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Research Article
Two new Erythrophylloporus species (Boletaceae) from Thailand, with two new combinations of American species
expand article infoSanthiti Vadthanarat‡, Mario Amalfi§, Roy E. Halling|, Victor Bandala, Saisamorn Lumyong#‡, Olivier Rasp駤
‡ Chiang Mai University, Chiang Mai, Thailand
§ Botanic Garden Meise, Meise, Belgium
| New York Botanical Garden, New York, United States of America
¶ Red Biodiversidad y Sistemática, Veracruz, Mexico
# Academy of Science, The Royal Society of Thailand, Bangkok, Thailand
¤ Service général de l’Enseignement universitaire et de la Recherche scientifique, Bruxelles, Belgium

Corresponding author: Olivier Raspé ( Olivier.Ras@mfu.ac.th )

Academic editor: Alfredo Vizzini
© 2019 Santhiti Vadthanarat, Mario Amalfi, Roy E. Halling, Victor Bandala, Saisamorn Lumyong, Olivier Raspé.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Vadthanarat S, Amalfi M, Halling RE, Bandala V, Lumyong S, Raspé O (2019) Two new Erythrophylloporus species (Boletaceae) from Thailand, with two new combinations of American species. MycoKeys 55: 29-57. https://doi.org/10.3897/mycokeys.55.34570
Open Access

Abstract

Erythrophylloporus is a lamellate genus in the family Boletaceae that has been recently described from China based on E. cinnabarinus, the only known species. Typical characters of Erythrophylloporus are reddish-orange to yellowish-red basidiomata, including lamellae, bright yellow basal mycelium and smooth, broadly ellipsoid, ellipsoid to nearly ovoid basidiospores. During our survey on diversity of Boletaceae in Thailand, several yellowish-orange to reddish- or brownish-orange lamellate boletes were collected. Based on both morphological evidence and molecular analyses of a four-gene dataset (atp6, tef1, rpb2 and cox3), they were recognised as belonging in Erythrophylloporus and different from the already known species. Two new species, E. paucicarpus and E. suthepensis are therefore introduced from Thailand with detailed descriptions and illustrations. Moreover, two previously described Phylloporus species, P. aurantiacus and P. fagicola, were also revised and recombined in Erythrophylloporus. A key to all known Erythrophylloporus species is provided.

Keywords

atp6, cox3, Taxonomy, Phylloporus, Pulveroboletus group, multigene phylogeny, Boletales, Southeast Asia

Introduction

Most fungi in the family Boletaceae are pileate-stipitate with poroid hymenophore, but some have a lamellate hymenophore. Lamellate Boletaceae are currently classified in four genera, Phylloporus Quél, which contains about 84 species worldwide, Phylloboletellus Singer from South America and Mexico, the two recently described genera Phylloporopsis Angelini et al., from the New World and Erythrophylloporus Ming Zhang & T.H. Li from Asia, each of which circumscribes only one species (http://www.indexfungorum.org, Farid et al. 2018; Zhang and Li 2018).

The genus Erythrophylloporus was recently described from China, with E. cinnabarinus Ming Zhang & T.H. Li as the type species. According to Zhang & Li (2018), the typical characters of the genus are orange to reddish-orange basidiomes, reddish-orange to yellowish-red lamellae turning greyish-green when bruised, bright yellow to orange yellow context staining blackish-blue to dark blue when exposed, bright yellow basal mycelium, smooth and broadly ellipsoid to nearly ovoid basidiospores and yellowish-brown pigmented cystidia. During our survey on the diversity of Boletaceae in Thailand, several collections of lamellate boletes were discovered. Some collections were recognised to belong to Erythrophylloporus by possessing yellowish-orange to deep orange to reddish-orange basidiomata with bright yellow basal mycelium and smooth basidiospores. We also found that two described Phylloporus species, P. aurantiacus Halling & G.M. Mueller from Costa Rica and P. fagicola Montoya & Bandala from Mexico (Halling et al. 1999, Montoya and Bandala 2011), share similar morphological characters with the genus Erythrophylloporus, but until now, have not been included in a molecular phylogeny. In this study, a combination of phylogenetic and morphological evidence indicated that our Thai collections were new species, that, together with the two aforementioned American Phylloporus species, belong in Erythrophylloporus. Therefore, we introduce two new species with detailed descriptions and illustrations and propose two new combinations. As some of the species we studied have some characters that do not fit with the protologue of the genus, we emend its description.

Materials and methods

Specimen collecting

Specimens were obtained and photographed from community forests and Doi Suthep-Pui National Park, Chiang Mai Province, northern Thailand during the rainy season in 2015 to 2016. The specimens were wrapped in aluminium foil and taken to the laboratory. After description of macroscopic characters, all specimens were dried in an electric drier at 45–50 °C. Examined specimens were deposited in the herbaria CMUB, MFLU, BKF or BR (Index Herbariorum; Thiers, continuously updated).

Morphological studies

Macroscopic descriptions were made based on detailed field notes and photos of fresh basidiomata. Colour codes follow Kornerup and Wanscher (1978). Macrochemical reactions (colour reactions) of fresh basidiomata were determined using 10% potassium hydroxide (KOH) and 28–30% ammonium hydroxide (NH4OH) in water. Microscopic structures were observed from dried specimens mounted in 5% KOH, NH4OH, Melzer’s reagent or 1% ammoniacal Congo red. A minimum of 50 basidiospores, 20 basidia and 20 cystidia were randomly measured at 1000× with a calibrated ocular micrometer using an Olympus CX51 microscope. The notation ‘[m/n/p]’ represents the number of basidiospores m measured from n basidiomata of p collections. Dimensions of microscopic structures are presented in the following format: (a–)b–c–d (–e), in which c represents the average, b the 5th percentile, d the 95th percentile and a and e the minimum and maximum values, respectively. Q, the length/width ratio, is presented in the same format. A section of the pileus surface was radially and perpendicularly cut at a point halfway between the centre and margin of the pileus. Sections of stipitipellis were taken from halfway up the stipe and longitudinally cut, perpendicularly to the surface. All microscopic features were drawn by free hand using an Olympus Camera Lucida model U−DA, fitted to the microscope cited above. For scanning electron microscopy (SEM), a spore print was mounted on to a SEM stub with double-sided tape. The sample was coated with gold, examined and photographed with a JEOL JSM–5910 LV SEM (JEOL, Japan).

DNA isolation, PCR amplification and DNA sequencing

Genomic DNA was extracted from fresh tissue preserved in CTAB or about 10–15 mg of dried specimens using a CTAB isolation procedure adapted from Doyle and Doyle (1990). Portions of the genes atp6, tef1, rpb2 and cox3 were amplified by the polymerase chain reaction (PCR) technique. The tailed primers ATP6-1M40F and ATP6-2M (Raspé et al. 2016) and the primer pairs EF1-983F/EF1-2218R (Rehner and Buckley 2005) and bRPB2-6F/bRPB2-7.1R (Matheny 2005) were used to amplify atp6, tef1 and rpb2, respectively. PCR conditions were the same as in Raspé et al. (2016). Part of the mitochondrial gene cox3 was amplified with the primers COX3M1-F and COX3M1-R (Vadthanarat et al. 2019), using KAPA2G™ Robust HotStart polymerase (Kapa Biosystems, Wilmington, MA, USA) and the following PCR programme: 2 min 30 s at 95 °C; 35 cycles of 25 s at 95 °C, 30 s at 48 °C, 30 s at 72 °C; 3 min at 72 °C. 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. Sequencing was performed by Macrogen Inc. (Korea and The Netherlands) 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 sequences were assembled in GENEIOUS Pro v. 6.0.6 (Biomatters) and introns were removed prior to alignment, based on the amino acid sequence of previously published sequences. All sequences, including sequences from GenBank, were aligned using MAFFT version 7 (Katoh and Standley 2013) on the server accessed at http://mafft.cbrE.jp/alignment/server/.

Maximum Likelihood (ML) phylogenetic tree inference was performed using RAxML-HPC2 version 8.2.10 (Stamatakis 2006) on the CIPRES web portal (Miller et al. 2009). The phylogenetic tree was inferred from a four-partitions combined dataset, using the GTRCAT model with 25 categories. Two Buchwaldoboletus and nine Chalciporus species from subfamily Chalciporoideae were used as the outgroup. Statistical support of clades was obtained with 1,000 rapid bootstrap replicates.

For Bayesian Inference (BI), the best-fit model of substitution amongst those implementable in MrBayes was estimated separately for each gene using jModeltest (Darriba et al. 2012) on the CIPRES portal, based on the Bayesian Information Criterion (BIC). The selected models were GTR+I+G for atp6 and cox3, SYM+I+G for tef1 and K80+I+G for rpb2. Partitioned Bayesian analysis was performed with MrBayes 3.2 (Ronquist et al. 2012) on the CIPRES portal. Two runs of five chains were run for 15,000,000 generations and sampled every 1,000 generations. The chain temperature was decreased to 0.02 to improve convergence. At the end of the run, the average deviation of split frequencies was 0.007058 and the Potential Scale Reduction Factor (PSRF) values of all parameters were close to 1. The burn-in phase (25%) was estimated by checking the stationarity in the plot generated by the sump command.

