and SEM. However, the latter species is different by white context that slowly turns yellow often from the margin toward the center, longer basidiospores (10–17 × 4.5–6 μm), likely forms association with oak in northern America (Farid et al. 2021).

Phylogenetically, H. inferius was most closely related to H. hortonii, H. rugosum, and an undescribed specimen (voucher HKAS 53421) from China. Hemileccinum hortonii, an American species, can easily be distinguished by its conspicuously pitted pileus, smooth to lightly pruinose stipe that sometimes has delicate reticulation on the upper half, pores that occasionally turn blue on when touched, and slightly longer and narrower basidiospores (12–15 × 3.5–4.5; Kuo and Ortiz-Santana 2020; Farid et al. 2021). For morphological comparison with H. rugosum see the above paragraph.

Discussion

In this study, the morphological and phylogenetic evidence highly supported establishing Rostrupomyces as a new genus of Boletaceae to accommodate Xerocomus sisongkhramensis. The most important morphological characters used to differentiate the new genus from other Boletaceae genera are: subscabrous stipe surface with scattered granulose squamules; hymenophore that is white in young basidiomes and becomes yellow in age; yellowish brown spore print; and broadly ellipsoid to ellipsoid basidiospores with smooth surface.

The character of subscabrous to scabrous stipe surface dotted with scattered granulose squamules is also present in other Boletaceae genera such as Hemileccinum (see in notes under Rostrupomyces), Leccinum Gray, Leccinellum Bresinsky & Manfr. Binde, Rugiboletus G. Wu & Zhu L. Yang, and Sutorius Halling, Nuhn & N.A. Fechner. Leccinum can be separated from Rostrupomyces by having a white to pallid to light brown hymenophore while Leccinellum has yellow hymenophore similar to Rostrupomyces. However, both genera are different from Rostrupomyces by a more or less pronounced color change of hymenophore, stipe surface, and/or context, which can stain red, brown, yellow, or occasionally blue when bruised. Leccinellum and Leccinum produce boletoid basidiospores which are also different from Rostrupomyces. Moreover, they are phylogenetically distant and placed in another subfamily, the Leccinoideae (den Bakker and Noordeloos 2005; Wu et al. 2016; Xue et al. 2019; Meng et al. 2021). Rugiboletus differs from Rostrupomyces by its strongly wrinkled pileus (especially when young), yellow or brown or reddish brown hymenophore that is unchanging or turns bluish when bruised, subfusiform basidiospores, and phylogenetically distant and placed in Pulveroboletus group (Wu et al. 2015; Kuo and Ortiz-Santana 2020). Sutorius Halling, Nuhn & N.A. Fechner, is different in having chocolate to reddish brown or purplish brown basidiomata, grayish or reddish brown or brownish orange hymenophore, context always with scattered reddish or violet or dark brown encrustations that are visible with the naked eye, reddish brown spore deposit, and narrowly ellipsoid to subcylindrical basidiospores (Halling et al. 2012; Vadthanarat et al. 2021). Like Rugiboletus, Sutorius is phylogenetically distant from Rostrupomyces, belonging to the Pulveroboletus group (Vadthanarat et al. 2021).

Xerocomoideae genera other than Rostrupomyces also produce smooth basidiospores, including Amylotrama, Aureoboletus, Alessioporus, Pulchroboletus, Rubinosporus, and Veloboletus. Moreover, while most Xerocomus and Phylloporus species produce basidiospores with bacillate surface, a few species produce smooth basidiospores (Neves and Halling 2010; Wu et. al. 2016; Chuankid et al. 2019). However, only Amylotrama and Rubinosporus present the same shape of basidiospore as Rostrupomyces, whereas the others produce more or less oblong to ellipsoid to fusiform basidiospores (Gelardi et al. 2014; Wu et al. 2016; Farid et al. 2017; Frank et al. 2017; Zhang et al. 2019; Crous et al. 2020; Lebel et al. 2022; Vadthanarat et al. 2022). Amylotrama comprises two species from Australia, which are completely different from Rostrupomyces by their sequestrate basidiomata (Lebel et al. 2022). Rubinosporus, differs by having a strikingly thin hymenophore, especially when young; golden yellow hymenophore; and dark ruby spore print (Vadthanarat et al. 2022). Aureoboletus differs by the pileus usually having a viscid surface especially when moist; and golden yellow hymenophore (Wu et al. 201; Zhang et al. 2019). Alessioporus is different by its reticulated stipe occasionally with a granular ring-like zone in the middle or lower half of the stipe, golden yellow hymenophore; blue staining of the stipe surface, hymenophore, and context; and distribution in Mediterranean Italy and subtropical USA (Gelardi et al. 2014; Frank et al. 2017). Pulchroboletus differs by the stipe surface with scattered red to reddish brown, occasionally with reticulum or longitudinal striations, and with a pseudo-annulus; golden yellow hymenophore; intense blue staining of the hymenophore and context; and occurrence only in Mediterranean Europe and tropical to subtropical America (Gelardi et al. 2014; Farid et al. 2017). The only Veloboletus species, is different by its basidiomata with a distinctive universal veil; blue staining of the pileus, stipe, hymenophore, and context, and distribution in Australia (Crous et al. 2020).

Tan et al. (2022) phylogeny was based on ITS and LSU sequences of only Xerocomus spp., and Phylloporus as outgroup, which resulted in the clustering of X. sisongkhramensis in Xerocomus. However, our phylogeny based on multiple protein-coding genes (atp6, cox3, tef1, and rpb2) and on a much wider taxon sampling of Boletaceae resolved X. sisongkhramensis in subfamily Xerocomoideae indeed, but distant from other Xerocomus species. Keeping X. sisongkhramensis would have rendered the genus polyphyletic. The erection of the new genus Rostrupomyces, which can also be morphologically separated from Xerocomus, was therefore necessary.

In the phylogeny, Rostrupomyces appeared sister to another monotypic genus, Rubinosporus (morphological comparison see in notes under Rostrupomyces sisongkhramensis). The two genera can be differentiated mainly by the spore print color, and color of hymenophore, two characters that do not vary between species in the same genus in Boletaceae. The characters have been primarily used to differentiate many genera in Boletaceae e.g., Sutorius, Cacaoporus, Hourangia, Baorangia (Halling et al. 2012; Wu et al. 2015; Zhu et al. 2015; Vadthanarat et al. 2019b). Additional morphological characters, including pileus color and stipe surface, could be useful to separate them. However, both genera so far comprise only a single species and the pileus color and stipe surface are found to be variable between species within the same genus. For example, in Boletus L. and Tylopilus P. Karst. the pileus color is variable from white, yellow, brown, orange, green, gray, and purple, and the stipe surface from even to reticulate to strongly reticulate (e.g., Cui et al. 2015; Wu et al. 2016; Li and Yang 2021). Hence, if more species in either of those two genera are described, the comparison between the two genera might need updating.

Rostrupomyces has been found so far on sandy loam to sandy clay loam soils at elevations lower than 1,000 m (165 to 915 m), in open dry dipterocarp and dipterocarp forest mainly dominated by ectomycorrhizal trees in family Dipterocarpaceae genera Anthoshorea (A. roxburghii), Dipterocarpus (D. obtusifolius, D. tuberculatus, D. intricatus), Pentacme (P. siamensis), and Shorea (S. obtusa), with scattered Fagaceae trees. In Thailand the Dipterocarpaceae tree species are mainly distributed in lowland (<800 m) to mid-elevation forests (800–1,200 m) whereas Fagaceae trees are mostly distribute in mid-elevation to highland forests (>1,200 m) (Gardner et al. 2007). During our surveys on the diversity of Boletaceae in Thailand, no Rostrupomyces collection was found in the forests above 1,000 m, where no Anthoshorea, Dipterocarpus, Pentacme, or Shorea trees were observed or mentioned as occurring. This suggests that the distribution of Rostrupomyces depends on the distribution of the mentioned tree genera, and they are inferred as the associated tree hosts of Rostrupomyces. However, a more detailed study is needed to confirm the specificity of its relationship with ectomycorrhizal hosts.

In this study, some specimens of Rostrupomyces sisongkhramensis were collected from community forests and the species was found to be consumed by local people, in Ubon Ratchathani and Sisaket provinces in lower northeastern Thailand. It is found on sale on roadsides and local markets, along with other Boletaceae in genera such as Baorangia, Boletus, Boletellus, Heimioporus, Retiboletus, Sutorius, and Tylopilus. The species is called “Hed Phueng Waan” in which the words “Hed Phueng” refer to bolete and “Waan” means sweet. It can also be called “Hed Phueng Kaw” in which the words “Kaw” means rice. The same local names are also applied to the other bolete species that are mostly white and have sweet taste after cooking such as Boletus spp. In this region a local name can be used for different mushroom species which present similarly striking morphological characters. Conversely, one mushroom species may have more than one local name. Rostrupomyces sisongkhramensis is also found in the northern parts of Thailand in Chiang Rai and Chiang Mai provinces. However, during our survey in the region, the species has never been found being collected or on sale for consumption by the locals. The protologue of this species (collections from upper northeastern Thailand) did not mention the edibility (Tan et al. 2022).