Results

Phylogenetic analysis

Twenty-five sequences were newly generated and deposited in GenBank (Table 1). The sequences from three specimens, OR0689, OR1135 (E. paucicarpus) and OR0615B (E. suthepensis), were not included in our phylogenic analyses because they were identical to the sequences of the type specimens of E. paucicarpus and E. suthepensis. The alignment contained 906 sequences (179 for atp6, 313 for tef1, 279 for rpb2, 135 for cox3) from 315 voucher specimens and was 2946 characters long (TreeBase number 24078). ML and BI trees showed similar topologies without any supported conflict (Bootstrap Support values, BS ≥ 70% and posterior probabilities, PP ≥ 0.90; Fig. 1). The four-gene phylogram indicated that the included taxa formed seven major clades, representing the Austroboletoideae, Boletoideae, Chalciporoideae, Leccinoideae, Xerocomoideae, Zangioideae and the Pulveroboletus group. Erythrophylloporus cinnabarinus (typus generis) grouped with the two new Erythrophylloporus species, E. paucicarpus and E. suthepensis, in a highly supported clade (BS = 100% and PP = 1). The two New World Phylloporus species (P. aurantiacus voucher REH7271 and P. fagicola voucher Garay215) also clustered in the Erythrophylloporus clade indicating that they are close relatives. Erythrophylloporus formed a clade sister to the genus Singerocomus T.W. Henkel & M.E. Sm. with high Bootstrap support (96%) but low posterior probability support (0.86) within the Pulveroboletus group. Some undescribed species formed two different generic clades in the Pulveroboletus group. Boletus p.p. spp. clade 1 contains two specimens, HKAS63598 and HKAS9660, named “Boletus sp.” in Wu et al. (2016), as well as two of our specimens, OR0832 and OR1002. Boletus p.p. sp. clade 2 contains a single African specimen, JD0693, sister to and morphologically different from Cyanoboletus.

Figure 1. 

Phylogenetic tree inferred from the four-gene dataset (atp6, rpb2, tef1 and cox3), including Erythrophylloporus species and selected Boletaceae using Maximum Likelihood and Bayesian Inference methods (ML tree is presented). The two Buchwaldoboletus and nine Chalciporus species in subfamily Chalciporoideae were used as outgroup. Most of the taxa not belonging to the Pulveroboletus group were collapsed into subfamilies. All generic clades, including one undescribed generic clade in Pulveroboletus group that were highly supported, were also collapsed. Bootstrap support values (BS ≥ 70%) and posterior probabilities (PP ≥ 0.90) are shown above the supported branches.

Table 1.
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List of collections used in this study, with origin and GenBank accession numbers. Newly generated sequences are presented in bold.

Species Voucher Origin atp6 tef1 rpb2 cox3 References
Afroboletus aff. multijugus JD671 Burundi MH614651 MH614700 MH614747 MH614794 Vadthanarat et al. 2019
Afroboletus costatisporus ADK4644 Togo KT823958 KT824024 KT823991 MH614795* Raspé et al. 2016; *Vadthanarat et al. 2019
Afroboletus luteolus ADK4844 Togo MH614652 MH614701 MH614748 MH614796 Vadthanarat et al. 2019
Aureoboletus catenarius HKAS54467 China – KT990711 KT990349 – Wu et al. 2016
Aureoboletus duplicatoporus HKAS50498 China – KF112230 KF112754 – Wu et al. 2014
Aureoboletus gentilis ADK4865 Belgium KT823961 KT824027 KT823994 MH614797* Raspé et al. 2016; *Vadthanarat et al. 2019
Aureoboletus mirabilis HKAS57776 China – KF112229 KF112743 – Wu et al. 2014
Aureoboletus moravicus VDKO1120 Belgium MG212528 MG212573 MG212615 MH614798* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Aureoboletus nephrosporus HKAS67931 China – KT990720 KT990357 – Wu et al. 2016
Aureoboletus projectellus AFTOL-ID-713 USA DQ534604* AY879116 AY787218 – *Binder & Hibbett 2006; Binder, Matheny & Hibbett, Unpublished
Aureoboletus shichianus HKAS76852 China – KF112237 KF112756 – Wu et al. 2014
Aureoboletus sp. HKAS56317 China – KF112239 KF112753 – Wu et al. 2014
Aureoboletus sp. OR0245 China MH614653 MH614702 MH614749 MH614799 Vadthanarat et al. 2019
Aureoboletus sp. OR0369 Thailand MH614654 MH614703 MH614750 MH614800 Vadthanarat et al. 2019
Aureoboletus thibetanus HKAS76655 China – KF112236 KF112752 – Wu et al. 2014
Aureoboletus thibetanus AFTOL-ID-450 China DQ534600* DQ029199 DQ366279 – *Binder and Hibbett 2006; Unpublished
Aureoboletus tomentosus HKAS80485 China – KT990715 KT990353 – Wu et al. 2016
Aureoboletus viscosus OR0361 Thailand MH614655 MH614704 MH614751 MH614801 Vadthanarat et al. 2019
Aureoboletus zangii HKAS74766 China – KT990726 KT990363 – Wu et al. 2016
Austroboletus cf. dictyotus OR0045 Thailand KT823966 KT824032 KT823999 MH614802* Raspé et al. 2016; *Vadthanarat et al. 2019
Austroboletus cf. subvirens OR0573 Thailand MH614656 MH614705 MH614752 MH614803 *Vadthanarat et al. 2019
Austroboletus eburneus REH9487 Australia – JX889708 – – Halling et al. 2012b
Austroboletus olivaceoglutinosus HKAS57756 China – KF112212 KF112764 – Wu et al. 2014
Austroboletus sp. HKAS59624 China – KF112217 KF112765 – Wu et al. 2014
Austroboletus sp. OR0891 Thailand MH614657 MH614706 MH614753 MH614804 Vadthanarat et al. 2019
Baorangia pseudocalopus HKAS63607 China – KF112167 KF112677 – Wu et al. 2014
Baorangia pseudocalopus HKAS75739 China – KJ184570 KM605179 – Wu et al. 2015
Baorangia pseudocalopus HKAS75081 China – KF112168 KF112678 – Wu et al. 2014
Baorangia rufomaculata BOTH4144 USA MG897415 MG897425 MG897435 MH614805* Phookamsak et al. 2019; *Vadthanarat et al. 2019
Baorangia major OR0209 Thailand MG897421 MG897431 MG897441 MK372295* Phookamsak et al. 2019; *Vadthanarat et al. 2019
Boletellus aff. ananas NY815459 Costa Rica – KF112308 KF112760 – Wu et al. 2014
Boletellus aff. emodensis OR0061 Thailand KT823970 KT824036 KT824003 MH614806* Raspé et al. 2016; *Vadthanarat et al. 2019
Boletellus ananas K(M)123769 Belize MH614658 MH614707 MH614754 MH614807 Vadthanarat et al. 2019
Boletellus sp. OR0621 Thailand MG212529 MG212574 MG212616 MH614808* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Boletellus sp. HKAS58713 China – KF112307 KF112759 – Wu et al. 2014
Boletellus sp. HKAS59536 China – KF112306 KF112758 – Wu et al. 2014
Boletus aereus VDKO1055 Belgium MG212530 MG212575 MG212617 MH614809* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Boletus albobrunnescens OR0131 Thailand KT823973 KT824039 KT824006 MH614810* Raspé et al. 2016; *Vadthanarat et al. 2019
Boletus botryoides HKAS53403 China – KT990738 KT990375 – Wu et al. 2016
Boletus edulis HMJAU4637 Russia – KF112202 KF112704 – Wu et al. 2014
Boletus edulis VDKO0869 Belgium MG212531 MG212576 MG212618 MH614811* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Boletus p.p. sp JD0693 Burundi MH645583 MH645591 MH645599 – Vadthanarat et al. 2019
Boletus p.p. sp. OR0832 Thailand MH645584 MH645592 MH645600 MH645605 Vadthanarat et al. 2019
Boletus p.p. sp. OR1002 Thailand MH645585 MH645593 MH645601 MH645606 Vadthanarat et al. 2019
Boletus pallidus BOTH4356 USA MH614659 MH614708 – MH614812 Vadthanarat et al. 2019
Boletus pallidus TDB-1231-Bruns – AF002142 – – AF002154 Kretzer and Bruns 1999
Boletus reticuloceps HKAS57671 China – KF112201 KF112703 – Wu et al. 2014
Boletus s.s. sp. OR0446 China MG212532 MG212577 MG212619 MH614813* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Boletus sp. HKAS59660 China – KF112153 KF112664 – Wu et al. 2014
Boletus sp. HKAS63598 China – KF112152 KF112663 – Wu et al. 2014
Boletus violaceofuscus HKAS62900 China – KF112219 KF112762 – Wu et al. 2014
Borofutus dhakanus HKAS73789 Bangladesh – JQ928576 JQ928597 – Hosen et al. 2013
Borofutus dhakanus OR0345 Thailand MH614660 MH614709 MH614755 MH614814 Vadthanarat et al. 2019
Buchwaldoboletus lignicola HKAS76674 China – KF112277 KF112819 – Wu et al. 2014
Buchwaldoboletus lignicola VDKO1140 Belgium MH614661 MH614710 MH614756 MH614815 Vadthanarat et al. 2019
Butyriboletus appendiculatus VDKO0193b Belgium MG212537 MG212582 MG212624 MH614816* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Butyriboletus cf. roseoflavus OR0230 China KT823974 KT824040 KT824007 MH614819* Raspé et al. 2016; *Vadthanarat et al. 2019
Butyriboletus frostii NY815462 USA – KF112164 KF112675 – Wu et al. 2014
Butyriboletus pseudoregius VDKO0925 Belgium MG212538 MG212583 MG212625 MH614817* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Butyriboletus pseudospeciosus HKAS63513 China – KT990743 KT990380 – Wu et al. 2016
Butyriboletus roseoflavus HKAS54099 China – KF739779 KF739703 – Wu et al. 2014
Butyriboletus roseopurpureus BOTH4497 USA MG897418 MG897428 MG897438 MH614818* Phookamsak et al. 2019; *Vadthanarat et al. 2019
Butyriboletus sp. HKAS52525 China – KF112163 KF112671 – Wu et al. 2014
Butyriboletus sp. HKAS59814 China – KF112199 KF112699 – Wu et al. 2014
Butyriboletus sp. HKAS57774 China – KF112155 KF112670 – Wu et al. 2014
Butyriboletus subsplendidus HKAS50444 China – KT990742 KT990379 – Wu et al. 2016
Butyriboletus yicibus HKAS55413 China – KF112157 KF112674 – Wu et al. 2014
Caloboletus calopus ADK4087 Belgium MG212539 KJ184566 KP055030 MH614820 Vadthanarat et al. 2018; Zhao et al. 2014a; Zhao et al. 2014b; Vadthanarat et al. 2019
Caloboletus inedulis BOTH3963 USA MG897414 MG897424 MG897434 MH614821* Phookamsak et al. 2019; *Vadthanarat et al. 2019
Caloboletus panniformis HKAS55444 China – KF112165 KF112666 – Wu et al. 2014
Caloboletus radicans VDKO1187 Belgium MG212540 MG212584 MG212626 MH614822* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Caloboletus sp. HKAS53353 China – KF112188 KF112668 – Wu et al. 2014
Caloboletus sp. OR0068 Thailand MH614662 MH614711 MH614757 MH614823 Vadthanarat et al. 2019
Caloboletus yunnanensis HKAS69214 China – KJ184568 KT990396 – Zhao et al. 2014a; Wu et al. 2016
Chalciporus aff. piperatus OR0586 Thailand KT823976 KT824042 KT824009 MH614824* Raspé et al. 2016; *Vadthanarat et al. 2019
Chalciporus aff. rubinus OR0139 China MH614663 MH614712 MH614758 – Vadthanarat et al. 2019
Chalciporus africanus JD517 Cameroon KT823963 KT824029 KT823996 MH614825* Raspé et al. 2016; *Vadthanarat et al. 2019
Chalciporus piperatus VDKO1063 Belgium MH614664 MH614713 MH614759 MH614826 Vadthanarat et al. 2019
Chalciporus rubinus AF2835 Belgium KT823962 KT824028 KT823995 – Raspé et al. 2016
Chalciporus sp. HKAS53400 China – KF112279 KF112821 – Wu et al. 2014
Chalciporus sp. HKAS74779 China – KF112278 KF112820 – Wu et al. 2014
Chalciporus sp. OR0363 Thailand MH645586 MH645594 MH645602 MH645607 Vadthanarat et al. 2019
Chalciporus sp. OR0373 Thailand MH645587 MH645595 MH645603 MH645608 Vadthanarat et al. 2019
Chiua sp. OR0141 China MH614665 MH614714 MH614760 MH614827 Vadthanarat et al. 2019
Chiua virens HKAS76678 China – KF112272 KF112793 – Wu et al. 2014
Chiua virens OR0266 China MG212541 MG212585 MG212627 MH614828* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Chiua viridula HKAS74928 China – KF112273 KF112794 – Wu et al. 2014
Crocinoboletus cf. laetissimus OR0576 Thailand KT823975 KT824041 KT824008 MH614833* Raspé et al. 2016; *Vadthanarat et al. 2019
Crocinoboletus rufoaureus HKAS53424 China – KF112206 KF112710 – Wu et al. 2014
Cyanoboletus brunneoruber OR0233 China MG212542 MG212586 MG212628 MH614834* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Cyanoboletus instabilis HKAS59554 China – KF112186 KF112698 – Wu et al. 2014
Cyanoboletus pulverulentus RW109 Belgium KT823980 KT824046 KT824013 MH614835* Raspé et al. 2016; *Vadthanarat et al. 2019
Cyanoboletus sinopulverulentus HKAS59609 China – KF112193 KF112700 – Wu et al. 2014
Cyanoboletus sp. OR0257 China MG212543 MG212587 MG212629 MH614836* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Cyanoboletus sp. HKAS76850 China – KF112187 KF112697 – Wu et al. 2014
Cyanoboletus sp. OR0322 Thailand MH614673 MH614722 MH614768 MH614837 Vadthanarat et al. 2019
Cyanoboletus sp. OR0491 China MH614674 MH614723 MH614769 MH614838 Vadthanarat et al. 2019
Cyanoboletus sp. OR0961 Thailand MH614675 MH614724 MH614770 MH614839 Vadthanarat et al. 2019
Erythrophylloporus aurantiacus REH7271 Costa Rica MH614666 MH614715 MH614761 MH614829 This study
Erythrophylloporus cinnabarinus GDGM70536 China – MH378802 MH374035 – Zhang and Li 2018
Erythrophylloporus fagicola Garay215 Mexico MH614667 MH614716 MH614762 MH614830 This study
Erythrophylloporus paucicarpus OR1151 Thailand MH614670 MH614719 MH614765 MH614831 This study