To date, 15 Hemileccinum species have been described worldwide, among which eight are originally from Asia (China: H. albidum, H. brevisporum, H. duriusculum, H. ferrugineipes, H. parvum, H. rugosum; Singapore: H. indecorum; and Thailand: H. inferius), two species from France in Europe (H. depilatum and H. impolitum), four species from North America (H. floridanum, H. hortonii, H. rubropunctum, and H. subglabripes), and a single species, H. brunneotomentosum, from Belize in Central America (Šutara 2008; Halling et al. 2015; Wu et al. 2016; Kuo and Ortiz-Santana 2020; Nitson and Frank 2020; Farid et al. 2021; Li et al. 2021; Liu et al. 2024). Three Hemileccinum species have been previously reported to occur in Thailand, namely H. depilatum (reported as Boletus depilatus Redeuilh), H. impolitum (reported as B. impolitus Fr.), and H. indecorum (Chandrasrikul et al. 2011; Vadthanarat et al. 2019b). The first two species were originally described from France and were then reported from Thailand based on morphological identification only. As we know that the distribution of Boletaceae species depends on the distribution of their hosts, the ecology and host specificity are important characters in distinguishing species in Boletaceae (den Bakker et al. 2004; Dentinger et al. 2010; Cui et al. 2015; Loizides et al. 2019; Gelardi 2020). It is therefore doubtful that European species are also present in Southeast Asia where the forests are dominated by different tree species or families. Unfortunately, no specimens associated with the reports of occurrence in Thailand are available for molecular analysis to compare with European specimens. Moreover, molecular analysis of several Hemileccinum specimens obtained in our study showed none of them belong to those European species. It is therefore reasonable to assume that the identifications of the Thai collections as H. depilatum and H. impolitum were not correct. The other recorded species, H. indecorum was originally described from Singapore in Southeast Asia (Horak 2011). Specimens collected from Thailand were identified based on both molecular and morphological evidences (Vadthanarat et al. 2019b). However, the full morphological description of this Thai collection has not yet been published. In the future, more detail on the species and more records of Hemileccinum will be reported.

Basidiospores with tiny warts and pinholes (when observed under SEM) are typical of Hemileccinum. However, a few Hemileccinum species produce basidiospores with smooth surface, including the new species (Kuo and Ortiz-Santana et al. 2020; Farid et al. 2021). This kind of exception is also found in other Xerocomoideae genera i.e., in Phylloporus and Xerocomus, In the latter two genera, most of the species produce basidiospores with bacillate surfaces, but a few produce smooth basidiospores (Neves and Halling 2010; Wu et. al. 2016; Chuankid et al. 2019).

A total of 39 new taxa (4 new genera and 35 new species), including those introduced in this paper, have been originally described from Thailand (Rostrup 1902; Yang et al. 2006; Desjardin et al. 2009; Choeyklin et al. 2012; Neves et al. 2012; Halling et al. 2014; Raspé et al. 2016; Vadthanarat et al. 2018; Chuankid et al. 2019; Phookamsak et al. 2019; Vadthanarat et al. 2019a, 2019b, 2020; Chuankid et al. 2021; Raghoonundon et al. 2021; Vadthanarat et al. 2021; Tan et al. 2022; Vadthanarat et al. 2022; This study). Our study on the diversity of Boletaceae in Thailand is still ongoing and is needed to uncover more new taxa and new distribution records for Thailand.

Acknowledgements

Authors are grateful for the permit number 0907.4/4769 granted by the Department of National Parks, Wildlife and Plant Conservation, Ministry of Natural Resources and Environment for collecting in Doi Suthep-Pui National Park.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research work was supported by a Postdoctoral Fellowship from Mae Fah Luang University to Santhiti Vadthanarat, and partially funded by the Mae Fah Luang University research grant 641A01003.

Author contributions

Conceptualization: SV, OR. Data curation: BR, SV. Formal analysis: SV. Funding acquisition: OR. Investigation: SV. Methodology: SV. Project administration: OR. Resources: BR, OR, SV. Software: SV. Supervision: OR. Validation: SL, OR. Visualization: SV. Writing – original draft: SV. Writing – review and editing: BR, OR, SL.

Author ORCIDs

Santhiti Vadthanarat https://orcid.org/0000-0002-9035-0375

Bhavesh Raghoonundon https://orcid.org/0000-0001-6671-2404

Olivier Raspé https://orcid.org/0000-0002-8426-2133

Data availability

All of the data that support the findings of this study are available in the main text.

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