Erythrophylloporus paucicarpus OR0689 Thailand MH614668 MH614717 MH614763 – This study
Erythrophylloporus paucicarpus OR1135 Thailand MH614669 MH614718 MH614764 – This study
Erythrophylloporus suthepensis SV0236 Thailand MH614672 MH614721 MH614767 MH614832 This study
Erythrophylloporus suthepensis OR0615B Thailand MH614671 MH614720 MH614766 – This study
Fistulinella prunicolor REH9880 Australia MH614676 MH614725 MH614771 MH614840 Vadthanarat et al. 2019
Fistulinella prunicolor REH9502 Australia MG212544 MG212588 MG212630 – Vadthanarat et al. 2018
Gymnogaster boletoides NY01194009 Australia – KT990768 KT990406 – Wu et al. 2016
Harrya atriceps REH7403 Costa Rica – JX889702 – – Halling et al. 2012b
Harrya chromapes HKAS50527 China – KF112270 KF112792 – Wu et al. 2014
Harrya chromapes HKAS49416 China HQ326840 HQ326863 – – Li et al. 2011
Harrya moniliformis HKAS49627 China – KT990881 KT990500 – Wu et al. 2016
Heimioporus cf. mandarinus OR0661 Thailand MG212545 MG212589 MG212631 MH614841* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Heimioporus japonicus OR0114 Thailand KT823971 KT824037 KT824004 MH614842* Raspé et al. 2016; *Vadthanarat et al. 2019
Heimioporus retisporus HKAS52237 China – KF112228 KF112806 – Wu et al. 2014
Heimioporus sp. OR0218 Thailand MG212546 MG212590 MG212632 – Vadthanarat et al. 2018
Hemileccinum depilatum AF2845 Belgium MG212547 MG212591 MG212633 MH614843* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Hemileccinum impolitum ADK4078 Belgium MG212548 MG212592 MG212634 MH614844* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Hemileccinum indecorum OR0863 Thailand MH614677 MH614726 MH614772 MH614845 Vadthanarat et al. 2019
Hemileccinum rugosum HKAS84970 China – KT990773 KT990412 – Wu et al. 2016
Hortiboletus amygdalinus HKAS54166 China – KT990777 KT990416 – Wu et al. 2016
Hortiboletus rubellus VDKO0403 Belgium MH614679 – MH614774 MH614847 *Vadthanarat et al. 2019
Hortiboletus sp. HKAS51239 China – KF112184 KF112695 – Wu et al. 2014
Hortiboletus sp. HKAS50466 China – KF112183 KF112694 – Wu et al. 2014
Hortiboletus sp. HKAS51292 China – KF112181 KF112692 – Wu et al. 2014
Hortiboletus sp. HKAS76673 China – KF112182 KF112693 – Wu et al. 2014
Hortiboletus subpaludosus HKAS59608 China – KF112185 KF112696 – Wu et al. 2014
Hourangia cf. pumila OR0762 Thailand MH614680 MH614728 MH614775 MH614848 Vadthanarat et al. 2019
Hourangia cheoi HKAS74744 China – KF112285 KF112772 – Wu et al. 2014
Hourangia cheoi Zhu108 China – KP136979 KP136928 – Zhu et al. 2015
Hourangia nigropunctata HKAS 57427 China – KP136927 KP136978 – Zhu et al. 2015
Hymenoboletus luteopurpureus HKAS46334 China – KF112271 KF112795 – Wu et al. 2014
Imleria badia VDKO0709 Belgium KT823983 KT824049 KT824016 MH614849* Raspé et al. 2016; *Vadthanarat et al. 2019
Imleria obscurebrunnea OR0263 China MH614681 MH614729 MH614776 MH614850 Vadthanarat et al. 2019
Imleria subalpina HKAS74712 China – KF112189 KF112706 – Wu et al. 2014
Lanmaoa angustispora HKAS74759 China – KM605155 KM605178 – Wu et al. 2015
Lanmaoa angustispora HKAS74765 China – KF112159 KF112680 – Wu et al. 2014
Lanmaoa asiatica HKAS54094 China – KF112161 KF112682 – Wu et al. 2014
Lanmaoa asiatica HKAS63603 China – KM605153 KM605176 – Wu et al. 2015
Lanmaoa asiatica OR0228 China MH614682 MH614730 MH614777 MH614851 Vadthanarat et al. 2019
Lanmaoa carminipes BOTH4591 USA MG897419 MG897429 MG897439 MH614852* Phookamsak et al. 2019, *Vadthanarat et al. 2019
Lanmaoa flavorubra NY775777 Costa Rica – KF112160 KF112681 – Wu et al. 2014
Lanmaoa pallidorosea BOTH4432 USA MG897417 MG897427 MG897437 MH614853* Phookamsak et al. 2019, *Vadthanarat et al. 2019
Lanmaoa sp. HKAS52518 China – KF112162 KF112683 – Wu et al. 2014
Lanmaoa sp. OR0130 Thailand MH614683 MH614731 MH614778 MH614854 Vadthanarat et al. 2019
Lanmaoa sp. OR0370 Thailand MH614684 MH614732 MH614779 MH614855 Vadthanarat et al. 2019
Leccinellum aff. crocipodium HKAS76658 China – KF112252 KF112728 – Wu et al. 2014
Leccinellum aff. griseum KPM-NC-0017832 Japan KC552164 JN378450* – – unpublished, *Orihara et al. 2012
Leccinellum corsicum Buf4507 USA – KF030435 – – Nuhn et al. 2013
Leccinellum cremeum HKAS90639 China – KT990781 KT990420 – Wu et al. 2016
Leccinellum crocipodium VDKO1006 Belgium KT823988 KT824054 KT824021 MH614856* Raspé et al. 2016; *Vadthanarat et al. 2019
Leccinellum sp. KPM-NC-0018041 Japan KC552165 KC552094 – – Orihara et al. 2016
Leccinellum sp. OR0711 Thailand MH614685 MH614733 MH614780 – Vadthanarat et al. 2019
Leccinum monticola HKAS76669 China – KF112249 KF112723 – Wu et al. 2014
Leccinum quercinum HKAS63502 China – KF112250 KF112724 – Wu et al. 2014
Leccinum scabrum RW105a Belgium KT823979 KT824045 KT824012 MH614857* Raspé et al. 2016; *Vadthanarat et al. 2019
Leccinum scabrum VDKO0938 Belgium MG212549 MG212593 MG212635 MH614858* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Leccinum scabrum KPM-NC-0017840 Scotland KC552170 JN378455 – – Orihara et al. 2016; Orihara et al. 2012
Leccinum schistophilum VDKO1128 Belgium KT823989 KT824055 KT824022 MH614859* Raspé et al. 2016; *Vadthanarat et al. 2019
Leccinum variicolor VDKO0844 Belgium MG212550 MG212594 MG212636 MH614860* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Mucilopilus castaneiceps HKAS75045 China – KF112211 KF112735 – Wu et al. 2014
Neoboletus brunneissimus HKAS52660 China – KF112143 KF112650 – Wu et al. 2014
Neoboletus brunneissimus HKAS57451 China – KM605149 KM605172 – Wu et al. 2015
Neoboletus brunneissimus OR0249 China MG212551 MG212595 MG212637 MH614861* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Neoboletus hainanensis HKAS59469 China – KF112175 KF112669 – Wu et al. 2014
Neoboletus junquilleus AF2922 France MG212552 MG212596 MG212638 MH614862* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Neoboletus magnificus HKAS54096 China – KF112149 KF112654 – Wu et al. 2014
Neoboletus magnificus HKAS74939 China – KF112148 KF112653 – Wu et al. 2014
Neoboletus sanguineoides HKAS55440 China – KF112145 KF112652 – Wu et al. 2014
Neoboletus sp. HKAS76851 China – KF112144 KF112651 – Wu et al. 2014
Neoboletus sp. OR0128 Thailand MH614686 MH614734 MH614781 MH614863 Vadthanarat et al. 2019
Neoboletus tomentulosus HKAS53369 China – KF112154 KF112659 – Wu et al. 2014
Neoboletus erythropus VDKO0690 Belgium KT823982 KT824048 KT824015 MH614864* Raspé et al. 2016; *Vadthanarat et al. 2019
Octaviania asahimontana KPM-NC-17824 Japan KC552154 JN378430 – – Orihara et al. 2016; Orihara et al. 2012
Octaviania asterosperma AQUI3899 Italy KC552159 KC552093 – – Orihara et al. 2016
Octaviania celatifilia KPM-NC-17776 Japan KC552147 JN378416 – – Orihara et al. 2016; Orihara et al. 2012
Octaviania cyanescens PNW-FUNGI-5603 USA KC552160 JN378438 – – Orihara et al. 2016; Orihara et al. 2012
Octaviania decimae KPM-NC17763 Japan KC552145 JN378409 – – Orihara et al. 2016; Orihara et al. 2012
Octaviania tasmanica MEL2128484 Australia KC552157 JN378437 – – Orihara et al. 2016; Orihara et al. 2012
Octaviania tasmanica MEL2341996 Australia KC552156 JN378436 – – 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 MG212597 MG212639 MH614865 Vadthanarat et al. 2019
Phylloporus bellus OR0473 China MH580778 MH580798 MH580818 MH614866* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus brunneiceps OR0050 Thailand KT823968 KT824034 KT824001 MH614867* Raspé et al. 2016; *Vadthanarat et al. 2019
Phylloporus castanopsidis OR0052 Thailand KT823969 KT824035 KT824002 MH614868* Raspé et al. 2016; *Vadthanarat et al. 2019
Phylloporus imbricatus HKAS68642 China – KF112299 KF112786 – Wu et al. 2014
Phylloporus luxiensis HKAS75077 China – KF112298 KF112785 – Wu et al. 2014
Phylloporus maculatus OR0285 China MH580780 MH580800 MH580820 – Chuankid et al. 2019
Phylloporus pelletieri WU18746 Austria MH580781 MH580801 MH580821 MH614869* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus pusillus OR1158 Thailand MH580783 MH580803 MH580823 MH614870* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus rhodoxanthus WU17978 USA MH580785 MH580805 MH580824 MH614871* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus rubeolus OR0251 China MH580786 MH580806 MH580825 MH614872* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus rubiginosus OR0169 China MH580788 MH580808 MH580827 MH614873* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus sp. OR0896 Thailand MH580790 MH580810 MH580829 MH614874* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus subbacillisporus OR0436 China MH580792 MH580812 MH580831 MH614875* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus subrubeolus BC022 Thailand MH580793 MH580813 MH580832 MH614876* Chuankid et al. 2019; *Vadthanarat et al. 2019
Phylloporus yunnanensis OR0448 China MG212554 MG212598 MG212640 MH614877* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Porphyrellus castaneus OR0241 China MG212555 MG212599 MG212641 MH614878* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Porphyrellus cf. nigropurpureus ADK3733 Benin MH614687 MH614735 MH614782 MH614879 Vadthanarat et al. 2019
Porphyrellus nigropurpureus HKAS74938 China – KF112246 KF112763 – Wu et al. 2014
Porphyrellus porphyrosporus MB97-023 Germany DQ534609 GU187734 GU187800 – Binder & Hibbett 2006; Binder et al. 2010
Porphyrellus sp. HKAS53366 China – KF112241 KF112716 – Wu et al. 2014
Porphyrellus sp. JD659 Burundi MH614688 MH614736 MH614783 MH614880 Vadthanarat et al. 2019
Porphyrellus sp. OR0222 Thailand MH614689 MH614737 MH614784 MH614881 Vadthanarat et al. 2019
Pulveroboletus aff. ravenelii ADK4360 Togo KT823957 KT824023 KT823990 MH614882* Raspé et al. 2016; *Vadthanarat et al. 2019
Pulveroboletus aff. ravenelii ADK4650 Togo KT823959 KT824025 KT823992 MH614883* Raspé et al. 2016; *Vadthanarat et al. 2019
Pulveroboletus aff. ravenelii HKAS53351 China – KF112261 KF112712 – Wu et al. 2014
Pulveroboletus fragrans OR0673 Thailand KT823977 KT824043 KT824010 MH614884* Raspé et al. 2016; *Vadthanarat et al. 2019
Pulveroboletus ravenelii REH2565 USA KU665635 KU665636 KU665637 MH614885* Raspé et al. 2016; *Vadthanarat et al. 2019
Pulveroboletus sp. HKAS74933 China – KF112262 KF112713 – Wu et al. 2014
Retiboletus aff. nigerrimus OR0049 Thailand KT823967 KT824033 KT824000 MH614886* Raspé et al. 2016; *Vadthanarat et al. 2019
Retiboletus brunneolus HKAS52680 China – KF112179 KF112690 – Wu et al. 2014
Retiboletus fuscus HKAS59460 China – JQ928580 JQ928601 – Hosen et al. 2013
Retiboletus fuscus OR0231 China MG212556 MG212600 MG212642 MH614887* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Retiboletus griseus MB03-079 USA KT823964 KT824030 KT823997 MH614888* Raspé et al. 2016; *Vadthanarat et al. 2019
Retiboletus kauffmanii OR0278 China MG212557 MG212601 MG212643 MH614889* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Retiboletus nigerrimus HKAS53418 China – KT990824 KT990462 – Wu et al. 2016
Retiboletus sinensis HKAS59832 China – KT990827 KT990464 – Wu et al. 2016
Retiboletus zhangfeii HKAS59699 China – JQ928582 JQ928603 – Hosen et al. 2013
Rhodactina himalayensis CMU25117 Thailand MG212558 MG212602, MG212603 – – Vadthanarat et al. 2018
Rhodactina rostratispora SV170 Thailand MG212560 MG212605 MG212645 – 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 OSC61484 Australia KC552109 JN378446 – – Orihara et al. 2016; Orihara et al. 2012
Royoungia reticulata HKAS52253 China – KT990786 KT990427 – Wu et al. 2016
Royoungia rubina HKAS53379 China – KF112274 KF112796 – Wu et al. 2014
Rubroboletus latisporus HKAS80358 China – KP055020 KP055029 – Zhao et al. 2014b
Rubroboletus legaliae VDKO0936 Belgium KT823985 KT824051 KT824018 MH614890* Raspé et al. 2016; *Vadthanarat et al. 2019
Rubroboletus rhodosanguineus BOTH4263 USA MG897416 MG897426 MG897436 MH614891* Phookamsak et al. 2019, *Vadthanarat et al. 2019
Rubroboletus rhodoxanthus HKAS84879 Germany – KT990831 KT990468 – Wu et al. 2016
Rubroboletus satanas VDKO0968 Belgium KT823986 KT824052 KT824019 MH614892* Raspé et al. 2016; *Vadthanarat et al. 2019
Rubroboletus sinicus HKAS68620 China – KF112146 KF112661 – Wu et al. 2014
Rubroboletus sp. HKAS68679 China – KF112147 KF112662 – Wu et al. 2014
Rugiboletus brunneiporus HKAS68586 China – KF112197 KF112719 – Wu et al. 2014
Rugiboletus brunneiporus HKAS83209 China – KM605144 KM605168 – Wu et al. 2015
Rugiboletus extremiorientalis HKAS63635 China – KF112198 KF112720 – Wu et al. 2014
Rugiboletus extremiorientalis HKAS76663 China – KM605147 KM605170 – Wu et al. 2015
Rugiboletus extremiorientalis OR0406 Thailand MG212562 MG212607 MG212647 MH614893* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Rugiboletus sp. HKAS55373 China – KF112303 KF112804 – Wu et al. 2014
Singerocomus inundabilis TWH9199 Guyana MH645588 MH645596 LC043089* MH645609 *Henkel et al. 2016; Vadthanarat et al. 2019
Singerocomus rubriflavus TWH9585 Guyana MH645589 MH645597 – MH645610 Vadthanarat et al. 2019
Spongiforma thailandica DED7873 Thailand MG212563 KF030436* MG212648 MH614894** *Nuhn et al. 2013; Vadthanarat et al. 2018; **Vadthanarat et al. 2019
Strobilomyces atrosquamosus HKAS55368 China – KT990839 KT990476 – Wu et al. 2016
Strobilomyces echinocephalus OR0243 China MG212564 MG212608 MG212649 – Vadthanarat et al. 2018
Strobilomyces strobilaceus RW103 Belgium KT823978 KT824044 KT824011 MH614895* Raspé et al. 2016; *Vadthanarat et al. 2019
Strobilomyces strobilaceus MB-03-102 USA DQ534607* AY883428 AY786065 – Binder and Hibbett 2006*, Unpublished
Strobilomyces mirandus OR0115 Thailand KT823972 KT824038 KT824005 MH614896* Raspé et al. 2016; *Vadthanarat et al. 2019
Strobilomyces sp. OR0259 China MG212565 MG212609 MG212650 MH614897* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Strobilomyces sp. OR0778 Thailand MG212566 MG212610 MG212651 MH614899* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Strobilomyces sp. OR0319 Thailand MH614690 MH614738 MH614785 MH614898 Vadthanarat et al. 2019
Strobilomyces sp. OR1092 Thailand MH614691 MH614739 MH614786 MH614900 Vadthanarat et al. 2019
Strobilomyces verruculosus HKAS55389 China – KF112259 KF112813 – Wu et al. 2014
Suillellus amygdalinus 112605ba USA – JQ327024 – – Halling et al. 2012a
Suillellus luridus VDKO0241b Belgium KT823981 KT824047 KT824014 MH614901* Raspé et al. 2016; *Vadthanarat et al. 2019
Suillellus queletii VDKO1185 Belgium MH645590 MH645598 MH645604 MH645611 Vadthanarat et al. 2019
Suillellus subamygdalinus HKAS57262 China – KF112174 KF112660 – Wu et al. 2014
Sutorius australiensis REH9441 Australia MG212567 JQ327032* MG212652 – *Halling et al. 2012a; Vadthanarat et al. 2018
Sutorius eximius REH9400 USA MG212568 JQ327029* MG212653 MH614902** *Halling et al. 2012a; Vadthanarat et al. 2018; **Vadthanarat et al. 2019
Sutorius ferrugineus HKAS77718 China – KT990789 KT990431 – Wu et al. 2016
Sutorius flavidus HKAS59443 China – KU974136 KU974144 – Wu et al. 2016
Sutorius rubriporus HKAS83026 China – KT990795 KT990437 – Wu et al. 2016
Sutorius sanguineus HKAS80823 China – KT990802 KT990442 – Wu et al. 2016
Sutorius sp. OR0378B Thailand MH614692 MH614740 MH614787 MH614903 Vadthanarat et al. 2019
Sutorius sp. OR0379 Thailand MH614693 MH614741 MH614788 MH614904 Vadthanarat et al. 2019
Tengioboletus glutinosus HKAS53425 China – KF112204 KF112800 – Wu et al. 2014
Tengioboletus reticulatus HKAS53426 China – KF112313 KF112828 – Wu et al. 2014
Tengioboletus sp. HKAS76661 China – KF112205 KF112801 – 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 – KF112284 KF112730 – Wu et al. 2014
Tylopilus alpinus HKAS55438 China – KF112191 KF112687 – Wu et al. 2014
Tylopilus atripurpureus HKAS50208 China – KF112283 KF112799 – Wu et al. 2014
Tylopilus balloui s.l. OR0039 Thailand KT823965 KT824031 KT823998 MH614905* Raspé et al. 2016; *Vadthanarat et al. 2019
Tylopilus brunneirubens HKAS53388 China – KF112192 KF112688 – Wu et al. 2014
Tylopilus felleus VDKO0992 Belgium KT823987 KT824053 KT824020 MH614906* Raspé et al. 2016; *Vadthanarat et al. 2019
Tylopilus ferrugineus BOTH3639 USA MH614694 MH614742 MH614789 MH614907 Vadthanarat et al. 2019
Tylopilus otsuensis HKAS53401 China – KF112224 KF112797 – Wu et al. 2014
Tylopilus sp. HKAS74925 China – KF112222 KF112739 – Wu et al. 2014
Tylopilus sp. HKAS50229 China – KF112216 KF112769 – Wu et al. 2014
Tylopilus sp. JD598 Gabon MH614695 MH614743 MH614790 MH614908 Vadthanarat et al. 2019
Tylopilus sp. OR0252 China MG212569 MG212611 MG212654 MH614909* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Tylopilus sp. OR0542 Thailand MG212570 MG212612 MG212655 MH614910* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Tylopilus sp. OR0583 Thailand MH614696 MH614744 – – Vadthanarat et al. 2019
Tylopilus sp. OR1009 Thailand MH614697 – MH614791 MH614911 Vadthanarat et al. 2019
Tylopilus vinaceipallidus HKAS50210 China – KF112221 KF112738 – Wu et al. 2014
Tylopilus vinaceipallidus OR0137 China MG212571 MG212613 MG212656 MH614912* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Tylopilus violaceobrunneus HKAS89443 China – KT990886 KT990504 – Wu et al. 2016
Tylopilus virens KPM-NC-0018054 Japan KC552174 KC552103 – – Unpublished
Veloporphyrellus alpinus HKAS68301 China JX984515 JX984550 – – Li et al. 2014
Veloporphyrellus conicus REH8510 Belize MH614698 MH614745 MH614792 MH614913 Vadthanarat et al. 2019
Veloporphyrellus gracilioides HKAS53590 China – KF112210 KF112734 – 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 – KT990850 KT990486 – Wu et al. 2016
Xanthoconium porophyllum HKAS90217 China – KT990851 KT990487 – Wu et al. 2016
Xanthoconium sinense HKAS77651 China – KT990853 KT990488 – Wu et al. 2016
Xerocomellus chrysenteron VDKO0821 Belgium KT823984 KT824050 KT824017 MH614914* Raspé et al. 2016; *Vadthanarat et al. 2019
Xerocomellus cisalpinus ADK4864 Belgium KT823960 KT824026 KT823993 MH614915* Raspé et al. 2016; *Vadthanarat et al. 2019
Xerocomellus communis HKAS50467 China – KT990858 KT990494 – Wu et al. 2016
Xerocomellus corneri HKAS90206 Philippines – KT990857 KT990493 – Wu et al. 2016
Xerocomellus porosporus VDKO0311 Belgium MH614678 MH614727 MH614773 MH614846 Vadthanarat et al. 2019
Xerocomellus ripariellus VDKO0404 Belgium MH614699 MH614746 MH614793 MH614916 Vadthanarat et al. 2019
Xerocomellus sp. HKAS56311 China – KF112170 KF112684 – Wu et al. 2014
Xerocomus aff. macrobbii HKAS56280 China – KF112265 KF112708 – Wu et al. 2014
Xerocomus fulvipes HKAS76666 China – KF112292 KF112789 – Wu et al. 2014
Xerocomus magniporus HKAS58000 China – KF112293 KF112781 – Wu et al. 2014
Xerocomus s.s. sp. OR0237 China MH580796 MH580816 MH580835 – Chuankid et al. 2019
Xerocomus s.s. sp. OR0443 China MH580797 MH580817 MH580836 MH614917* Chuankid et al. 2019; *Vadthanarat et al. 2019
Xerocomus sp. OR0053 Thailand MH580795 MH580815 MH580834 MH614918* Chuankid et al. 2019; *Vadthanarat et al. 2019
Xerocomus subtomentosus VDKO0987 Belgium MG212572 MG212614 MG212657 MH614919* Vadthanarat et al. 2018; *Vadthanarat et al. 2019
Zangia citrina HKAS52684 China HQ326850 HQ326872 – – Li et al. 2011
Zangia olivacea HKAS45445 China HQ326854 HQ326873 – – Li et al. 2011
Zangia olivaceobrunnea HKAS52272 China HQ326857 HQ326876 – – Li et al. 2011
Zangia roseola HKAS75046 China – KF112269 KF112791 – Wu et al. 2014
Zangia roseola HKAS51137 China HQ326858 HQ326877 – – Li et al. 2011

Taxonomy

Erythrophylloporus Ming Zhang & T.H. Li, Mycosystema 37(9): 1111–1126 (2018)

Description

Basidiomata stipitate-pileate with lamellate hymenophore, small to medium-sized; Pileus subhemispheric to convex when young becoming convex to plano-convex to plano-subdepressed when old, dry, pruinose or velutinous, subtomentose to tomentose, yellowish-orange to red; pileus context vivid yellow to yellowish-orange. Hymenophore lamellae, slightly thick, decurrent, deeply yellowish-orange to deep orange or reddish-orange to orange red or brownish-orange to red. Stipe central to slightly excentric, cylindrical or clavate, yellowish- to reddish-orange to yellowish red, with scattered yellowish- to reddish-orange to red scales on surface, with bright yellow basal mycelium; stipe context solid, yellow to reddish-yellow or yellow with olivaceous brown. Staining none or slightly reddening or greening or gradually bluing or dark violet, greyish to blackish-blue when bruised on the basidiomata or context or lamellae. Spore print olivaceous brown. Basidiospores ovoid or ellipsoid to broadly ellipsoid to subovoid, thin-walled, with non-bacillate surface. Basidia clavate to narrowly clavate. Cheilocystidia and pleurocystidia present, subcylindrical or narrowly conical to narrowly fusiform to ventricose with slightly or obtuse apex, thin-walled, sometimes thick-walled, originating more or less deeply in the sub hymenium or from hymenophoral trama, hyaline or sometimes containing yellowish-brown pigments. Pileipellis a subcutis to cutis to trichoderm to palisadoderm, composed of thin to slightly thick-walled hyphae. Clamp connection absent in all tissues.

Typus species

Erythrophylloporus cinnabarinus Ming Zhang & T.H. Li.

Known Distribution

Asia (China and Thailand), North America (Mexico) and Central America (Costa Rica).

Remarks

Erythrophylloporus is easily distinguished from other lamellate Boletaceae genera by a combination of the following characters: the intense orange to red colour of the pileus and lamellae; bright yellow basal mycelium; ovoid or ellipsoid to broadly ellipsoid to subovoid basidiospores with non-bacillate surface; pleurocystidia originating more or less deeply in the subhymenium or from hymenophoral trama.

Erythrophylloporus paucicarpus Raspé, Vadthanarat & Lumyong, sp. nov.

MycoBank No: 823605
Figs 2A, 3A, 4A and 5

Holotype

THAILAND, Chiang Mai Province, Mae On District, Huay Kaew, 18°52'0"N, 99°17'30"E, elev. 700 m, 16 August 2016, O. Raspé & S. Vadthanarat, OR1151, (holotype: CMUB, isotype: BR).

Etymology

from Latin “pauci-” meaning few and “carpus” meaning fruits or what is harvested, refers to the low number of basidiomata produced.

Description

Basidiomata stipitate-pileate with lamellate hymenophore, small to medium-sized. Pileus 2.3–5.5 cm in diameter, plano-convex with involute margin at first becoming almost plane to slightly depressed with inflexed to straight margin, irregularly and coarsely crenate in age, sometimes with low and broad umbo and a few to several verrucae, especially when young; surface more or less even, tomentose, dull, slightly moist, colour distribution patchy with red to brownish-orange (9B8 to 9C8), brownish-red (10E8 to 10D8) becoming orange-red to orange (8B/C8 to 6B7) at the margin when old, abruptly paler at the margin. Pileus context 3–4 mm thick half-way to the margin, tough, colour distribution even, yellow (3A6) to yellowish-orange (4A5), slowly reddening when exposed, especially at the centre and above lamellae. Stipe 2.4–4.5 × 0.7–1.3 cm, central or sometimes slightly eccentric, clavate with strigose base, straight to curved, terete, even, dull, dry, tomentose, yellowish-orange (4–5A7–8) to orange (6–7A7–8) with orange to yellowish-orange (7B/C7–8 to 4A7–8) coarse scales, with bright yellow (2A6–7) basal mycelium. Stipe context solid, fleshy fibrous, yellow marbled with olivaceous brown (4D5, 5D5). Hymenophore lamellate; lamellae decurrent, close, thick, 40–42 lamellae, with 4–6 different lengths of lamellullae, 2–4.5 mm wide half-way to margin, somewhat anastomosing, especially near the stipe, yellowish-orange (4-5A6-7) with orange to red tinge, slightly reddening when bruised. Odour rubbery; Taste not recorded. Spore print olive-brown (4E7).

Figure 2. 

Habits of Thai Erythrophylloporus species A E. paucicarpus B E. suthepensis. Scale bars: 1 cm.

Macrochemical reactions. KOH on pileus and stipe surface deep red at first, then red-brown to brown, with pale orange aura on the pileus; brown on pileus context, dark red-brown on stipe context; brownish-orange on hymenophore. NH4OH on pileus first red, then orange; on pileus context bluing at first then with a greenish tinge; on stipe surface and context briefly bluing; no reaction on hymenophore.

Basidiospores [208/4/4] (5.9–)6.1–6.8–7.5(–8) × (4.1–)4.6–5.1–5.5(–6) µm, Q = (1.2–)1.23–1.33–1.48(–1.56); from the type (OR1151) (6–)6.3–6.8–7.5(–7.8) × (4.6–)4.8–5.2–5.5(–6) µm, Q = (1.2–)1.22–1.31–1.48(–1.56), N = 88, broadly ellipsoid to ellipsoid, smooth under light microscope and SEM, yellowish to pale brown in water, hyaline in 5% KOH, thin-walled, inamyloid. Basidia 4–spored, (37.8–)38–45.6–54.7(–54.8) × (6.2–)–6.3–8–9.5(–9.6) µm, narrowly clavate to subcylindrical, attenuated towards the base, clampless, hyaline to yellowish hyaline in water, Melzer’s reagent and 5% KOH; sterigmata up to 5.5 µm long. Cheilocystidia (35.4–)35.5–49.9–61.8(–61.9) × (3.9–)3.9–6–7.7(–7.7) µm, narrowly fusiform with obtuse apex, projecting up to 30 µm, thin-walled, smooth, yellowish hyaline in water, in 5% KOH and NH4OH, inamyloid. Pleurocystidia (66.9–)67.4–80.3–93.5(–94.7) × (8.8–)8.9–11.7–16.1(–16.2) µm, abundant, narrowly conical with obtuse, somewhat prolonged apex, projecting up to 32 µm, thin-walled, smooth, yellowish hyaline in water, in 5% KOH and NH4OH, arising more or less deeply in the subhymenium or from hymenophoral trama, inamyloid. Hymenophoral trama subregular near the pileus context becoming slightly divergent near the edge, 87–238 µm wide, widest near the pileus context then getting narrower when close to the edge, composed of clampless hyphae 4.5–8 µm wide, yellowish hyaline in water, hyaline in 5% KOH and NH4OH. Pileipellis a palisadoderm to trichoderm 83–165 µm thick, composed of slightly thick-walled, cylindrical hyphae, terminal cells 16–46 × 4–6.5 µm with rounded apex, hyaline or yellowish hyaline to yellowish-orange hyaline hyphae ornamented with scattered fine epiparietal encrustation when observed in water, hyaline to yellowish hyaline in 5% KOH and NH4OH, inamyloid. Pileus trama composed of slightly thick-walled, strongly interwoven hyphae, 4.5–8.5 µm wide, inamyloid. Stipitipellis a disrupted palisadoderm perpendicular to the stipe axis, 63–145 µm thick, composed of slightly thick-walled, slightly rough, cylindrical, yellow to yellowish-orange in water, yellowish hyaline hyphae in 5% KOH and NH4OH, terminal cells 13–57 × 3–8 µm, cylindrical to irregular hyphae with rounded to notched apex; wall covered by dispersed fine encrustations when observed in water. Caulocystidia not seen. Stipe trama composed of parallel hyphae, densely packed, 4–8.5 µm wide; hyphae wall covered by dispersed encrustations when observed in water. Clamp connections not seen in any tissue.

Figure 3. 

Scanning electron micrographs of basidiospores from Thai Erythrophylloporus show smooth surfaces A E. paucicarpus B E. suthepensis. Scale bars: 1 µm.

Habit and habitat

On soil, mostly solitary in dipterocarp forest dominated by Dipterocarpus tuberculatus, D. obtusifolius, Shorea obtusa, S. siamensis, Quercus spp. and Lithocarpus spp.

Known distribution

Currently known only from Chiang Mai Province, northern Thailand.

Additional specimens examined

– THAILAND, Chiang Mai Province, Muang District, Doi Suthep-Pui National Park, 18°48'05"N–98°55'40"E, elev. 800 m, 17 May 2015, O. Raspé, OR0615A (CMUB, BKF, BR); Mae Taeng District, Baan Tapa, 19°08'29"N, 98°45'47"E, elev. 1035 m, 4 August 2015, O. Raspé & A. Thawthong, OR0689 (MFLU, BR); Mae On District, Huay Kaew, 18°52'12"N, 99°18'12"E, elev. 780 m, 15 August 2016, O. Raspé & S. Vadthanarat, OR1135 (CMUB, BR).

Remarks

E. paucicarpus is characterised by the following combination of features: orange to brownish- to orange-red basidiomata, yellowish-orange lamellae that turn slightly red when bruised; pileus context yellow to yellowish-orange that slowly reddens when exposed and mostly occurring as solitary basidiomata.

In the inferred molecular phylogeny, E. paucicarpus clustered close to E. suthepensis and E. cinnabarinus (65% BS and 1 PP), but the two species are different from E. paucicarpus in that they have darker lamellae which are orange to orange red or brownish-orange. Moreover, spores of E. paucicarpus are wider and longer (5.9–8 × 4.1–6 µm) than those of E. suthepensis (4.6–5.9 × 3.5–4.5 µm) and, on average, longer than those of E. cinnabarinus (5.5–7 × 4.5–5.5 µm) (Zhang and Li 2018). Erythrophylloporus paucicarpus also differs from both species by the slight reddening of the context and lamellae when exposed or bruised, whereas E. suthepensis context seems unchanging when exposed and lamellae turn blue when bruised. In E. cinnabarinus, the context slowly turns dark violet, blackish-blue to dark blue when exposed and lamellae turn greyish-blue, or greyish-green when bruised (Zhang and Li 2018).

Figure 4. 

Origin of pleurocystidia (white arrow), more or less deep in the subhymenium or from hymenophoral trama A E. paucicarpus B E. suthepensis – hymenium (H), subhymenium (SH), Scale bars: 25 µm (A–B).

Erythrophylloporus paucicarpus is different from the two New World species by the reddening of the context, whereas in E. fagicola, it turns blue and, in E. aurantiacus, the colour remains unchanged when exposed. Moreover, E. fagicola has somewhat thick-walled (0.8–3.5 µm) pleurocystidia (Montoya and Bandala 2011), which are not found in E. paucicarpus. Although the basidiospores of E. paucicarpus and E. aurantiacus are similar in size (E. aurantiacus = 6.0–7.5 × 4–5.5 µm), they differ in shape, being more ovoid in E. aurantiacus than in E. paucicarpus. Erythrophylloporus paucicarpus also differs from E. aurantiacus by macro-chemical reactions. In the latter, the pileus surface and pileus context are unchanging with NH4OH (Halling et al. 1999), while in E. paucicarpus, the pileus becomes orange to red and the pileus context initially turns blue then with a greenish tinge.

Erythrophylloporus suthepensis Vadthanarat, Raspé & Lumyong, sp. nov.

MycoBank No: 823606
Figs 2B, 3B, 4B and 6

Holotype

THAILAND, Chiang Mai Province, Muang District, Doi Suthep-Pui National Park, 18°48'47"N, 98°55'56"E, elev. 645 m, 25 August 2015, S. Vadthanarat, SV0236, (holotype CMUB, isotype BKF, BR).

Etymology

Refers to the type locality Doi Suthep.

Description

Basidiomata stipitate-pileate with lamellate hymenophore, small-sized. Pileus (1.0–)2.5– 3.5 cm in diameter, subumbonate with involute margin at first, becoming convex to plano-convex with inflexed margin; surface even with some small pustules, tomentose, dull, slightly moist, yellow (3–4A4– 5) becoming light orange to orange-red (5–6A5–7 to 7–8A–B7–8) with patches of light yellow to light orange (4–5A5–6) becoming brownish-orange to dull red (7B–C8 to 8B–D8) with age, the colour of the margin when young clearly paler than the rest of the pileus, bluing when bruised. Pileus context 2–3 mm thick half-way to the margin, tough, yellowish-orange (4A5), unchanging when bruised. Stipe 2.5– 4.5 × 0.3– 0.8 cm, central, slightly curved, terete, dull, dry, yellowish-orange (2A6–7) with greyish-orange (5–6 B 7–8) coarse scales at first, then light yellow or reddish-yellow to brownish-orange (4A/B5–6 to 7C6) with brownish-red to reddish-dark brown (7F4–5, 8C7–8, 8F5–7) scales, sub-bulbous, with bright yellow to greyish-yellow (2A6–7 to 3A/B5–6) sparse basal mycelium that extends half-way up the stipe. Stipe context solid, tough, reddish-yellow (4A6) near the pileus then paler to light yellow (4A5) near the base, unchanging when bruised. Hymenophore lamellate; lamellae decurrent, subdistant, slightly thick, with sinuate edge, of varying lengths, 26–34 lamellae, with 4–6 different lengths of lamellullae, 4–5 mm wide half-way to margin, brownish-orange (7C7–8) with deep yellow to orange (4–5A7–8) edge, bluish-grey when looking tangentially to the surface, bluing when bruised. Odour rubbery. Taste mild with rubbery texture. Spore print olivaceous brown (4F5).

Macrochemical reactions. KOH orange-brown on pileus and stipe surface; yellowish-brown on pileus and stipe context and hymenophore. NH4OH yellowish-brown on pileus and stipe surface and hymenophore; yellowish on pileus and stipe context.

Basidiospores [218/4/2] (4.6–)4.8–5.2–5.7(–5.9) × (3.5–)3.6–4–4.3(–4.5) µm, Q = (1.15–)1.21–1.32–1.44(–1.57); from the type (SV0236) (4.6–)4.8–5.2–5.7(–5.9) × (3.5–)3.6–3.9–4.4(–4.5) µm, Q = (1.15–)1.21–1.32–1.43(–1.57), N = 80, broadly ellipsoid to subglobose, smooth under light microscope and SEM, yellowish to pale brown in water, hyaline in 5% KOH, thin-walled, inamyloid. Basidia 4-spored, (24.7–)25.3–31.1–35.8(–35.9) × (5.3–)5.3–6.6–7.5(–7.5) µm, narrowly clavate to subcylindrical, attenuated towards the base, clampless, hyaline to yellowish hyaline in water, Melzer’s reagent and 5% KOH; sterigmata up to 4.5 µm long. Cheilocystidia (37.3–)37.9–51–63.8(–64.1) × (5.3–)5.4–8.5–12.4(–13.7) µm, narrowly conical to narrowly fusiform with obtuse apex, projecting up to 25 µm, thin-walled, smooth, yellowish-hyaline in water, hyaline in 5% KOH and NH4OH, inamyloid, more or less forming a sterile edge . Pleurocystidia (46.5–)49.2–68.9–95.2(–99.3) × (9.3–)9.6–12.6–18.9(–20) µm, abundant, narrowly conical with obtuse apex, projecting up to 28 µm, thin-walled, mostly yellowish hyaline in water and hyaline in 5% KOH and NH4OH, some containing yellowish-brown to dark brown pigments in water and yellowish-pale brown in 5% KOH and NH4OH, inamyloid, arising more or less deeply in the subhymenium or from hymenophoral trama. Hymenophoral trama subregular near the pileus context becoming slightly divergent near the edge, 46–192 µm wide, widest near the pileus context then getting narrower when close to the edge, composed of clampless hyphae 2.5–7.5 µm wide, pinkish-red hyaline in water, especially at the centre of the trama, yellowish hyaline to hyaline in 5% KOH and NH4OH. Pileipellis a palisadoderm to trichoderm 71–119 µm thick, composed of slightly thick-walled, cylindrical to irregular hyphae with fine encrustation on the wall, terminal cells 12–46 × 3.5–9 µm with pointed to notched apex or sometimes truncated apex, with 6–15(–28) µm short cells at the base, hyaline or yellowish-orange hyaline to orange hyaline hyphae with scattered fine encrustation on the wall when observed in water, hyaline to yellowish hyaline in 5% KOH and NH4OH, inamyloid. Pileus context composed of slightly thick-walled, strongly interwoven hyphae, 5–8.5 µm wide, inamyloid. Stipitipellis a disrupted palisadoderm perpendicular to the stipe axis, 47–123 µm thick, composed of slightly thick-walled, cylindrical to irregular hyphae with fine encrustations on the wall, yellow to yellowish-orange, intermixed with mostly yellowish hyaline to yellowish-brown hyphae in 5% KOH and NH4OH, terminal cells 14–47 × 4–8.5 µm with variously notched apex. Caulocystidia mixed in a group with the stipitipellis hyphae, same shape and size as the pleurocystidia, dark brown in water, paler in 5% KOH and NH4OH. Stipe context composed of parallel, densely packed, 4–9.5 µm wide hyphae, hyphae wall with scattered fine encrustations when observed in water. Clamp connections not seen in any tissue.

Figure 5. 

Microscopic features of Erythrophylloporus paucicarpus A basidiospores B basidia C cheilocystidia D pleurocystidia E pileipellis F stipitipellis. – Scale bars: 10 µm (A–B); 50 µm (C–F). All drawings were made from the type (OR1151).

Habit and habitat

On soil, gregarious (up to 10 basidiomata) in dipterocarp forest dominated by Dipterocarpus tuberculatus, D. obtusifolius, Shorea obtusa and S. siamensis, mixed with scattered fagaceous trees.

Known distribution

Currently known only from Doi Suthep-Pui National Park, Chiang Mai Province, northern Thailand.

Additional specimens examined

– THAILAND, Chiang Mai Province, Meuang District, Doi Suthep-Pui National Park, 18°48'05"N, 98°55'40"E, elev. 800 m, 17 May 2015, O. Raspé, OR0615B (CMUB, BKF, BR).

Remarks

Erythrophylloporus suthepensis is characterised by the following combination of features: yellow to light orange to orange red to brownish-orange to dull red pileus; brownish-orange lamellae with deep yellow to orange edge; the colour of the lamellae appears more bluish-grey when observed from an oblique angle to the surface; pileus surface and lamellae turning blue when bruised; some pleurocystidia containing yellowish-brown to dark brown pigments in water; basidiospores that are smaller or shorter (4.6–5.9 × 3.5–4.5 µm) than the other Erythrophylloporus species (E. aurantiacus = 6.0–7.5 × 4–5.5µm; E. cinnabarinus = 5.5–7 × 4.5–5.5 µm; E. fagicola = 6.5–11 × 4–7.5 µm; E. paucicarpus = 5.9–8 × 4.1–6 µm) (Halling et al. 1999, Montoya and Bandala 2011, Zhang and Li 2018).

Morphologically, E. suthepensis is quite similar to E. cinnabarinus in that they have similar colours in pileus and lamellae; the lamellae in both species also turn more or less blue to dark blue when bruised. Erythrophylloporus suthepensis and E. cinnabarinus are also similar, based on some pleurocystidia containing yellowish-brown to dark brown pigments, but those features are not found in E. paucicarpus and in the two New World Erythrophylloporus species (Halling et al. 1999, Montoya and Bandala 2011). However, the pleurocystidia containing brown pigments seem to be more frequent in E. cinnabarinus, which also has, on average, larger basidiospores than E. suthepensis (Zhang and Li 2018).

The pinkish-red hymenophoral trama of E. suthepensis was not found in either E. paucicarpus or in the two American Erythrophylloporus species. In our observation of the two American specimens (E. aurantiacus voucher REH7271 and E. fagicola voucher Garay215), we found that the hymenophoral trama was yellowish hyaline when observed in water. The original description of E. cinnabarinus does not mention the colour of the hymenophoral trama and we could not obtain a specimen to observe this character. However, other morphological characters and phylogenetic evidence are enough to differentiate E. suthepensis from E. cinnabarinus.

Our phylogenetic analyses of a four-gene dataset revealed that Phylloporus aurantiacus from Costa Rica and P. fagicola from Mexico clustered in the Erythrophylloporus clade with high support (BS = 100% and PP = 1). Both species possess the distinctive morphological characters of Erythrophylloporus, which include yellowish-orange to reddish-orange basidiomata, orange to orange brown lamellae, bright yellow basal mycelium, ovoid or ellipsoid to broadly ellipsoid basidiospores with smooth surface and subcylindrical to subfusoid to ventricose cheilocystidia and pleurocystidia (Halling et al. 1999, Montoya and Bandala 2011). Therefore, the following two new combinations are proposed:

Figure 6. 

Microscopic features of Erythrophylloporus suthepensis A basidiospores B basidia C cheilocystidia D pleurocystidia E pileipellis F stipitipellis showing some dark caulocystidia mixed with slightly rough, cylindrical to irregular hyphae. – Scale bars: 10 µm (A–B); 50 µm (C–F). All drawings were made from the type (SV0236).

Erythrophylloporus aurantiacus (Halling & G.M. Muell.) Raspé & Vadthanarat, comb. nov.

MycoBank No: 823607

Basionym

Phylloporus aurantiacus Halling & G.M. Mueller, Mycotaxon 73: 64 (1999)

Specimen examined

– COSTA RICA. Near town of Palo Verde, elev. 1600 m, 11 June 1994, Halling 7271 (NY).

Erythrophylloporus fagicola (Montoya & Bandala) Raspé & Vadthanarat, comb. nov.

MycoBank No: 823608

Basionym

Phylloporus fagicola Montoya & Bandala, Mycotaxon 117: 10 (2011)

Specimen examined

– MEXICO. Veracruz: Mpio. Acatlán, Acatlán Volcano, 29 September 2009, Garay 215 (XAL).

Key to the species in Erythrophylloporus

1 Growing in North or Central America 2
– Growing in Southeast Asia or in tropical to subtropical China 3
2 Bluing of the context when exposed; basidiospores ellipsoid to oblong, obtuse, 6.5–11 × 4–7.5 µm; pleurocystidia somewhat thick-walled (0.8–3.5 µm thick) E. fagicola
– Context unchanging when exposed; basidiospores ovoid to subellipsoid, 6.0–7.5 × 4–5.5 µm; pleurocystidia thin-walled E. aurantiacus
3 Yellowish-orange lamellae slightly reddening when bruised; context slowly or slightly reddening when exposed E. paucicarpus
– Brownish-orange or orange, deep orange, reddish-orange to orange red lamellae bluing to greyish-green when bruised; context unchanging to gradually turning dark violet, blackish to dark blue 4
4 Basidiospores 4.6–5.9 × 3.5–4.5 µm, broadly ellipsoid to subglobose; cystidia mostly hyaline, only some containing yellowish-brown to dark brown pigments. E. suthepensis
– Basidiospores 5.5–7 × 4.5–5.5 µm, broadly ellipsoid, ellipsoid to nearly ovoid; cystidia usually containing yellowish-brown pigments E. cinnabarinus

Discussion

Both phylogeny and morphology support the placement of the two new species from Thailand, E. paucicarpus and E. suthepensis in the genus Erythrophylloporus. Phylogenetically, both species were highly supported in the Erythrophylloporus clade close to E. cinnabarinus (typus generis). Morphologically, they are characterised by having yellowish-orange to reddish- to brownish-orange basidiomata with bright yellow basal mycelium and smooth, ellipsoid, broadly ellipsoid to subglobose basidiospores. The other lamellate Boletaceae in Phylloporus, Phylloboletellus and Phylloporopsis are solely similar to the new species by having a lamellate hymenophore instead of a poroid hymenophore. However, Phylloporus differs from Erythrophylloporus species by having whitish- to yellowish-pale brown basidiomata with yellow to golden-yellow lamellae, with off-white to whitish to yellow basal mycelium and most species in the genus have basidiospores with more or less bacillate ornamentation under SEM (Neves & Halling 2010, Neves et al. 2012, Zeng et al. 2013). The single Phylloboletellus species, Ph. chloephorus Singer differs from Erythrophylloporus by having longitudinally ridged basidiospores (Bandala et al. 2004). The sole species of Phylloporopsis, Phy. boletinoides, differs by having beige to olive-cream or olive buff lamellate to subporoid hymenophore, with anastomosing and interveined gills and basal mycelium whitish to yellowish (Farid et al. 2018). Moreover, those genera are phylogenetically distant from Erythrophylloporus. (Bandala et al. 2004, Neves & Halling 2010, Neves et al. 2012, Zeng et al. 2013, Farid et al. 2018).

Interestingly, Phylloporus coccineus Corner, described from Singapore (Corner 1970), is similar to Erythrophylloporus species, in that it produces crimson to scarlet, lamellate basidiomata with orange to orange-red lamellae and yellow basal mycelium, broadly ellipsoid to subglobose and smooth basidiospores. It probably should also be transferred to Erythrophylloporus, but we refrain from doing so until specimens become available for molecular study. According to the protologue of P. coccineus, it differs from the newly described Asian species of Erythrophylloporus by having larger basidiospores (7.5–10 × 6.5–8 µm), larger cheilocystidia (70–120 × 10–18 µm) and larger caulocystidia (up to 200 × 10–16 µm) (Corner 1970).

Erythrophylloporus species formed two clades, an Asian species clade (BS = 65% and PP = 1) and a New World species clade (BS = 100% and PP = 1) (Fig. 1). The Asian one contains three species, E. cinnabarinus, E. paucicarpus and E. suthepensis, while the American clade contains the remaining two species E. aurantiacus and E. fagicola. Erythrophylloporus aurantiacus and E. fagicola seem to be genetically very close to each other, much closer than the species in the Asian clade. Only morphological differences between the two species were used to separate them from each other. Erythrophylloporus fagicola produces larger basidiospores than E. aurantiacus and pleurocystidia are somewhat thick-walled (0.8–3.5 µm thick) in E. fagicola, whereas they are thin-walled in E. aurantiacus and the latter has non-staining context, whereas the former has a cyanescent context. However, the descriptions were based on a limited number of collections and more samples are desirable to verify whether the morphological traits observed are good characters differentiating the two species or merely extremes of a continuum in morphological variation within a single species.

Regarding the phylogenetic affinities of Erythrophylloporus, Zhang and Li (2018) reported that it was likely close to the genus Rugiboletus G. Wu & Zhu L. Yang and Lanmaoa G. Wu & Zhu L. Yang, based on a multilocus dataset of nrLSU, tef1, rpb1 and rpb2, although this relationship was not supported in their phylogram. In our phylogeny, based on a multilocus dataset of atp6, tef1, rpb2 and cox3, with wider taxon sampling, Erythrophylloporus also clustered within the Pulveroboletus group, but was sister to Singerocomus with high bootstrap support (96%) but relatively weak posterior probability support (0.86). Singerocomus contains three species, S. atlanticus A.C. Magnago, S. inundabilis (Singer) T.W. Henkel and S. rubriflavus T.W. Henkel & Husbands that have some similar morphological characters to Erythrophylloporus, including red-orange to red pileus and light yellow basal mycelium. The three existing Singerocomus species are clearly different from all known Erythrophylloporus species by having a poroid, non-cyanescent hymenophore (Henkel et. al. 2016, Magnago et al. 2018). However, the hymenophore structure (lamellate vs. poroid) is not sufficient to separate genera in Boletaceae. Phylloporus currently contains both lamellate and poroid species, although some poroid species have already been transferred to another genus, Hourangia (Zhu et al. 2015). Phylogenetic analyses, including the remaining poroid Phylloporus species, are needed to verify their taxonomic position.

Erythrophylloporus putatively forms ectomycorrhizal associations with trees in family Fagaceae, including the genera Fagus, Lithocarpus and Quercus (Neves and Halling 2010, Montoya and Bandala 2011, Zhang and Li 2018). The two Thai Erythrophylloporus species were found in forests dominated by Dipterocarpaceae trees, mainly Dipterocarpus, including D. tuberculatus, D. obtusifolius and Shorea, including S. obtusa and S. siamensis. However, some Quercus and Lithocarpus trees (Fagaceae) were also observed in the vicinity and could also be the ectomycorrhizal partners. Further study is needed to confirm the ectomycorrhizal relationships of Erythrophylloporus.

Acknowledgements

Financial support from the Graduate School, Chiang Mai University, is appreciated. The work was partly supported by a TRF Research Team Association Grant (RTA5880006) to SL and OR. OR is grateful to the Fonds National de la Recherche Scientifique (Belgium) for travel grants. 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.

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